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Table of Contents.

MeROLTSBERG, Cari. Notes on-a visit to Easter Island .....-...-.:..
. HacerMan, T. H. Beitrige zur Geologie der Juan Fernandez-Inseln

. QuENsEL, P. Additional Comments on the Geology of the Juan Fernandez
RM cab cet rans. x, ie |x) Co, Sarg AE. oP vo a ements ea ot al

. SkoTrsBERG, C. A Geographical Sketch of the Juan Fernandez Islands . . .

5. —— Derivation of the Flora and Fauna of Juan Fernandez and Easter Island

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HE NATURAL HISTORY

JUAN FERNANDEZ
EASTER ISLAN D

ee V9 1 q |
GEOGRAPHY; GEOLOGY,
ORIGIN OF ISLAND LIFE | f

PART E> ns ae

‘C. SKOTTSBERG: Notes on a visit to Easter Island.

EASTER ISLAND
pol, afler the map of the Chilean Hydr. Office
1:200 000

Height ur meters above sea level.
x Observation spot

2. ee

1. Notes on a visit to Easter Island.
By
CARL SKOTTSBERG.

With 14 plates, 1 map, and 3 text figures.

While working on the Juan Fernandez Islands, our party obtained per-
mission to accompany the Chilean corvette »General Baquedano» on her cruise
to Easter Island, in 1917. A short and very popular account of our visit
appears in my book »Till Robinson-6n och varldens ande» (1918). Although
the purpose of our survey was purely biological, no scientist visiting the famous
island can help taking a vivid interest in the archaeological remains, and we
occasionally made a few observations. However, I have refrained from writing
anything on the subject, as I had to wait for the publication of the results
obtained by Mr. and Mrs. SCORESBY ROUTLEDGE of the British »Mana» Ex-
pedition. Last year Mrs. ROUTLEDGE published a most interesting account of
their work. on the island (The Mystery of Easter Island), where the ancient
monuments of all kinds are amply described and illustrated. A second volume
will follow, containing the detailed descriptions of the prehistoric remains.

I willingly admit that this little paper will appear rather unnecessary
since the British Expedition has explored the place with such a wonderful
accuracy. But it is Mrs. ROUTLEDGE’s excellent narrative which has induced
me to collect a few notes and to. use them as a basis for a discussion of some
interesting points. I have also found it worth while to add a number of my
photographs, which may be of some value.

The »General Baquedano» sailed from Iquique on May 27th, 1917. She
carried a Government commission presided over by Bishop RAFAEL EDWARDS,
a prominent Chilean ecclesiastic, who went to continue his studies on native
conditions and to distribute a large amount of materials, clothing etc. among
the members of the little island colony. Capitan de fragata J. T. MERINO was
in command of the vessel, and he as well as the bishop and the officers of the
ship did all in their power to assist us in our undertaking.

After a rather uneventful cruise our vessel anchored in La Pérouse Bay
on June 15th, and the same day we made our first excursion along the north
coast. We were bound for Hanga Roa (Cook’s Bay) but were detained in La
Pérouse on account of adverse winds; finally we resoived to cross the island

4 CARL SKOTTSBERG

on horseback, and arrived at Mataveri, the seat of the farm house, on the 19th.
In the meantime we had made some excursions in the northeastern part, where
Mt Katiki was ascended. At Mataveri we were cordially received by Mr.
Percy EDMUNDS, the manager, and were invited to to take up our quarters in
his house. Quite naturally, the natives were in a state of great excitement
over the arrival of the vessel with their much beloved bishop, the missionaries
(two Capuchin Brethren) and the many useful articles reported to be onboard;
and consequently they were rather unwilling to render us any assistance. It
was only through the kind intervention of the bishop that we were able to get
an important part of our scientific baggage, which had been left on the beach
at La Pérouse, transported to Mataveri. From our headquarters the district
round Hanga Roa and Hanga Piko etc. was visited and several trips under-
taken to Rano Kao and Orongo, the famous stone village. Further, our work
was extendend to Rano Aroi and Mt. Terevaka, the highest mountain, and
also to the south coast and to Rano Raraku, the image mountain.

We had expected to remain at least three weeks on the island, and greatly
regretted that we were ordered onboard already after a fortnight’s stay, espe-
cially as my capacity for hard work had become reduced on account of illness.
The »Baquedano» left Easter Island on July Ist.

GENERAL NOTES ON THE GEOGRAPHY OF EASTER ISLAND

The topographical features of the island are fairly well illustrated on the
accompanying map, the result of Chilean Navy surveys of later years. The
position of the observation spot in Hanga Roa is given as Lat. 27° 08' 06"S.,
Long. 109° 25’ 54" W. Mrs. ROUTLEDGEs map is based on U. S. Hydro-
graphic Office chart no. 1119, from which it differs in the position of some of
the mountains and in the geographical names. There are certain discrepancies
between this map and the Chilean one, and the difference between the latter
and the U. S. chart are still more considerable, especially in the configuration
of the northwestern part of the island. There has been some confusion in the
placing of the names, but I take it for granted that all the names used by
Mrs. R. are properly spelt and rightly placed.

The island is known to be wholly volcanic. There are no signs of recent
action, save for a couple of tepid springs below high water mark reported to,
but never seen by us. It is rather curious that both THOMSON (Smiths. Inst.
Ann. Rep. 1889, Washington 1891) and AGAssiz (Mem. Mus. Comp. Zool.
Cambridge 33, 1906) should discuss volcanic eruptions and great earthquakes
as a possible reason for the destruction of the megalithic monuments and for
the disappearance of the greater part of the population, as all signs of recent
catastrophes are entirely absent. On the other hand, not a few craters are
well preserved; sometimes they are arranged on distinct lines suggesting lines
of less resistance in the older, more or less horisontal basaltic beds, which form
the bulk of the island. The tufas and ashes of the numerous cones present a
great variation of colour contributing to lessen the monotony of the scenery.
The attention of the visitor is especially drawn to the three great Kavo. Rano

NOTES ON A VISIT TO EASTER ISLAND 5

is the native name for a mountain which contains water. R. Kao forms the
broad southwestern promontory. The greatest height of the rim was found by
the writer to be 316 m., as the result of three aneroid observations (differences
in temperature duly considered) at different occasions. AGASSIZ has 1 327 feet
or 403 m.; the Chilean map, 400 m. The crater lake measures, according to
COOKE, 2085 feet across (634 m.); this figure may be the result of a careful
observation, but seems too small. Its surface was found by me to be 120 m.
above sea level; THOMSON and COOKE (Smiths. Inst. Ann. Rep. 1897, Wash-
ington 1899), say, respectively, that it is 700 and 600 feet below the crater
rim, thus according to their figures for the latter corresponding to a height of
190 or 160 m.; taking 316 m. as the starting point we get 106 or 134 m. I
am sorry that we were unable to examine the thickness of the peat that
covers the sheet of water save for some irregular pools, which do not appear
to have decreased much in size since the photographs of the Albatross Ex-
pedition were taken. No reliable figure for the depth of the lake has been
obtained; according to COOKE, Mr. SALMON tried to sound, but at a depth
of 300 feet the line broke without having reached the bottom. I need not tell
that according to the belief of the islanders, the pool in Rano Kao belongs to
the category of famous lakes without a bottom.

The lake is partly surrounded by stands of a very robust bullrush, an
endemic variety of the wide-spread Scirpus riparius, called paschalis by Dr.
KUKENTHAL.

The country NE of Rano Kao is hilly, one of the cones being known as
Punapau (Plate 1), the seat of the hat quarry. The northwestern corner of the
island is occupied by the highest mountain, the Terevaka (Plate 2), non seldom
veiled by a bank of clouds. This name is not mentioned in Mrs. ROUTLEDGE'’s
book, where the entire high land in question is called Rano Aroi. But the
latter name only applies to the crater on the southeast slope of the mountain.
On the Chilean map appears Cerro Terevaka, separated from Rano Aroi (or
Roi) by a shallow depression, and both names were recognized by the native
JUAN TEPANO, who accompanied us to this place. The top of Terevaka did
not present any marked signs of being a crater; the height was found to be
530 m., which I believe is nearer to the truth that the figure 770 put down on
the Chilean map. I had expected to find some notable difference in the flora
of the highland, but was rather disappointed. The cryptogams, mosses and
lichens, however, played a much greater part here than in the lowlands, where
they are of a very slight importance. Rano Aroi is a very modest and shallow
rano and cannot at all be compared with the grand R. Kao. The height was
found to be 425 m. The lake is overgrown with vegetation. There is a gap
in the east wall through which, after prolonged rains, the water flows down
to another pool, which empties itself into a long, narrow fissure, crossed by
the track from La Pérouse Bay to Hanga Roa. This fissure does not seem to
have been eroded by water but suggests a volcanic origin.

The land along the north coast, from the hills backing Anakena Cove to
Katiki, is a rather flat basaltic plain, with occasional outcrops of hard rock
and strewn with innumerable sharp-edged stones, partly hidden by the coarse
grass and making walking disagreeable, more so for a person in a state of ill-

6 CARL SKOTTSBERG

health. The slope of Katiki (another name not found on Mrs. ROUTLEDGE’s
map, but frequently used) is comparatively gentle. The top was found to be
412 m. high (300 m. on the Chilean map must be wrong); it presents a rather
striking appearance, forming a shallow circular basin, perfectly dry and with
a flat bottom 5—6 m. below the rim, which is 75—80 m. across. On the north
slope is a succession of three cones, of which the northernmost is gradually
eaten away through the action of the sea. The one nearest Katiki, Vaintu
Rova, is of a light yellowish colour; the hight is 310 m. On the south slope
we came across a deep fissure, containing rain’ water and surrounded by a fine
growth of ferns. The natives, of course, all know this rare watering-place, and
I guess this is the well spoken of by Captain Cook in his second voyage:
»Towards the eastern end of the island they met with a well whose water was
perfectly fresh, being considerably above the level of the sea; but it was dirty,
owing to the filthiness or cleanliness (call it what you will) of the natives, who
never go to drink without washing themselves all over as soon as they have
done . ..» (the edition in Everyman’s Library, p. 163). North of Vaintu Rova
stands the somewhat lower Tea-tea, the »white mountain».

SW of Katiki the famous image mountain, Rano Raraku, is situated, so
ably described and illustrated by Mrs. ROUTLEDGE, who gained an intimate
knowledge of this unique place. It is shown on Plate 5. Between this rano
and the hills east of Hanga Roa there is an extensive plain, only broken by
a few higher eminences.

The visitor, even if he be not a geographer, cannot fail to notice the
absence of every trace of valley or ravine caused by the action of running
water. It is almost with surprise that one learns the figure for rainfall, 1 218
mm., the average of 8 years’ observations. This is, indeed, no small amount,
surpassing that of Juan Fernandez, where erosion has modelled the entire island
into a system of deep valleys and sharp ridges. But in Easter Island there is
no stream, no brook; only in the crater lakes water is always found. The
great scarcity of water makes the high development of the ancient culture
quite astonishing. The climate is warmer in Easter Island than in Juan Fer-
nandez, the evaporation undoubtedly much greater, the winds at least equally
frequent. Anybody will note the rapid disappearance of the water; after a
heavy rain, the ground may become soaking wet; nevertheless, some hours
later, it is perfectly dry, the result of the combined forces of the burning sun,
the strong winds, and the extreme porosity of the soil. Occasionally, water is
encountered by digging deep holes; but to dig through the hard rocks must
have been too difficult a matter for the natives. Subterraneous streams are
reported, and are, of course, to be expected; and running water has played an
important part in the formation of the numerous caves round the coast.

The vegetation is extremely poor; if one comes from Juan Fernandez,
the contrast is very striking. The island is destitute not only of wood, but of
trees, except for a few specimens in the crater of Rano Kao, where the last
stunted dwarfs of the famous toromiro (Sephora toromiro) still linger, in com-
pany with mahute (Sroussonetia papyrifera), hau-hau (or jau-jau, spanish j),
called 7rcumfetta? by FUENTES [I believe that it is 7. semztrzloba\ and ti (Cor-

NOTES ON A VISIT TO EASTER ISLAND 7

dyline terminalis f:a).' From the earliest descriptions is seen that the island
was never wooded; planted Eucalypts, Melia azedarach and other subtropical
trees do pretty well, however.

In his »Informe» (Memorias del Ministerio de Relaciones esteriores, culto
i colonizacion, Santiago 1892), P. P. TORO says: »En otro tiempo formé (i. e.
the toromiro) sin duda bosques pues en diversas partes de la isla se ven todavia
innumerables i tupidos troncos secos de dos a tres metros de alto. Parece in-
dudable que esos bosques naturales han desaparecido, secandose la mayor parte
de los arboles a consecuencia principalmente de la introduccion de animales
vacunos i ovejunos que han quebrado las plantas o les han comido la corteza.»

THOMSON and especially COOKE also speak of the numerous groups of
trees of small dimensions: »In other parts of the island may be seen, in places
in considerable numbers, a hardwood tree, more properly bush or brush, called
by the natives toromiro, all or nearly all dead and decaying by reason of
being stripped of their bark by the flocks of sheep which roam at will all over
the island»,

When Cook visited the place in 1774, he certainly did not come across
anything like a forest, for he expresses himself thus: »the country appeared
quite barren and without wood». Both ROGGEVEEN and FORSTER (A voyage
round the world. London 1777, Vol. I) assert that there was nothing like a
forest on the island. FORSTER gives a good description of the general appear-
ance. of the place; he mentions about ten species of plants, among them paper
mulberry, Hibiscus populneus and Mimosa: whether Hibiscus still occurs, I
cannot tell, but the Mimosa is certainly Sophora; this was the largest tree,
and a very small one: »there was not a tree upon the island, which exceeded
the height of ten feet». In some places, on the hills, Sophora seems to have
formed small shrubberies. FORSTER did not estimate the flora to comprise
more than twenty species, including the cultivated plants. On board the »Ba-
quedano», Bishop EDWARDS showed me a letter from Brother EUGENIO EYRAUD,
the missionary of the island, to his Superior General in Valparaiso, dated
December, 1864. He writes: »la vegetacion toda de yerbas y arbustos, fal-
tando los arboles y plantas elevadas». It is possible that Brother EUGENIO
includes the toromiro within his »arbustos». They cannot have been of any
considerable size, or he would not have pointed out that trees were missing.
A toromiro of 3 m. decidedly has the look of a small tree, not of a shrub,
especially in a place where there are no larger plants to compare it with.
TORO must be mistaken when he believes that there had grown forests on the
island not long before his arrival. Mr. COOKE does not tell if he saw the
numerous trunks himself or if he was only told of their existence.

1 In Anuario Hidrogr. de la Mar. de Chile (30) 1916, p. 55 C. DE LA Maza mentions a
small indigenous tree which he calls »tumahitiv. »La madera del tumahiti es bastante dura y
mas o menos flexible. Lo utilizan los canacas para construir yugos, arados etc. Es el unico
arbol que crece libremente en toda la isla.» The fruit is described as oval-shaped, yellow,
1 cm. long, with a stone within, and of bitter taste. There is, however, no tree growing freely
over the island. The description of the fruit suggests J/e/éa, but this is called szofazji (vide
F. FuEnTEs, Resena botanica de la Isla de Pascua. Inst. Centr. Meteor. y geofis. de Chile.
No. 4. Santiago 1913). Tumahiti sounds like a corruption for te mahute, the paper mulberry,
which certainly cannot supply materials for the implements mentioned.

8 CARL SKOTTSBERG

The dwarf trees now existing are, as has already been stated, almost
wholly confined to the crater of Rano Kao and are on the verge of extinction.
They grow not far from the lake, where the steep slope is covered with very
large blocks, which prevent the sheep from reaching them. Mr. EDMUNDs told
me about some trees which grow along the steep bluff of the eastern headland;
unfortunatly, we were prevented from visiting the place, what I hope some
future naturalist will do.

The greatest part of the island is covered with grass, wide-spread species,
either Polynesian or introduced from the old world via Chile, Tahiti or other
places. Occasionally ferns are found, also outside the rano, W/crolepia strigosa
being the most common. We discovered two species of Ophzoglossum, to which
the natives attribute medicinal qualities. Among the cryptogams are some
endemic species. A detailed account of the Flora will appear in the volume
dedicated to the Botany of the expedition.

The terrestrial Fauna is very poor; no indigenous mammals or land birds
are known. ‘There are two species of lizards, both of wide range. ‘The insect
fauna is remarkably poor; some species have been introduced by man, e. g.
cockroaches and flies, which have increased enormously.

ARCHAEOLOGICAL REMAINS: THE AHU

These structures, the burial-places — but not the only ones — of the is-
landers, have been called »terraces» by most authors. Such a word tells us
very little, while the word aku is a proper technical term, strongly and justly
recommended by Mrs. ROUTLEDGE for regular use.

There are several kinds of ahu. The most striking is the image-ahu,
which carried the now fallen statues of stone. We know that all the images
were purposely upset as a result of internal warfare, During our staty in La
Pérouse Bay we devoted some time to the inspection of the ahu. One not far
from the landing-place, close to the beach, was said to be one of the best
preserved, although not one of the largest and having supported one statue
only. This ahu was measured and described (see Plates 3—4 and text fig. 1).
The central part is 23,5; m. long and protrudes 4 m. in front of the wings; as
it stands on somewhat higher ground, it rises above the wings, in spite of the
front wall being rather low or, about 1,25 m. The front wall has a foundation
of small stones and flat slabs, followed by large, very well wrought blocks;
one of these was 2,3 m. long, 0,9 high, and 0,45 broad, another 2,45 long and
1m. high. They are closely fitted. The space behind this wall was filled
with boulders, the surface paved with larger, flat stones, making a level
platform.

The wings are larger than the centre, the east 31,2, the west 32 m., giving
as total length of the ahu 86,, m. Their front wall is as high as or even
higher (about 2 m.) than that of the centre and is similar in construction,
without being so neatly finished. The central part of the ahu, behind the plat-
form, had been disturbed, a stone wall of 7,6 m. stretching obliquely along the
fallen image. The ahu slopes gently inwards. This slope, which could be

—

NOTES ON A VISIT TO EASTER ISLAND 9

traced inland about 12 m. (measured from the front wall), is divided by a wall
(not visible above the surface), as described and figured by Mrs. R. The sea-
ward part has a surface of boulders almost free from vegetation, while behind
the dividing-line it looks like an old pavement with grass between the stones.
According to Mrs. R., the vaults for bones are found in this part. In the case
in question an open vault was to be seen in the seaward part of the west
wing; it measured I,9 X 0,7 m. with a depth of 1 m. Two slabs had covered
it. We do not know if the ahu were built with many vaults, or if accomoda-
tions for the bones of the deceased were arranged on each occasion.

eet Eben Peal RUS oc de oh Se Gia { the rage Pete lhld tans

4 A /y
tte =the wf Ly yeaa

soe

CLA TATA ae HUD taretny 7

TTL A

Fig. 1. Diagrammatic sketch of an ahu west of the landing-place in La Pérouse Bay.
Scale 1:750. s fallen image, # hat, wz stone-wall, g vault.

In the centre of the front platform one large image or moaz had stood
(Plate 4). It had been brought down by undermining the foundation stone.
This moai was the only one carefully measured:

Total length 10,2; m. Width across shoulders 3,2 m.
Length of body 6,55 m. » » head 2,6 m.

» » head 2,52 m. » > neck I.g m.

> » neck I,2 m. Circumference round shoulders 7,9 m.
Width of body ‘at base 2,7 m. » » neck 5,2 m.
Thickness of body at base 1,6 m. Length of ear 2,4 m.

Close by, the hat or crown rested flat upon the ground; its height was
1,85, its circumference 7, the greatest and smallest diameters 2,5; and 1,85 m.,
respectively. It is of the usual red stone from Punapau, but not finished. The
finished hat had a knob on top and an oval depression below; this one showed
neither on the exposed side. According to Mrs. R. the hats were finished after

having arrived at the ahu; I suppose this one never adorned the image. Could

it not be possible that the stone wall spoken of above was part of a construc-

tion on which the hat was to be rolled up to the top of the image?

Naturally, I have tried to identify the ahu described here with one of

those mentioned by Mrs. R. Its position at once suggested the Paro, of which

a sketch is communicated by Mrs. R. As far as I can see, this must be the

same (if so, it will undoubtedly be recognized by Mrs. R. from my photographs),

but there are some differences that I am unable to account for. In the sketch,
there is no trace of the stone wall alongside the image, the two parts of this
latter are far too much apart, and the hat seems to rest on its cylindrical
side. Then, there is the size of the image: Mrs. R. gives 32 feet, adding

10 CARL SKOTTSBERG

that it is the largest one ever found on an ahu and the last one to be upset.
Are there two exactly similar ahu close to each other, each with an unusually
large image? I think not. The measurements were taken by my wife and
myself with a tape 25 m. long and the figures committed to paper on the spot.

It could possibly also be the same ahu as no. 34 Punahoa of THOMSON
l. c., p. 505. He also gives the total length of the single moai as 32 feet.
However, the entire structure is said to have a length of 175 feet and a width
of 8 feet, which figures must be entirely incorrect if ahu Paro is meant.

By what kind of apparatus or devices the statues were transported from
Rano Raraku to the coast, in some cases to rather inaccessible places, remains
a mystery. The natives possessed strong cordage, and Mrs. R. has made out
that long lines were used occasionally, but veritable hawsers would have been
needed to drag the statues along over the ground in the manner imagined by
THOMSON. We have seen that there is evidence against the island ever having
produced good-sized timber suitable for rollers. THOMSON thinks that, after a
smooth road had been constructed, »the images were dragged by means of
ropes made of indigenous hemp»; »seaweed and grass made excellent lubri-
cants», He could »clearly see how it was accomplished with a large force of
able-bodied men» (p. 498). I must confess that I find it less easy to understand
how the work was done, for the least obstacle would become a serious one;
and the roads must have been made as smooth as a floor in order to serve
the purpose, the images being rather fragile. Mrs. R. has traced the few high-
ways leading from Rano Raraku to the coast; but if really the images were
dragged up to the numerous ahu all round the island, these roads cannot have
sufficed, but an elaborate network of very smooth paths was required, of which
all traces would have disappeared. It is true that seaweeds are plentiful, but
there is no species of any considerable size and I fail-to see how the quantities
required could have been brought together. It is astonishing that no tradition
on the means of transport survives. According to Mrs. R. the natives in-
variably offered one explanation: that the images were transported by the aid
of supernatural forces.

On p. 486 THOMSON discusses the possibility of a transport by sea. Near
a group of ahu he discovered a fine landing-place made by art, »admirably
adapted to the landing of heavy weights». From old drawings we know what
the aboriginal canoes were like — not a single one, as far as I know, has been
preserved to our days — but they were not strong enough to support any very
heavy weight. One might suggest that large rafts were built but, on the ether
hand, there are several ahu which are unaccessible from the shore.

Still, there is another metod to be reckoned with, although further specu-
lation on this matter may appear pretty useless. Some sort of a sledge-like
apparatus could have been constructed without the need of timber of any con-
siderable size. A sledge would slide quite well over the grass, provided that
the road was cleared from stones. A great number of people could be simul-
taneously engaged in pulling, while, if rollers were used, the image must have
been more difficult to handle. Once arrived at the ahu, a sloping causeway
could have. been built, from which the image was lowered down in position,
or, the same method could have been used that was applied when raising the

SE ee eee ee

NOTES ON A VISIT TO EASTER ISLAND Bh

images on Rano Raraku. Still, this was easier, as the statue was steadied by
the hole in the ground. For details, see Mrs. R., p. 189. Also compare what
is said above on ahu Paro: if a slope was built for the image, the same one
might have been used for bringing the crown to the top, and perhaps the wall
mentioned above will have to be explained in another and more natural manner
than the one indicated. Anyhow, to erect the statues on the platform in the
precise position required, turning their backs to the sea, must have demanded
not only great skill but also perseverance, a quality not very characteristic of
the present native population.

It is the merit of Mr. and Mrs. R. to have made out the various types of
-ahu and to have pointed out that not a few have been rebuilt. Some structures
of this kind were noted by us. A »poe-poe», not far from the western slope
of Mt. Katiki, was even sketched (another by Mrs. R., fig. 95); a stone pillar
stands on the surface, which is covered by grass.

THE IMAGE MOUNTAIN

Rano Raraku has been called the most interesting spot in the island.
Truly, the sight of this wonderful mountain with its quarries and statues is one
not likely te be forgotten. The place has been admirably well surveyed by
Mr. and Mrs. R., making. it quite unnecessary for the present writer to say
anything on the subject, but he thinks that the reproduction of some of his
photographs will be found pardonable. Plate 5 gives a total view of the
mountain from the SW, plate 6 is a familiar sight from the outer slope. On
plate 7 is shown one of the very largest statues in the quarries, no. 41 in Mrs.
R.’s diagram, also figured by her, Fig. 49; and by AGassiZ, Plate 39. Neither
of these shows more than the left part of the image. It must be about 16 m.
long, while the largest statue is just over 20. Mrs. R. finds it difficult to
believe that the latter was ever made to be launched; the same, then, can be
said of no. 41. It is hard to see where the limit for the capacity of the an-
cient islanders should be drawn. Surely there are images in the quarries that
are’ little more than rock-carvings, but I am not prepared to include the two
just mentioned under such a heading.

The two prostrate statues on Plate 8 are nos. 64—65 in Mrs. R.’s dia-
gram; they show the narrowed base. Plate g is also from the inside of the
crater, a quarry high up in the gap. It represents the heads of two images
(possibly nos. 15—-16 of the diagram), one but roughly modelled, one finished.
The first has a large »wart» on its cheek; perhaps this applies to the case
mentioned by Mrs. R. on p. 181, where it is stated that the unexpected occur-
rence of large and hard nodules in the rock could cause the whole work to
be abandoned.

It is the great triumph of Mrs. R. to have unveiled the mystery of the
scattered statues outside the mountain, which were formerly believed to have
been dropped on their way to the coast and left lying. We know now that

1 GEISELER (resp. WEISSER) describes a statue of 23 m. length (Die Oster-Insel. Berlin
1883, p. 9). ;

12 CARL SKOTTSBERG

neither the images standing on the mountain nor those found scattered over
the island were ever intended for the ahu, but that the latter lined the roads
leading from the mountain to the coast. I cannot add anything to the ex-
planation of the standing statues. If they were put up to celebrate »bird-
men», it seems quaint that not the names of these heroes but those of the
workers should have become attached to them, but such is the tradition, as
told by Mrs. R.

The rude stone implements (toki) used by the sculptors are often found.
In Hanga Roa we came across a large and very well wrought stone adze ofa
rather international type, but not found in any of the accounts on the island.
It had been picked up on the seashore at low water and is quite incrustated
with the shells of animals. It measures 20 cm. (Plate 14, fig. 1). A few stone
chisels were also obtained; two are figured on Plate 14, fig. 2, 3.

REMAINS OF HOUSES AND PLANTATIONS

Foundations of old houses are seen in many places, and several were
noted on our excursion to Mt. Katiki. Of one a sketch was made, also showing
the paved area in front, but without foundation-stones for a porch. None of
these dwelling-houses are left. Between Mataveri and Hanga Roa are a couple
of grass huts (Plate 10) which give us a faint idea of what the old houses
were like. They are small, lack the stone foundation, and have the entrance
at one end.

There is another kind of structure in the shape of low, very strongly
built towers of stone which cannot fail to arouse the curiosity of the visitor.
The present people do not seem to be sure as to their former use. A fine
tower at the landing-place in Hanga Ho Orno (La Pérouse) is shown on Plate
II, another is figured and described by Mrs. R. (p. 218, fig 87), a third one,
in a ruined state, by THOMSON (p. 484). The first-mentioned has a height of
3,4 m. and is 6,5;—7 m. across at the base. The only entrance is 0,65 m. high
and 0,9 wide. The roof is vaulted inside; outside, the wall ends in a girdle of
stones. The size of the stones bears witness of the prehistoric era and of the
makers of the great ahu. Mrs. R., on the authority of some resident, explains
these structures as »look-out towers whence watchers on land communicated the
whereabouts of the fish to those at sea; these contained a small chamber
below which was used as a sleeping apartment» (p. 218).

It sounds strange that these solid towers should have had no other pur-
pose than to serve as look-out stations for fishermen. In order to keep a good

look-out, presumably to follow the movements of shoals — it has not been
proved that there is any fish here of the social type, and Mrs. R. states that
»fish are not plentifulo — an observer must seek an eminence dominating a

considerable space of water. THOMSON tells us that from the towers the move-
ments of the turtles were watched. The observer must keep outside the
tower, or on the top of it, not a very comfortable place. Really, the tower
itself would have been little more than a refuge in bad weather and during
the night, but for such a purpose a much simpler structure would indeed

NOTES ON A VISIT TO EASTER ISLAND 13

suffice. Thus the importance of the tower with its lower apartment appears
to stand in no reasonable proportion to the vast amount of labour required to
build it. We should perhaps remember that the permanent dwelling-houses
were much more fragile. The tower suggests some kind of fortress, with a
chamber for stores or treasures: it would be easy to defend the entrance.
Speaking of the narrow entrances to the Orongo houses, THOMSON remarks
(p. 483): »The low contracted entrances were used here as well as elsewhere
for defence. Factional fights were common, and it was necessary that every
house should be guarded against surprise and easily defended». He adds:
»Another reason might be found for making the openings as small as possible,
in the absence of doors to shut out the storms». But, at least at Orongo,
there were plenty of slabs suitable for doors if wanted.

ST LF
.@ a: Rina

ie WN
- NI f J, Oo
ANA
cont M3 oe) ro
Sotelfess si re

2 wo ®2 y %
fy MY) /1/ Ye Wij WAY ny’ VF

, Ay
77 .

Fig. 2. Diagrams of old plantations; @ seen from above; 4 three types of shelter, in section.
1 = bananas, 2 = Melia azedarach, 3 = Andropogon halepensis. Scale 1: 250.

On the other hand, an enemy could pull down the roof over the occupants.
Also, we should expect to find a communication between the tower and the
underground chamber, which we have not found. So, after all, this theory may
not hold good. Perhaps prisoners were locked up in the towers, where they
could be easily guarded till the hour arrived to put them to death. But it is
also necessary to consider whether these buildings may not have had some
relation to unkown rites. Did they have a ritual purpose, we do not need to
wonder about their elaborate structure, as natives may invest any amount of
labour in connection with religious or other ritual buildings.

There is a description and figure of a similar building in La PEROUSE’s
Voyage, reproduced by STOLPE in Ymer, 1883. He states that they are only
found on the top of Rano Raraku, which is, of course, a mistake. They were
oval in shape; close by was the underground chamber with its separate en-
trance. In several respects, especially concerning the ahu, there are great dis-

14 CARL SKOTTSBERG

crepancies between the statements and illustrations of the old navigators and
the results obtained by modern explorers. A critical examination of the old
stories would be welcome, and it is to be expected that Mrs. ROUTLEDGE will
undertake to scrutinize the entire literature. There are, e. g., in La PEROUSE’s
and PINART’s narratives designs of ahu which do not at all correspond to
modern descriptions or photographs.

In the vicinity of Hanga Ho Orno we saw many remains of native planta-
tions: ‘They are of ‘several types. One, seen “in fig. 2 a, is probablggyones
later date, as the material has been taken from an ahu, the front wall of

Sais
Li Hhty A
a

Fig. 3. a. Two »bird-men» on rock at Orongo (height of the rock 1,6—1,7 m.) &. Incised marks
on door-poost at Orongo.

which forms the back wall of the garden. Circular miniature gardens are
represented in fig. 26. The need of shelter and moisture is well unterstood.
Melia was said to be grown for the sake of the timber. Probably it is of recent
introduction.

ORONGO AND THE BIRD CULT

Mrs. ROUTLEDGE devoted much time to the survey of the Orongo village,
and as a detailed plan was made and every house measured and described, I
shall content myself with a few short remarks.

The last house (if I remember right) towards the gap of the crater rim,
close to the sculptured rocks, had one door-post with incised carvings left,

NOTES ON A VISIT TO EASTER ISLAND 15

vide fig. 3 6. This house was’ pointed out to us ‘as the house of Ariki. The
ariki was the chief of the Miru clan, the authority on the script (i. e. the
variki-mau», vede R. p. 241; all Miru were also called ariki). Now, the same
design was found on a skull in the possession of the schoolmaster, Mr. I. Vives,
and this skull was attributed to an ariki. Unfortunately, the owner did not
want to part with his treasure. The design is unlike the one figured by Mrs.
R. (fig. 96) of another Miru skull.

The Bird Cult is described, with full details, by Mrs. R. Special attention
is paid to the rock carvings. I sketched a couple of the »bird-men» (fig. 3 a).
Their meaning is not known with certainty. Mrs. R. believes them either to
represent one of the egg-gods (they were spoken of as »Make-make») or made
to immortalize the bird-men, the winners of the egg-race; she finds the latter
explanation more probable. I have not been able to form an independent
opinion, The same carvings are seen on a flat stone opposite Orongo, marking
the place where the path descends into the crater of Rano Kao.

All that is left of prehistoric remains, at least of the large ones, will
remain on the island. Shortly befgre our visit a law was passed prohibiting the
removal of statues etc,, so that we had to abandon our idea of bringing home
a small image presented to us by one of the residents. The »Mana» was just
in time to rescue the small but unique statue from Motu Nui.

WOOD CARVINGS

The famous wooden statues as well as other pieces of carving are gone
from the island for ever. What is offered to passing visitors is not worth
mentioning. The art is gone. One old moai-miro, in a very much decayed
state, had been discovered in a cave after the departure of the »Mana». It was
presented to Bishop Epwarps. In 1908, while staying at Valparaiso, a Swedish
captain, Mr. G. KARSTROM, who had been shipwrecked on Easter Island many
years before, presented me with two beautiful wooden images, one of which
is in the Etnographical Museum in Stockholm; the other is owned by a private
person.

HOUSEHOLD GOODS, WEAPONS, ETC.

Very little of this kind is now to be encountered. Sticks used for net-
knitting are available, and so are baskets or rather bags made of bullrushes
(figured by THOMSON on Plate 51). There are still some people skilled in the
preparation of tapa cloth from the mahute and of strings from the hau-hau,
and we had samples made for the collection. Curiously enough, GEISELER
does not mention the latter plant, but states that all the cordage, fishing-nets
etc. were made from the bullrushes.

It is generally stated that the islanders never possessed any earthen-ware.
Contrary to this, RUTLAND (Transactions New Zeal. Inst. 29, 1896) says that

16 CARL SKOTTISBERG

the earliest discoverers had seen »rude earthen-ware» on the island, a statement
due to some misinterpretation. Of stone implements, besides the toki, and the
stone adze and chisels, we got one fish-hook, very neatly wrought but unfor-
‘tunately not complete, as the point is missing (Plate 14, fig. 4). There is a
drawing of one of these hooks in THOMSON’s report (Plate 58). Another curious
article is the spherical stone ball, Fig. 5 on Plate 14. It shows two holes
which communicate so that a string can be passed through, and may have
been worn as an ornament. No explanation was offered. Perhaps it is a
»fetish-stone». THOMSON has described and figured many such stones, but
none of them present any likeness to this one.

The object on Plate 14, Fig. 6 is not, as might be suspected, a broken
spear-head or szafaa, but has been given its present shape on purpose. It fits
well into the hand and may have been used as a knife or scrape. But if it was
used with a handle, my explanation may not be satisfactory. Spear-heads are
commonly found in the soil and also manufactured to satisfy the demand of
visitors. Two, of an ordinary type and apparently old, are seen on Plate 14,
Figs. 7, 8. According to THOMSON there were at least nine kinds, all with
different names, a statement well needing the corroboration of Mrs. R.

ORIGIN OF THE PEOPLE

The history of Easter Island is full of mystery, but I think that Mrs. R.
has come pretty near the solution of some of the problems. She has drawn
some important conclusions from the legendary traditions still alive. <A tale
of two different races and two successive colonizations runs through the old
legends. The anthropological evidence seems to be in favour of a double origin,
Melanesian and Polynesian. The comparative studies of the Bird Cult in the
Solomon Islands and Easter Island (by H. BALFOUR, vzde Mrs. R.) seem nothing
less than convincing. The bird represented in the numerous carvings, paintings
etc. of Easter Island is not the holy bird of this place, but the frigate bird,
worshipped in the Solomon Islands. The bird figures were called penguins by
LEHMANN (Essai dune bibliographie, Anthropos, 1907), which undoubtedly
must be a mistake, especially as penguins hardly ever visit these waters.

If we sum up the results obtained, there is evidence that the Easter Is-
landers are of a twofold origin and that, after the Melanesian immigration, a
Polynesian immigration followed. The population now tends to assume a
multicoloured aspect; there has been a late influence from Tahiti (so we were
told) and various white men have contributed towards the »amelioration» of the
race. The young girl figured on Plates 12—13 was said to be of »pure Easter
Island race», but whether representing a Melanesian or Polynesian type, I am
unable to tell.

A critical examination of the language would be of interest. Many words
are the same as in the Maori or other Polynesian tongues, such as maunga
(mountain), szahute (paper mulberry), 7“ (Cordyline), cusmara (sweet potato) a. o.
A large vocabulary, collected by Padre ROUSSEL, was published in Santiago,

NOTES ON A VISIT TO EASTER ISLAND 17

1917, but it must be used with much criticism, as it contains many Tahitian
words and also corrupted English, French or Spanish; good examples are anzo
(agneau) and mutone = sheep, hzmene (hymn) = to sing, teperanate = serpent,
tokint = stockings, “aporo (diablo) = devil, vretute = virtue, given without
reservation. A closer look reveals that the material is not at all so rich as the
number of words would indicate, for the author has invented hundreds of ex-
pressions for ideas wholly unfamiliar to the aboriginal soul, by combining the
words and extending their meaning in a most improper manner, e. g. expres-
sions for cabin, desert, doctrine, palace, river, saint, W. C. etc. etc. to quote
a few obvious examples, of which scores could be given. This is, I believe,
a common missionary method to enrich the language with ideas and expressions
necessary for the translation of religious and other books, but otherwise never
used by the natives.

Concerning the name of the sweet potato, see below. It has been ad-
vanced as indicating an American influence previous to the Columbian era.
RUTLAND (I. c.) thinks that the ancient monuments bear witness of a constant
communication between the island and Peru and Mexico: »from hence architects
of Easter Island may have been derived».

CULTIVATED PLANTS

If we knew the history of the cultivated plants, many a mystery related
to the history of mankind would be solved. But, unfortunately, discussion
often begins with the original home of the wild parents of these plants, and
there it also ends.

The first record of domesticated plants in Easter Island is that of Roc-
GEVEEN, the discoverer of the island or, at least, the first white man to set
his foot upon it. He makes the following statement on p. 120 (De Reis van
JacoB ROGGEVEEN. Worken uitgeven door de Linschoten-Vereenigung 4.
IgII): »en toegebragt worden alles wat sy hadden, bestaende en boomvrugten,
aardgewasch en hoenderen», that is, tree-fruits, soil-fruits (rootcrops) and hens;
and, farther down: »want na verloop van een kleynen tijd bragten sy eene
menigte van suykerriet, hoenderen, ubaswortelen en bananas», that is sugar-
cane, bananas and ubas-roots. But what is wéas? Most likely the same word
as the Malesian ubi (uwi, huwi), yams (Dzoscorea alata), now called uf in the
island. All these plants are of Old World origin and have spread from the
Indo-malayan region over the Pacific. According to FRIDERICI, the same word,
in a corrupted form, is current in South America: »Dieses Wort schlagt eine
Briicke iiber den grossen Ozean: es gehdrt als Op unter der Bezeichnung 'siisse
Kartoffel’ zum Sprachschatz der Chimu, des kiistenbewohnenden Kulturvolkes
westlichen Siidamerikas» (Wiss. Ergebn. seiner amtl. Forschungsreise nach dem
Bismarck-Archipel im Jahre 1908). But the bridge in question seems to be weak.

The word cumara is used for sweet potato (/pomaea batatas or Batatas
edulis) from New Zealand through Polynesia to Easter Island. According to
CHEESEMAN (Manual of the New Zealand Flora) the Maori introduced the plant
from Polynesia when they colonized the country (supposingly 1350—1400), and
it was described by SOLANDER as Convolvulus chrysorhizus, now reduced to

2 — 20199. The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I.

18 CARL SKOTTISBERG

a synonym of I. batatas. This plant is universally considered to be of Central
American origin, although wild plants are not found nowadays, nor is the more
precise locality known where they grew. We must consider whether the same
species’ was not originally a native both in America and in the Polynesian region
or, whether the cultivated forms were not derived from more than one wild
species, so that it is unnecessary to suppose that the sweet potato was intro-
duced to Polynesia from the American coast. Yams is obtained from forms of
several wild species characteristic of different continents.

If this theory holds good, we should expect to find different names for
_the sweet potato on the two sides of the Pacific. But according to R. LENZ
(Diccionario etimoldjico. Santiago 1910), the word cwmara is found in the
Quichua language; it is not indicated as the principal name of the sweet potato,
which is apzchu, but nevertheless used, according to this author, for a »clase
parecida» of the camote, thus for some form of the same plant. From this
fact some people would conclude that, as the plant is American and called
cumara by the Quichua, it was introduced to Polynesia under the same name
long before the Columbian era. It is useless to discuss this matter any further
till we know more of the history of the camote and also, whether the word
cumara in Quichua really applies to the true sweet potato and, if such be the
case, belongs to the original Quichua language or has been introduced through
the Europeans. If old communications existed between America and Polynesia,
many other proofs must be found. Much has been written about old land-
bridges across the ocean, considered by some naturalists to be indispensable
for the explanation of the distribution of animals and plants. But generally
their existence was supposed to have ceased long before the age of Man. Only
HALLIER (Uber frithere Landbriicken, Pflanzen- und Vélkerwanderungen zwi-
schen Australasien und Amerika. Mededeel.’s Rijks Herb. Leiden 13, 1912)
gives them a longevity sufficent to let people march across. I am afraid that
such bridges rest on a very unstable foundation.

To return to the sweet potato, we have seen that it is not mentioned by
ROGGEVEEN as enisting in Easter Island in 1722. Cook and FORSTER found
it in cultivation. At that time also Broussonetia, Thespesia (also Triumfetta?)
and toromiro were cultivated in addition to taro, bananas and sugar-cane.
According to tradition all of them were brought by Hotu Matua’s party, the
first settlers. The barahti mentioned by F. VIDAL GORMAzZ, Jeografia nautica,
p. 177 (Anuario Hidrogr. de la Marina de Chile, 7) is, to judge from the de-
scripton, the same as the hau-hau. The calabash mentioned by THOMSON,
p. 535 is Lagenaria vulgaris. At present, the following food-plants are cultiv-
ated: sugar, wheat, Indian corn, taro, pineapple, yams, bananas, white mul-
berry, figs, maniok, oranges, lemons, grapes, peaches, quince, plums, beans,
sweet potatoes, tomatoes, melons, artichokes and lettuce, but several of these
only on a very small scale and exclusively in the garden of Mataveri. Some
tobacco is also grown. I do not know what THOMSON means by the »two
varieties of indigenous hemp», as there is no plant of this kind either in a
cultivated or abandoned state. The cordage has always been prepared from
the hau-hau, as far as I have been able to ascertain. Nor does Mrs. R. refer
to any such plant, nor to the hau-hau.

NOTES ON A VISIT TO EASTER ISLAND 19

THE FUTURE OF THE ISLAND

The power of resistance of the Easter Island people was definitely crushed
through the Peruvian slave raids, and through missionaries and farmers they
lost the strength which lies in the possession of an aboriginal culture. Their

removal to Hanga Roa, where a village was formed, was very unlucky, as it

meant giving up many small plantations and induced the people to lead a
parasitic life, expecting everything from their new rulers. Although they have
left so many wonderful monuments to bear witness of earlier busy days and
of a people of warriors, they are now, with few exceptions, lazy beggars. In
part this may be due to their pronounced feelings of animosity against the
intruders, as they regard themselves as the true possessors of the island. It
appears that ever since the establishment of a farming company the state of
affairs has never been lucky, and Mrs. R. has an interesting tale to tell of an
anxious time. I do not at all believe that the present manager is to blame,
for we got the impression that he is as well liked as any white man in his
position can expect to be. In Chile, nobody seems to have taken much notice
of the distant colony till Bishop EDWARDS entered the field. During his first
visit, in 1916, he informed himself of the state of things, and he returned in
1917 invested with powers to put everything right if he could. Among other
things he wanted to take up war against the leprosy.’ Not quite 5 % of the
population suffer from this disease; they are confined to a colony some distance
from Hanga Roa. Apparently it is not very contagious, for the isolation is
not quite effective. The surgeon of the »Baquedano», Dr. G. LONGO, examined
almost every soul, but only one or two new cases were discovered. As acco-
modations for the most advanced cases had been wanting, the vessel this time
brought materials for the construction of a small hospital which was to be
erected by the new »subdelegado» or governor. Captain MERINO carried in-
structions to examine the claims against the company, and a meeting was held
where the natives put forth their demands. I understood that the Company
was said to have taken possession of more land than it was entitled to and
that the natives wanted it to be restored. Officers went round with natives
who indicated the seats of their former homes and fields, and parts of the land
were measured. The scheme was, I think, that certain parts should be restored
to the old owners, that the village should be abandoned, and that the natives
should move into »the camp» in order to become selfsustaining. A certain
amount of native labour should be granted to the manager at a fixed rate of
pay. I have had no chance to learn how far the realisation of this humanitary
scheme has advanded; nor would I venture to foretell if it is likely to meet
with success.

EASTER ISLAND AS A FIELD OF PSEUDOGEOGRAPHICAL SPECULATION

Finally, I shall make a few remarks in addition to what Mrs. R. tells us
(p. 290) of the theosophists’ views of Easter Island, which are based on errors

1 The surgeon of the »Mohican», Dr. Cooke, does not mention this malady as existing

in the island in the year 1886. It was imported from Tahiti.

20 CARL SKOTTSBERG

regarding the existing monuments. Last year a small book appeared, entitled
»Det sunkne kontinent (Atlantis)», by a Norwegian, C. SUND (Copenhagen 1919),
where also the supposed Pacific continent is spoken of. As might be expected,
Easter Island forms an important item. With my permission, two of my photo-
graphs were reproduced. No doubt Mr. SUND regards himself as excused for
his mistakes, for he has quoted various obscure authors; but it must be regretted
that he should not happen to draw from a single reliable source, not even
from my popular description, which was known to him. Mr. SuND tells us of
the Egyptian influence in Easter Island, of the enormous foundation walls and
ruins of temples; almost every mountain had sculptured designs of goods, fishes
and pyramids, the cave paintings were in the Toltec or Egyptian style, etc.
There are 300 tablets with script on the island (if it were but true!), waiting
to be deciphered. On the mountain terraces are fortresses with walls up to
80 feet high. The pyramid is the architectonical principle, built as the Egyptian
one, even with the same kind of cement. All materials, bricks, glass, porcelain,
everything was known in Easter Island; religion, symbols and habits were the
same as in Egypt, only, the culture of the island was older. There are fan-
tastic groups of statues roundabout, gods of hard store with faces up to 25
feet high, in the highlands there are images on high stone pillars or staircase-
like foundations, and with square hats of stone, most of them covered with
script in a probably forgotten language. Round them are the remains of large
walls and buildings, so they probably stood in vast temple-yards. And so
forth. No wonder that Mr. SUND draws the most surprising conclusions. Now,
this must not be taken too seriously and will do no harm in scientific circles.
The general reader, however, will get a rather curious idea of Easter Island.
I dare say the place is remarkable enough in itself and need not be glorified
by such fantastic inventions.

Finally, I wish to express my sincere gratitude for kind assistance to the
Commander and Officers of the »General Baquedano», to Bishop RAFAEL ED-
WARDS, Mr. PERCY EDMUNDS, Mr. I. VivEs and Baron ERLAND NORDENSKIOLD.

Explanation of Plate 14.

1. Stone adze, no: 19, I: 307; not quite t/e.

2. Stone chisel, no. 19. 1. 320; almost 9/1.

3 » Pe ENO SOs 1.5 20s PEN:

4. Fish-hook of stone, no. 19. 1. 325; almost %/s.

5. Stone ball, no. 19. 1. 309, not quite nat. size.

6. Knife or scrape? no. 19. I. 315, not quite nat. size.
7a mopeat-nedad, No. /19. 1. 313-6273.

8. » PMOL Os al 30 Ti Be

The originals in the Museum, Gothenburg.

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Nat. Hist. Fuan Fernandes and Faster lst. Vol. J. PLATE

Photo by K. Backstrém

Same girl as in Plate 12.

Nat. Hist. fuan Fernandez and Easter Isl. Vol. J.

Rese

1A.

Photo by O. Thulin
Stone implements. For explanation, see last page of text.

THE NATURAL HISTORY
SPreLTUAN FERNANDEZ
Sey) EASTER ISLAND

PDOEPBED MEY DR: CARL SKOTTSBERG

VOL. I
GEOGRAPHY,,;GEOLOGY,
ORIGIN OF ISLAND, LIPE

PART TI

2. T. H. HAGERMAN: Beitrage zur Geologie der Juan Fernandez-
Inseln.

3. P. QUENSEL: Additional Comments on the Geology of the Juan
Fernandez Islands.

4. C. SKOTTSBERG: A Geographical Sketch of the Juan Fernandez
Islands.

UPPSALA 1954

ALMQVIST & WIKSELLS BOKTRYCKERI AB

INE’

BO’

2. Beitrage zur Geologie der Juan Fernandez-Inseln.

Von

TOR H. HAGERMAN.

Mit 12 Textfiguren.

Die Schwedische Pazifik-Expedition 1I916—1917 unter der Leitung von
Professor C. SKOTTSBERG brachte unter anderem eine Gesteinsammlung von
den Juan Fernandez- und Oster-Inseln zuriick, die dem Mineralogischen Institut
der Hochschule zu Stockholm zur Bearbeitung ubergeben wurde. Bei der von
mir vorgenommenen Untersuchung von etwa 50 Handstiicken von den Juan
Fernandez-Inseln stand mir ausserdem QUENSEL’s Material von einer friiheren
Beschreibung! desselben geologischen Gebietes zur Verfigung.

Die Juan Fernandez-Inseln liegen zwischen 33 und 34° S. Br., 660 km W.
von Valparaiso. Die Inselgruppe besteht aus zwei grosseren Inseln, Masatierra,
Flacheninhalt ca. 95 qkm und W. von derselben Masafuera, 85 qkm. Nahe der
erstgenannten liegt eine kleine Insel St. Clara, 5 qkm.

Der Gebirgsgrund dieser Inseln besteht ausschliesslich aus Effusivgesteinen.
Eine exakte Altersbestimmung derselben kann kaum gemacht werden, da keine
Sedimentgesteine vorhanden sind. QUENSEL? nimmt an dass die vulkanischen
Gesteine kaum Alter als jungtertiar sein dirften, und es ist seitdem nichts be-
kannt geworden, was fiir eine veranderte Auffassung sprache.

Die Inselgruppe ist einer kraftigen Erosion ausgesetzt gewesen, sodass die
urspriinglichen Vulkankegel nicht mehr zu erkennen sind. Besonders auf Masa-
fuera, wo die Wasserscheide weit nach W. verschoben liegt, hat sich eine aus-
gesprochene Cafion-Landschaft gebildet. Die wilden Terrainformen sind deutlich
aus den zahlreichen Photographien zu erkennen, von denen viele in SKOTTSBERG'Ss
Reisebeschreibung*, wie auch in Vol. I und II dieses Werkes ver6ffentlicht
worden sind.

Masatierra.

Die Untersuchung des Materials von Masatierra hat erneut bestatigt, was
QUENSEL bereits hervorhebt, namlich, dass die Gesteine untereinander chemisch
und mineralogisch nahe verwandt sind, und sich im wesentlichen nur strukturell
voneinander unterscheiden. Sie kdénnen als verschiedene Erstarrungsformen ein

1 P. D. QUENSEL, Die Geologie der Juan-Fernandez-Inseln. Bull. Geol. Ups., Vol. XI,
Pp. 253-290.

aoe. Cp. 250.

3 C, SKOTTSBERG, Till Robinsonén och varldens ande. Stockholm 1918.

3—248. The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I.

22 TOR H. HAGERMAN

und desselben Magmas betrachtet werden; spaterhin sind allenthalben stellen-
weise sekundare Veranderungen infolge hydrothermaler Prozesse entstanden, die
abweichende Ausbildungen hervorgerufen haben.

Die Gesteine sind im primaren Zustande durchwegs mehr oder weniger olivin-
reiche Basalte mit der Zusammensetzung: Olivin, Pyroxen, Plagioklas, Magnetit,
Ilmenit und oft ein wenig Glasbasis.

Charakteristisch fur alle diese verschiedenen Teilen der Insel entnommenen
Proben ist besonders die Zusammensetzung des Pyroxens. Dieser besteht aus
einem Titanaugit, sofort erkennbar an seiner schwach rotvioletten Farbe und
starken Dispersion der optischen Achsen. Doch ist zu bemerken, dass derselbe
nicht zu den extremsten Typen gehort.

Auch die Feldspate weisen eine konstante Zusammensetzung auf. Sie sind
fast alle zwillingsgebildet nach dem Albit- oder Karlsbader-Gesetz. In Schnitten
senkrecht zu M zeigen die Albitlamellen eine max.-Auslésung von 32—33°, in
einigen Einzelfallen diese Werte mit héchst 2° variierend. Der Feldspat ist also
ein Labrador von einer Durchschnittzusammensetzung Ab, .An;..

Da die Basalte also mineralogisch einander nahe verwandt sind, wurden sie
hauptsachlich nach der Struktur in folgende Typen eingeteilt:

basaltische Laven, (teils dichte, teils grobkérnig doleritische, teils schlackige)
Tuffe, hydrothermale Umwandlungsprodukte.

Basaltische Laven.

Diese Gesteine bilden, wie friihere Verfasser bereits betont haben, den Haupt-
bestandteil der Insel. Auf Grund des vorliegenden Materials konnte man den
Verlauf der verschiedenen Lavastrome und deren Neigungsverhaltnisse nicht be-
stimmen und so auch keine Klarheit tber die Eruptionsstellen erlangen.

Die Fundstatten der dem Verfasser zur Untersuchung vorliegenden Hand-
stiicke sind ziemlich gleichmassig uber die Insel verteilt. Von den zu den Laven
gehorenden Gesteinsproben sind ungefahr ?/, sehr pordse und schlackige Typen,
wahrend !/, (6 Stck.) dichte, dabei gleichmassigere und feinkérnige Gesteine
darstellen.

Die Struktur dieser Gesteine ist im allgemeinen hypokristallin porphyrisch.
In einigen Fallen, besonders bei den dichten Typen, kommt es vor, dass Glas-
basis ganz fehlt.

Mit Bezug auf die mineralogische Zusammensetzung der basaltischen Laven
sind folgende Mineralien beobachtet worden: Olivin nebst dessen Umwandlungs-
produkten, die vorgenannten Pyroxene und Plagioklase sowie Erzmineralien (Mag-
netit und Ilmenit).

Die Einsprenglinge sind Olivine und ihre Umwandlungsprodukte sowie
Feldspat. Die Augitkorner kénnen sich zuweilen der Grodsse der Einsprenglinge
nahern.

Die Olivineinsprenglinge erreichen ihre grésste durchschnittliche Ausdehnung,
ca. 2 mm, in Proben, die von dem nordlichen Ufer der Padrebucht stammen.
Gerade diese Einsprenglinge zeigen meistens in einer scharf begrenzten Zone
die von QUENSEL! friither beschriebene Iddingsitumwandlung. Aus Dunnschliffen
der genannten Gesteine ist klar ersichtlich, wie die Umwandlung von den Ran-

he 3 jo, Ao.

BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ-INSELN

dern nach der Mitte zu ausgegangen ist und sich 0,o2—0,o3 mm in den Olivin
hineinerstreckt hat. An manchen Stellen ist die Umwandlung langs der Spalt-
risse vor sich gegangen, wahrend anderweitig die Durchgange merkwurdigerweise
vollkommen unveranderte Teile der Olivinkerne durchqueren. (Vergl. Fig. 1.)
In der Gesamterscheinung des obengenannten Praparates méchte der Verfasser
die Iddingsitbildung als das Resultat einer von aussen kommenden chemischen
Beeinflussung ansehen. WASHINGTON! verweist die Iddingsitbildung bis auf mag-
matischen Ursprung zuriick. Man braucht vielleicht nicht so weit zu gehen, da
dieselbe ebensogut einer hydrothermalen Umwandlungsperiode zugeschrieben wer-
den kann.

Fig. 1. An den Randern iddingsitumwandelte Olivine. Olivinbasalt von der Padrebucht. —
Vergr. 56x. Photo HJ. OLSSON.

In einem anderen Gestein von der Padrebucht, das etwas feinkOrniger ent-
wickelt ist, treten auch Feldspateinsprenglinge von 0,;—1 mm Lange auf, neben
Resten von Olivinen. Der Olivinumwandlungsprozess hat hier ein anderes Pro-
dukt hervorgebracht, namlich gewohnlichen Serpentin. Dies ist auch bei den der
Grundmasse angehorenden Olivinkérnern der Fall. Dieselben sind vollkommen
als Serpentin ausgeflossen. Auch der Augit scheint an den Randern etwas ange-
griffen zu sein. Beinahe identisch entwickelt sind zwei andere feinkornige Basalte,
der eine von der Mitte des siidlichen Ufers, am Fuss des Yunque, der andere von
einem Gebirgsriicken (385 m i. M.) SW. von Tres Puntas, W. von der Villagra-
Bucht, herstammend. Iddingsit- und Serpentinumwandlungen der Olivine sind,
wie ich besonders hervorheben mdéchte, niemals in ein und demselben Gestein
gleichzeitig angetroffen worden.

Die beiden erwahnten Gesteine von der Padrebucht und an beiden Seiten
von Villagra zeigen eine eigentiimliche primare Struktur der gleichkornig ent-
wickelten Grundmasse, indem die Plagioklase mit einer mittleren Ausdehnung

1 Italian petrogr. sketches. Journ. Geology. 4 (1896), p. 835—836.

24 TOR H. HAGERMAN

von 0,2 mm sich vielfach radialstrahlig mit den Augitindividuen geordnet haben.
Ist man erst einmal auf diese spharolitahnlichen Bildungen aufmerksam gewor-
den, so findet man sie haufig hauptsachlich in den feinkérnigsten Proben dieser
Gesteine ausgebildet. Besonders schone Beispiele hiervon zeigt ein Handstiick,
das der »Pyramide», einem Berggipfel ungefahr in der Mitte der Insel unweit
des SELKIRK-Denkmals, entnommen wurde. (Fig. 2.)

Eine sehr ahnliche Erscheinung ist von REITER! beschrieben worden. Er
schmolz 45% Albit, 45% Augit und 10% Magnetit zusammen. »Der Schliff

ES, Pyroxen (= Plagicklas GF Magnetit

Fig. 2. Plagioklas-Augit-Spharolit. Basalt von der »Pyramide».— Vergr. 250%.
Zeichnung vom Verf.

einer durch 7 Stunden abgekihlten Schmelze zeigt eine sphariodale Anordnung
der Kristalle, in dem magnetitreiche Kerne von Glaspartien mit einzelnen aus-
geschiedenen Augit- und Plagioklasleisten und Kristalliten umgeben sind. Die
Wiederholung des Versuches bei 30-stiindiger Abkuhlung ergab eine Schmelze
mit kérnig-porphyrischer Struktur.» Zweifelsohne ist dies auch in dem vorliegenden
Falle zutreffend, indem die spharolitfiihrenden Laven einer raschen und ungestorten
Abkihlung ausgesetzt gewesen sein dirften.

In Anschluss an diese Spharolite seien hier die in Fig. 3 abgebildeten kreuz-
formig liegenden Olivinkristalle erwahnt. Der Dimnschliff entstammt einem etwas
grobkornigen Gestein von Bahia Cumberland. Wie aus der Figur deutlich her-

1 H. H. REITER, Experimentelle Studien an Silikatschmelzen. Neues Jahrbuch. Beil. Bd. 22
(1906), p. 197.

BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ-INSELN 25

vorgeht, handelt es sich um eine skelettartige Ausbildung der Kristallindividuen.
Eine gesetzmassige Verwachsung der verschiedenen Individuen habe ich nicht
nachweisen konnen.

Im Vaqueriatal tritt, wie aus der untenstehenden Photographie (Fig. 4) er-
sichtlich, ein fast horizontal liegendes Gestein auf. Nach ihrer grobkristallinischen
Struktur zu urteilen, sind diese Basalte
méglicherweise als intrusiv aufzufassen.’
Das nur an zwei Seiten zugeschlagene
Handstiick ist vorziiglich durch Schrump-
fung unter rechtem Winkel zerkliiftet.
(Fig. 5.) U. d. M. zeigt dasselbe ein un-
verandertes hochkristallinisches Aussehen.
Reichlich albitlamellierte Feldspatleisten,
durchschnittlich ca. 2 mm lang, bedin-
gen mit Olivin- und Titanaugitkristallen
eine ophitische Struktur. Ausser diesen
Mineralien habe ich nebst Magnetit hier
und da ein Biotitkorn gefunden. Moglicher-
weise erstreckt sich dieser Basalt bis zur
Cumberland Bay, wo eine ahnliche Aus-
bildung von QUENSEL? beschrieben wurde.
Auch das Material SKOTTSBERG’s enthalt
eine ahnliche Probe von dort, in losem
Block gefunden. Der Mineralbestand der
beiden letztgenannten Handstiicke ist der- _.. k ne

Z § ¥ } Fig. 3 a. Zentrisch angeordnete Olivine. —
selbe wie jener der Vaqueriaprobe nur Vergr. 56x. Verf. phot.
mit dem Unterschied, dass kein Biotit vor-
handen ist. Auf Grund seiner grobkristallinischen Struktur muss das obenerwahnte
Gestein zu den doleritischen Basalten gerechnet werden.

Feinkorniger, aber im tibrigen dem vorgenannten Gestein vollig gleich, ist
der bei » Tres Puntas» genommene Basalt. Die Handstiicke bestehen aus langen,
schmalen, dreiseitigen Prismen.

Unter den schlackigen Laven weisen einige eine auffallende Analogie zu
rezenter Oberflachenbildung auf. Besonders ist dies der Fall bei einem sehr po-
rosen glasreichen Gestein vom Ufer s. von Yunque. Der vorerwahnte, auf dem
Gipfel der »Pyramide» befindliche dichte Basalt hat ein schlackiges und glasiges
Lavabett als Unterlage.

In einigen anderen der schlackigen Gesteine sind die Locher mehr oder we-
niger mit Opal, Chlorit, Serpentin und Calcit ausgefiillt.

1 Vergl. jedoch QUENSEL p. 263—264.
SAL C2 pai263:

26 TOR H. HAGERMAN

Tuffe.

Unter dem mitgebrachten Material befinden sich zwei Proben von ausge-
sprochenen Tuffen. Der eine, ein pordses, dichtes Gestein, stammt von El] Puente,
dem Istmus zwischen der Padrebucht und Carbajal und ist ein Palagonittuff mit
einigen sporadischen Augit- und Magnetitk6rnern. Der andere Tuff stammt von
dem nordlichen Ufer der Padrebucht, von wo einige umgewandelte Olivinba-
salte (s. S. 23) herrihren. Ausser Augit und Magnetit enthalt derselbe einige
groéssere vollkommen reine Olivinkorner in einem teilweise kryptokristallinisch aus-
sehenden Glase. Stellt man diese verschiedenen Bildungen aus der Nahe der

b. Detail von Fig. 3 a.— Vergr. 170 x. Verf. phot.

Le «|
oq”
Oo

Padrebucht zusammen, so gelangt man zu der Auffassung, dass dieses Gebiet frische
Spuren vulkanischer Tatigkeit aufweist. Vergleicht man die obengenannten Tuffe
mit den von QUENSEL! beschriebenen roten Tuffen von der Cumberland Bay, so
scheinen die letztgenannten nicht so empfindlich gegen Verwitterung zu sein,
wie besonders die Palagonittuffe.

Hydrothermale Bildungen.

Ein aragonithaltiges Gestein vom Ufer gleich stidlich vom Yunquegipfel diirfte
als hydrothermal umgewandelter Basalt angesehen werden. (Fig. 6.) Das Hand-
stiick ist ein von weissen Streifen durchzogenes scharfgriines Gestein, das u. d. M.

1L.c. p. 266.

BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ-INSELN

grosse Augitkristalle in einer vollig zerflossenen Serpentinmasse zeigt. Das Pra-
parat ist von Aragonitbandern durchzogen. Dieses Gestein muss als ein stark
umgebildeter Olivinfels bezeichnet werden. QUENSEL! hat ganz frische Gesteine
von letztgenanntem Typus angetroffen und beschrieben. Vielleicht kann die Ara-
gonitbildung hier eine Andeutung geben, auf welche Weise der Olivin chemisch
umgewandelt worden ist.

Auf derselben Stelle wurde auch eine reine Kalksinterbildung gefunden, was
darauf hinweist, dass diese Gegend in spaterer Zeit postvulkanischen Prozessen
hydrothermaler Natur ausgesetzt gewesen ist.

In Fig. 7 ist eine eigentiimliche Bildung dargestellt, wie sie auf dem offenen

Fig. 4. Das Vaqueriatal. Wasserfall tiber den saulenférmig abgesonderten doleritischen Basalt.

Plateau bei Puente vorkommt. Es sind lange, in einem »Sandfeld» aufrechtste-
hende rohrahnliche Bildungen, Aragonit, Pyroxen, Magnetit sowie etwas Olivin
in einem Zement von Karbonat enthalten. Die wahrscheinlichste Deutung dieser
Phanomene ist wohl, dass mit Calciumkarbonat gesattigte thermale Gewasser tuber
eine Vegetationsdecke geflossen sind, wobei Wurzeln etc. mit einer Kruste von
oben angegebener Zusammensetzung uberzogen wurden.

Santa Clara.

An dem Siidende von Masatierra liegt die kleine Insel Santa Clara. Von
Santa Clara selbst ist keine Probe mitgebracht, dagegen von der kleinen Insel
Morro de los alelies, die bei tiefstem Wasserstand mit der Hauptinsel zusammen-

a Ex ps 205-

28 TOR H. HAGERMAN

hangt. Sowohl in Diinnschliff wie in Handstiick
zeigt dieses Gestein, das als fast vertikale Gange
auftritt, eine vollige Ubereinstimmung mit einem
der dichten Basalte von Masatierra vom Gipfel des
Cerro Negro SO von Yunque, 190 m i. d. M. Ur-
sprungliche Plagioklaseinsprenglinge von bis zu 2
mm Lange sind oft so stark kaolinisiert, dass beim
Schleifen nur die Hohlraume iibrig geblieben sind
und dem Gestein ein falsches, schlackiges Aus-
sehen verleihen. Hierbei ist interessant, dass der
Feldspat der Grundmasse sich frisch beibehalten
hat. Die Olivine sind natirlich vollig in Serpentin
umgewandelt. Das ganze Praparat ist von Ilme-
nitskeletten durchwachsen.

PU ais)

q Masafuera.
Fig. 5. Handstiuck vom Vaqueria-

gang. Verf. phot. Etwa 180 km westlich von Masatierra erhebt

sich die Insel Masafuera. Abweichend von Masa-

tierra in Bezug auf die einheitliche Mineralzusammensetzung der Gesteine liefert

Masafuera Beispiele petrographisch weit verschiedener Typen. Basalte mit den

dazugeh6renden Gangformen von ungefahr gleichem Mineralbestand wie die auf

Masatierra vorkommenden gibt es zwar auch hier, ausserdem finden sich aber

auch an Erzmineralien stark iibersattigte Basalte, sowie den Trachytandesiten sich

nahernde Gesteine. Am interessantesten ist jedoch das Vorkommen von reinen
Alkaligesteinen, wie z. B. die von QUENSEL angefihrten Natrontrachyte.

Natrontrachyt.

Leider ist das einzige mitgebrachte Handstiick dieses Gesteins von einem
losen Block am Fuss der Steilwand von Tierras Blancas abgeschlagen. SKOTTS-
BERG hat indessen miindlich berichtet, dass zahlreiche Blocke desselben Gesteines
in den Talusbildungen von Tierras Blancas vorkommen, und dass, soweit er ver-
stehen konnte, dasselbe hellgraue Gestein den ganzen oberen Teil der Steilwand
bildet; seiner Kartenskizze nach zu urteilen tritt dasselbe bereits 400 m u. d. M.
auf. Dies ist von Bedeutung fiir das Feststellen der Eruptionsfolge, die spater kurz
erwahnt werden soll.

Das Gestein besteht aus gleichmassigen K6rnern und ist sehr reich an Feld-
spat. Die Feldspatleisten erreichen eine Lange von 0,3—0,4 mm. Sie sind gut
parallelorientiert und verleihen dem Gestein eine trachytoidale Struktur. Da Albit-
zwillinge nicht vorhanden sind und der Feldspat durch die Anlagerung der dinnen
Individuen unscharfe Begrenzungen zeigt, konnte eine genaue Bestimmung des-
selben nicht ausgefiihrt werden. Die Lichtbrechung halt sich im allgemeinen etwas
ber Kollolith (n= 1,535), stellenweise ist das Relief jedoch ganz verschwunden.
Um eine nahere Kenntnis von den Feldspaten zu bekommen, ist eine Alkalibestim-
mung des Gesteins ausgefthrt worden. Diese ergab 3,45 % KO und 7,34 % Na,O.
Dies wiirde einem Gehalt von 20,44 % Ortoklas und 62,27 % Albit im Gestein ent-
sprechen. Auf Grund der Lichtbrechungsverhiltnisse diirfte jedenfalls neben einem
Kali-Natronfeldspat auch ein saurer Plagioklas der Oligoklasreihe vorhanden sein.

BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ-INSELN 29

Eine geringere Menge Pyroxen tritt ebenfalls auf. Die durchschnittliche
Ausdehnung desselben ist 0,1 mm, die Farbe ist gelbbraun, die kristallographische
Ausbildung schlecht entwickelt. Eine Ausloschung von c:x= 40 (ungef.) deutet
auf Augit. Schliesslich war auch Magnetit vorhanden, der Ofters fliessende Be-
grenzung der graubraunen Glasbasis gegentiber zeigt.

< Aragonitband, von
einem Augitindivi-
duum uberquert

Fig. 6. Hydrothermal umgewandelter Olivinfels von dem Yunque.
— Vergr. 12,5. Verf. phot.

Fig. 7. Sinterbildung von Puente. Massstab in Cm, Vert, phot.

Der von QUENSEL! beschriebene Natrontrachyt ist ungefahr 1200 m u. d. M.
gefunden worden und zeigt bei einem Vergleich einige Abweichungen. Das Hand-
stiick scheint im Gegensatz zu dem obenbeschriebenen etwas verkieselt zu sein.
U. d. M. bemerkt man sofort, dass der Pyroxen hier abweichend von dem vorer-
wahnten farblos ist. QUENSEL bezeichnet denselben als Diopsid. Ferner ist die
farbige Glasbasis nicht vorhanden.

208 Gop) "28 3;

30 TOR H. HAGERMAN

Andesit.

Von dem Berggipfel »Las Torres», 1370 m i. d. M., und in losen Blocken
von dem Bergriicken NO von der genannten Stelle 1200 m u. d. M. sind Hand-
stiicke von einem feldspatreichen aschgrauen Gestein mitgebracht worden. Obwohl
etwas olivinreicher, stehen dieselben den von QUENSEL! beschriebenen, einem
1100 m i. d. M. liegenden Niveau entnommenen Trachytandesiten sehr nahe. Als
Einsprenglinge kommen Feldspat und Olivin vor, u. d..M. zeigt der erstgenannte
eine Zusammensetzung von AbsgAngs und erreicht eine Korngrosse von 3—4 mm.
Die Olivine sind etwas kleiner. Sie werden im allgemeinen nur ca. I mm in
Diameter und sind wenig verandert, nur an den Randern zeigt sich eine schwach
gelbe Farbe, wo die Umwandlung begonnen hat. In der Grundmasse dominiert
der Plagioklas mit einer Ausdehnung von ca. 0,1 mm. Die Pyroxene und Magnetite
sind noch kleiner, im allgemeinen nur 0,o5 mm. Die Magnetite sind vollig idio-
morph. Die Klassifizierung hiehergehorender Gesteine ist etwas unsicher. Die
trachytoidale Struktur kénnte auf einen gewissen, nicht wahrnehmbaren Alkali-
gehalt in der Grundmasse deuten. Geniigende Griinde, sie als Trachytandesite zu
bezeichnen, liegen jedoch nicht vor.

Sehr interessant ist ein bei Correspondencia (1420 m i. d. M.) genommenes
Gestein. Mikroskopisch zeigt dieses Handstuck ein aschgraues, porphyrisches
Aussehen, doch sind einige Partien bedeutend dunkler. Als Einsprenglinge kom-
men Feldspat und Olivin vor. U. d. M. zeigt der erstgenannte eine Zusammen-
setzung Ab,An;, und erreicht eine Korngrésse von durchschnittlich 2 mm _ bei
einer max.-Lange von 5 mm.

Der Olivin ist vollig frisch, die Einsprenglinge treten aber in zwei verschie-
denen Entwicklungen auf. Dies steht im Zusammenhang damit, dass das Gestein,
wie bereits erwahnt, nicht vollig homogen ist. Die dunkleren Partien erweisen
sich bei mikroskopischer Untersuchung als bedeutend magnetit- und ilmenitreicher
als der ubrige Teil des Gesteins.

In den dunkleren Schlieren finden sich nun Olivine mit Magnetiteinschliissen
vollgesteckt. Besonders an den Randern ist der Magnetit so reichlich vorhanden,
dass die Olivinkorner véllig opak sind. Sowohl aus diesem Grunde als auch in-
folge der abgerundeten Form der Mineralk6rner scheint es, als ob diese Olivin-
kérner einer kraftigen Resorption ausgesetzt gewesen waren. Naheliegend ist nun,
dass diese dunkleren Schlieren mit ihrem grésseren Eisengehalt Riickstande auf-
geloster Bruchstiicke sind, welche urspriinglich zu dem Typus gehorten, die einem
schlackigen Basalt vom Gipfel des Inocentas entsprechen, der spater beschrieben
werden soll.

In einer Entfernung von kaum 1!/, mm von einem der erwahnten Olivin-
korner treten Individuen des anderen Typus auf. Diese sind ganz einschlussfrei,
véllig idiomorph mit scharfen Begrenzungsflachen, erreichen einen Durchschnitt bis
zu 4 mm und entsprechen vermutlich der intratellurischen Olivingeneration des
Hauptgesteins.

Der Feldspat in der Grundmasse der helleren Schlieren tritt in Stengeln von
ca. 0,14 mm Lange auf und verleiht durch seine Parallelorientierung dem Gestein
eine Fluidalstruktur. Die Zusammensetzung desselben ist Ab,gAn;,. Im tbrigen
enthalt die Grundmasse Magnetit und Pyroxen.

Die Grundmasse der dunkleren Partien unterscheidet sich von der obenge-

a nCs pe 252.

BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ-INSELN 31

nannten durch ihren Gehalt an Ilmenit, leicht erkennbar an seinem tafelformigen
Habitus. Ausserdem kommen hier feine Nadeln vor, welche aus einem ziemlich
stark lichtbrechenden Mineral bestehen. Dasselbe ist pleochroitisch von braun-
gelber bis gelbgriiner Farbe und weist parallele Ausléschung auf. Wegen der
kleinen Dimensionen der Korner konnte eine sichere Bestimmung derselben nicht
ausgefiihrt werden. Mit grosster Wahrscheinlichkeit liegt hier nur eine feinblattrige
Ausbildung von I|menit vor.

Basaltische Laven.

Auf dem Uferplateau an der Ostseite der Insel bei dem Casastal steht ein
feinkérniger Basalt mit porphyrischen Feldspat- und Olivineinsprenglingen an.
Die Feldspate erreichen eine Lange von 0,9 mm und erweisen sich als Plagio-
klase mit einer Zusammensetzung von Ab,jyAngy und stimmen also mit dem Feldspat
der Masatierra-Basalten iiberein. Die von QUENSEL! erwahnten, stark basischen
Feldspatkerne habe ich nicht angetroffen. Die Olivinkorner erreichen in diesem
Praparat eine Grésse von 0,3—0,4 mm und sind etwas iddingsitumgewandelt. Von
den Mineralien in der Grundmasse werden die Plagioklase am gréssten, 0,08 mm.
Der Pyroxen ist dagegen so klein, dass eine nahere Bestimmung sich nicht aus-
filhren liess. Er erscheint in kleinen, viereckigen, farblosen Kornern, meistens
zusammen mit dem Magnetit.

Diesem Gestein sehr nahe verwandt ist dasjenige, welches am Ufer des Mono-
Tales ansteht. Makroskopisch sind die beiden Gesteine einander sehr ahnlich.
U. d. M. tritt jedoch ein Unterschied auf, und zwar indem die Grundmasse des
Monobasaltes hier bedeutend mehr Olivin enthalt, weshalb man dieses Gestein
auch wegen der zahlreicheren Olivineinsprenglinge als einen Olivinbasalt bezeichnen
muss, wahrend sich das erstgenannte den Feldspatbasalten nahert.

In Quebrada del Ovalo, ungefahr 150 m ii. d. M., steht eine saulenformige
Basaltkuppe an, die dem Tal seinen Namen gegeben hat. Sie ist oben horizon-
talzerkliiftet, wahrend weiter unten eine prismatische Vertikalzerkliftung ansetzt.
Vorausgesetzt, dass die Kuppe aus ein und demselben Gesteine besteht, kann
dies als ein gutes Beispiel der von IDDINGS? dargetanen. Veranderung in der
Richtung fiir den kleinsten Schrumpfwiderstand innerhalb des erstarrten Gesteins
angesehen werden.

Die Handstiicke sind den unteren Teilen der 20—30 m hohen Saule ent-
nommen und bestehen aus einem porphyrischen Olivinbasalt. Die Einsprenglinge
sind Olivine von ca. 0,6 mm Durchmesser. Dieselben sind an den Randern dunkel-
rot, kaum durchleuchtend, was wahrscheinlich durch einen Gehalt an freiem Fe,O,
verursacht wird. Untenstehende Photographie (Fig. g) stellt einen dieser tibrigens
sehr sparlich vorkommenden Einsprenglinge dar.

Die Grundmasse ist der des vorstehend erwahnten Olivinbasalts vollig gleich,
nur etwas gréber. Der Plagioklas, ein Labrador, erreicht eine Lange von ca.
0,12 mm, das ganze Praparat ist parallelorientiert. Besonders hervorgehoben sei,
dass das Olivin in der Grundmasse nicht rotpigmentiert ist. Im tbrigen findet
sich Magnetit und der farblose Pyroxen.

IPS.) Pp. 270-
2 J. P. IppinGs, The columnar structure in the igneous rocks of Orange Mountain, N. J.
Amer. Journ. 31 (1886), p. 321.

32 TOR H. HAGERMAN

Der héchste Berg auf Masafuera ist der Inocentes. Von dem Gipfel, ca. 1500
m ii. d. M., wurde ein rétlicher, sehr schlackiger porphyrischer Basalt mitgebracht,
der sich besonders durch seinen hohen Gehalt an Erzmineralien auszeichnet.

Unter dem Mikroskop erwies sich die Grundmasse als hyalopilitisch. Die
Einsprenglinge bestehen hauptsachlich aus grossen, im ersten Augenblick voll-
standig opaken Kornern von ca. 3 mm Durchmesser. Bei genauerer Untersuchung
erweisen sich jedoch mehrere als stellenweise durchsichtig. Diese Teile besitzen

Fig. 8 a. Das heterogene Gestein von Correspondencia. Dunklere Partien oben und unten
sichtbar. — Vergr. tox. Photo. E. DAHLSTROM.

die hohen Interferenzfarben des Olivins, dazu weisen die Korner auch im ubrigen
den Habitus des Olivins auf. Bei Beobachtung in konvergentem polarisiertem
Licht bei einem Achsenaustritt wurde keine Kriimmung des Achsenbalkens beob-
achtet, weshalb der Achsenwinkel nahezu 90° sein diirfte. Dies deutet auf eine
fiir Basalte normale Zusammensetzung von Olivin mit verhaltnismassig niedrigem
Eisengehalt.

Kand. S. LANDERGREN fihrte eine Eisen- und Titanbestimmung der Ge-
steinsarten aus, wobei folgende Werte erhalten wurden: 18,82 % Fe,O3 und 3,59 %
TiO,. Dies entspricht einem Gehalt von 6,81 % Ilmenit und 15,44 % Magnetit.
Diese Zahlen sind natiirlich zu hoch, da Fe auch in den geringen Mengen Olivin,
Pyroxen und Glasbasis enthalten ist, die sich in dem Gestein befinden. Es ist

BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ-INSELN 33
jedoch besonders interessant, dass, obgleich das Magma eisenibersattigt war, das
Eisen die Konstitution des Olivins nicht nennenswert beeinflusste.

Von solchen magnetitiibersattigten Olivinen spricht Doss! in einer Beschrei-
bung von Basalten aus Syrien: »In diesen Gesteinen beherbergt der porphyrische
Olivin eine derartige Menge von Magnetitkérnern, dass dieselben meist die Halfte,
zuweilen ungefahr 4/; des ganzen Kristalldurchschnittes einzunehmen scheinen». An
einer spateren Stelle schreibt er: »Das Extrem hiervon tritt dann ein, wenn der

Fig. 8 b. Detail von Fig. 8 a (bei —).— Vergr. 210x. Zu beachten ist die parallele
Anordnung des Magnetits. Photo. E. DAHLSTROM.

Olivinkrystall einen breiten, vollig opaken, schwarzen Saum von Magneteismen
besitzt». Das Phianomen stimmt mit dem vorliegenden vollkommen wberein, nur
dass hier der entgegengesetzte Fall vorliegt, indem das ganze Korn vollstandig
opak sein kann, auch in sehr diinnen Praparaten.

REITER? hat sich mit der Zusammenschmelzung von Olivin und Magnetit
beschaftigt und sagt: »Bei Abkihlung tritt eine gewisse Ubersattigung ein. Vom
Magnetit scheidet sich ein Teil ab, dann wird der ubersattigte Olivin ausge-
schieden ...

1 B. Doss, Die basaltischen Laven und Tuffe der Provins Hauran und vom Diret et-Tulil
in Syrien. T.M. P.M. Bd. 7 (1886), p. 483—484.
aoe Gps 232:

34 TOR H. HAGERMAN

Wir erhalten hiemit also Magnetite in zonarer Anordnung in den nachher
entstehenden Olivinen eingeschlossen;.. .»

Die umstehende Photographie (Fig. 10) zeigt, wie der Magnetit parallel an-
geordnet ist, sodass die durchsichtigen Partien in Streifen auftreten.

Unter den Einsprenglingen kommt, wenn auch selten, hier und da ein Feld-
spatindividuum vor, dessen Zusammensetzung ungefahr Ab,yAngg ist; es handelt
sich also um einen Labrador, unbedeutend abweichend von dem Feldspate der
ubrigen Gesteinsarten.

Die basischen Plagioklase der Grundmasse zeigten zum Teil unscharfe Be-
grenzungen, durch die Anlagerung der diinneren Individuen verursacht.

Die Pyroxene sind bedeutend kleiner und erreichen einen Durchschnitt von
ca. 0,o2—0,o3 mm, wenn auch einzelne stengelige Individuen etwas grosser werden

Fig. 9. Rotpigmentierte Olivineinsprenglinge. Basalt von El Ovalo. — Vergr. 56.
Photo. Hj. OLSSON.

kénnen. Von diesen kommen zwei verschiedene Ausbildungen vor. Der eine (nor-
male) ist mit den in den beschriebenen basaltischen Gesteinen auftretenden Py-
roxen indentisch und vermutlich als Diopsid zu betrachten, da er beinahe farblos
und kaum pleochroitisch bei einer Ausléschung von ca. 45. ist. Der andere Typus
ist braungelb und tritt besonders um die Locher herum, niemals zusammen mit
dem erstgenannten Typus auf, stellt aber wahrscheinlich nur eine Pigmentierung
desselben dar. Auch hierin zeigt das Gestein eine grosse Analogie mit den vor-
erwahnten, von Doss beschriebenen Basalten. Er sagt auf Seite 481: »Hier be-
sitzen die in der Nahe der von Kalkspat ausgefullten Hohlraume gelegenen Augite
eine goldgelbe Farbe;...>».

Im iibrigen besteht die Grundmasse aus kleinen, idiomorphen Magnetitkornern,
Ilmenit und Glasbasis. Der ganze Gebirgskamm soll aus diesem ausgesprochen
basischen Gestein bestehen.

Bereits mikroskopisch zeigt ein anderes Handstiick, das vom Strandabhange
nahe des Casatales abgeschlagen ist, eine auffallende Abnlichkeit mit dem Ino-

BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ-INSELN 35
centesgestein. (Laut Angabe von SKOTTSBERG soll das ganze Ostliche Ufer aus
diesem Gesteinstypus bestehen, abwechselnd mit den obenerwahnten Feldspat-
basalten.)

Auch hier treten magnetitfiihrende Olivine auf, wenn auch sparlicher. Die
Feldspate sind dieselben und besteht der einzige Unterschied darin, dass das
Gestein glasreicher ist, weshalb der in dem letztgenannten Gestein vorkommende
Pyroxen nicht mit Bestimmtheit hat wahrgenommen werden konnen. Nimmt man
an, dass dieses Gestein einer rascheren Abkihlung ausgesetzt gewesen ist als
das vorgenannte, kann man dasselbe diesem gleichstellen und mdglicherweise
eine Andeutung iiber eine nicht unbedeutende Ausbreitung der fraglichen stark
basischen Gesteinsart finden.

Fig. 10. Magnetitiibersattigte Olivinsprenglinge im Basalt von Inocentes. Verf. phot.

Umstehende Tabelle ist eine Zusammenstellung uber die Veranderlichkeit
der effusiven Masafuera-Gesteine im Verhaltnis zum Niveau der Fundstatten. Die
Gesteine sind nach abnehmender Basisitat geordnet und dabei das Material SKOTTS-
BERG’s wie auch dasjenige QUENSEL’s beriicksichtigt.

Zwar fallen die Lavabetten etwas nach NNO ab, doch dirfte mit Riicksicht
auf den geringen Umfang der Insel die umstehende Tabelle ein gutes Bild der
Eruptionsfolge gewahren, wobei die altesten Gesteine beim Meeresniveau, also
links beginnen.

Vor allem verstdésst der stark basische Basalt aus der Hohe von 1500 m U.
d. M. gegen die unter Zugrundlegung der Eruptionsfolge vorgebrachte Theorie,
dass die Gesteine hier gravitativ differentiert sein sollen. Die Gesetzmassigkeit,
die QUENSEL bei diesen Gesteinen gefunden zu haben glaubte — mit den sauren
und alkalireichen als den jiingsten — scheint mir schwerlich mit diesen Beob-
achtungen in Ubereinstimmung gebracht werden zu k6nnen.

36 TOR H. HAGERMAN

| |
Natrontrachyte ... | O | | x | |
| |
5 | | | |
WWAMNGESILGis = osts-sp. ers | x O fon ire)
| | |
| |
Basaniteerst.., tei | fe
| Feldspat- | |
Basalte siti ae va ”© ; r | | | |
[owe Olvin, [oolo |_| _| ae
| Erziubersattigte Ba- | |
lie) Seer ares | Chisel | | fo)
! | | | | |
° I 2 3 4 500 6 7 8 9 1000 =II 12 13 14 1500
m ii. d. M.

Die Verteilung der Effusivgesteine der Insel Masafuera.
© bezeichnet Das Material SKOTTSBERG’s.
x » » > QUENSEL’s.

Es ist selbstverstandlich denkbar, dass die in den tieferen Teilen des Zufluss-
kanales versunkenen schweren Magmas zuletzt herausgestossen wurden, doch spricht
gegen diese Annahme das bereits in einer Hohe von 400 m i. d. M. auftretende
alkalische Gestein. .

Schliesslich ist aus der Tabelle zu ersehen, dass aus einer Ho6hbe von 500—
1000 m tberhaupt kein Material untersucht ist. Die Masafuera Gesteine sind
jedenfalls in Hinblick auf den Mineralbestand so verschiedenartig, dass eine nahere
Untersuchung der noch unbekannten Hohenlagen wichtige Aufschliisse ergeben
diirfte.

First printed February 25th, 1024.
Reprinted without change April 13th, 1054.

Additional Comments on the Geology of the
Juan Fernandez Islands.

B y

PERCY QUENSEL.

Contents.

Page
Introduction .. . ene e Mee Ses ge Pe ot Toot Ton ok sree ee ee GT
Main Geological menares ars ete GOR S Gia ty Se Pe ee eee tree
Petrology of the Volcanic Roeeeiane Pas tees ere ere ATT Sk Pk
MEE EL ee or) oe cyst ee MOO ase fy ce tees. Sey AE et ee es coe ae
ACE rm eg Mg 52 oy Pus es Bos! USL A os Te ee

Regional Relations.
Tectonic Connections 74
Petrographic Connections 17
General Conclusions Be oh en ete eae Gaim, Nee ee cee oe eee ee
MME SEIMeNtS: 050i os Gets hc elit Tee ee ee eee
MEME TEALYSOS es Loic) Fl hed | ayy se ay ens eae (oe | Goins Hele ae ERE ee eee
DEEPA CAPN ye A ce wi ee 8d to A. Ee A A cee ec

Introduction.

The Juan Fernandez Islands consist of Masatierra, Masafuera and Santa
Clara. In many publications the name Juan Fernandez has been used to denote
Masatierra only, the other islands then being indicated by the names as above.

The largest island, Masatierra, situated 660 km from and nearly due west
of Valparaiso, measures 95 square km. Masafuera, 170 km further westward, is
64 square km. The small island Santa Clara, close to Masatierra, is only
square km in area.

During more or less casual visits to Juan Fernandez stray observations have
been recorded on geological features of the islands. In all cases they refer to
Masatierra. It may be of interest in this connection to give a summary recount
thereof.

The first samples of volcanic rocks of Masatierra were, as far as known,
collected by Lord COCHCRANE in January 1823. He was returning to Europe
after 5 years service as admiral in Chilean service during the war of independ-
ence. The ship at his disposal, ‘Colonel Allen’, touched at Masatierra for two
days. Mrs MARIA GRAHAM, a passenger on board, has in her diary given the
following details, recorded by THOMAS SUTCLIFFE in his book Crusoniana: “Lord

3 — 516795 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I.

38 PERCY QUENSEL

Cochcrane brought from the summit (1.500 feet) a piece of black porous lava;
and under it he found some dark hardened clay full of cells, the inside of which
appear slightly vitrified. The island seems chiefly composed of this porous lava;
the strata of which, being crossed at right angles by a very compact black lava,
dip on the eastern side of the island about 22° and on the west side 16°, pointing
to the centre of the island as an apex” (1, p. 198).

In 1830 C. BERTERO published some observations under the title “Notice
sur l'Histoire naturelle de l’ile Juan Fernandez’. With regard to geological ques-
tions he says: “Je pense qu'un géologue n’y trouverait que du basalte dans les
états, méme dans celui de la plus parfaite décomposition; plusieurs blocs sont
parsemeés d'une cristallisation particuliere, a laquelle on donne, je croix, le nom
dolivine ... Il n’y a pas de trace de volcan; les pierres qu'on prend pour
de la lave, et dont quelques-unes ressemblent assez aux scories ou de la pierre
ponce, ne sont, a mon avis, que du basalte décomposeé; on trouve aussi cette
roche sous forme spherique, et composée de couches concentriquesmmsa
p. 345; compare in the latter respect Fig. 10 on page 52 of this paper).

A. CALDCLEUGH, who accompanied Captain P. PARKER KING on the sur-
veying voyages of H. M.S. Adventure and Beagle on their first expedition
1826—1830, read before the Geological Society of London on Jan. 5th, 1831, a
statement on “The geology of the island of Juan Fernandez’. In the Proceedings
of that year the following account of Caldcleugh’s discourse is given: “‘The
author could discover no trace of a volcano, said to exist here by former visitors;
all the rocks, according to him, consist of basaltic greenstone and trap of various
mineralogical structure, both amorphous and vesicular, together with trappean
concretions, no other contained minerals being observable except olivine and
chaux carbonatée métastatique. It is further mentioned that the basalt in parts
is almost columnar, and in others has a peaked and serrated outline, the mass
being, here and there, traversed by dykes. Owing to the peculiar character of
this basalt, and especially from the great quantity of olivine, the author compares
its age with that of the basalt of Bohemia, the Rhine, the Vivarrais and Beaulieu
in Provence” (3, p. 256, also published in the Phil. Mag. and Annals of Philosophy,
Vole rs3i. sp.5220):

Captain King recapitulates Caldcleugh’s narrative, as given above, with the
addition: “In captain HALUL’s interesting journal, there is a list of geological and
mineralogical specimens, of which one from Masafuera is named vesicular lava’
(4, p. 304). The ultimate destiny of these specimens is unknown.

Members of the Dumont d’Urville expedition, when visiting Masatierra in
1838, collected and specified several different samples of the lavas from the island
(5 jor 114). The material for the new analyses of basalts from Masatierra, which
LACROIX recently caused to be made and which will be referred to later on, are
evidently from this collection, as Lacroix says they were made from specimens
collected by the Dumont d’Urville expedition. G. GRANGE records some observa-
tions as follows: “Toutes les roches appartiennent a diverses variétés de trapp
et de diorite basaltique amorphes et vésiculaires, on ne trouve dans ces roches
volcaniques aucun autres minéraux que de lolivine et de la chaux metastatique.

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 39

La roche basaltique s'y présente le plus souvent en couches superposées, quel-
quefois en escarpements interrompus et fractionnés et sont traversés par des dikes
d’éjections plus modernes. Le basalt forme des pics élevés dans l’ile et sur quelques
points prend une disposition prismatique fort remarquable’’ (6, p. 39).

In 1896 L. H. PLATE published a paper ‘Zur Kenntnis der Insel Juan Fernan-
dez’. Plate was a zoologist and the paper deals with the zoology of Masatierra,
but some introductory remarks refer to geological observations from which the
following instances may be quoted: “Das Gestein der Insel ist ausschliesslich
vulkanischer Natur und besteht aus schwarzer, basaltischer Lava, der an einzelnen
Stellen weissliche oder rothliche Tuffe eingelagert sind ... Die Insel fallt fast iber-
all mit senkrechten Wanden, deren Hohe zwischen 100 und 300 m schwankt, gegen
das Meer zu ab und nur in den Hafen der Ansiedelung (Bahia Cumberland), dem
Puerto Ingles, dem Puerto Frances und der Bahia de la Vaqueria erstrecken sich
die Thaler bis an die Kite, so dass man in diesen Stellen ohne Mihe vom
Meer in das Innere der Insel vordringen kann... Die Lavawande sind deutlich
geschichtet ... Betrachtet man nun vom Meer aus eine (solche) Lavawand, so
erkennt man sofort dass sie geschichtet ist; denn sie wird in ganzer Ausdehnung
von zahlreichen horizontalen Linien durchsetzt ... Diese horizontalen Linien sind
wohl der Ausdruck des successiven Aufbaues der Insel. Aus einem submarinen
Krater ergossen sich Lavastrome und breiteten sich auf dem Grund der Oceans
aus. Die Eruptionen wiederholten sich haufig, und so floss eine Lavaschicht uber
die andere, um spater zum Teil tiber die Oberflache des Meeres gehoben zu
werden... Ohne Zweifel war die Insel in friiheren Erdperioden sehr viel grosser.
Eine Untersuchung des Meeresbodens zwischen Masatierra und Masafuera wird viel-
leicht spater den Beweis bringen, dass die beiden Inseln, welche jetzt 92 Seemeilen
von einander liegen, urspriinglich nur eine einzige bildeten oder doch wenigstens
die héchsten Punkte desselben submarinen Plateaus darstellen und daher gleich-
zeitig entstanden sein miissen” (7, p. 221).

The observations, hitherto recorded, have hardly more than historical interest.
As little attention had been paid to the geology of the Juan Fernandez Islands
before the last decade of the past century, these casual comments have, however,
here been included to evade oblivion.

The first observations of any importance regarding the rocks of Masatierra
are based on specimens, collected by members of the Challenger Expedition in
1876. A. RENARD has given a description thereof in a paper titled “Rocks of
the Island of Juan Fernandez’. (This name here signifies Masatierra.) He says
that “the rocks which have been submitted to examination all belong to the
basalt type, and it seems probable that the whole island is made up of those
rocks that we are about to describe. The rocks, which form the central mass of
the island, appear in the specimens as dolerites or as common basalts ... Among
the specimens collected on the coast of Juan Fernandez it is necessary to mention
a greyish very scoriaceous rock from which stand out large crystals of plagioclase
of a waxy and milky appearance. This rock is a dolerite with large vesicules. Under

40 PERCY QUENSEL

the microscope the fundamental mass, in which the plagioclase crystals are embedded,
has a dolerite structure. The felspar crystals show large extinction angles (38°—41 )
which may be compared with those of bytownite. The sections of the mineral
are cracked and pervaded with zeolite matter, which forms an irregular network.
This matter which looks slightly grey, when seen by ordinary light, remains
obscured between crossed nicols... The olivine, of which large sections are seen,
is uniformly changed into a red hematite; these sections, however, still show
extinctions like those of the unaltered olivine’ (8, p. 176). The characteristic
change of olivine to iddingsite in the basalts, which will be treated below in
some detail, is here evidently noted by Renard, though by him named hematite.
For the rest his description mostly refers to the colour and texture of differ-
ent samples of dolerite and basalt.

In 1886 L. DARAPSKY published a short report on detached rounded lumps
of magnesite from Bahia del Padre in Masatierra, locally known as “‘piedras de
campana’”’ but named “‘Glockenstein’”’ by the author (9). R. POHLMANN later (1893)
described this singular formation in rather more detail and discussed its origin (10).

An account of some geological and petrographical observations by J]. SCHULZE
and R. POHLMANN, participants in FEDERICO JOHOW’s expedition to the islands
in 1891, has been published by the latter as an introduction to Johow’s monograph:
‘Estudios sobre la Flora de las islas de Juan Fernandez’ (11, p. 1).

As a member of the Swedish Magellanian Expedition of 1907—1909 I had
the opportunity, together with Professor SKOTTSBERG, of visiting the islands in
1908 and later described the rock specimens then collected in a paper: ‘Die
Geologie der Juan Fernandezinseln’ (12). In 1916—17 Skottsberg again visited the
islands and collected specimens from some new localities. These were subsequently
described by T. HAGERMAN under the title ‘Beitrage zur Geologie der Juan
Fernandez-Inseln’ (13).

Since Skottsberg’s visit in 1917, no further exploration in the field has been
published concerning the geology of the islands in question. And, with the exception
of four recent analyses of lavas from Masatierra from collections made by members
of Dumont d’Urville’s ‘Voyage au Pole Sud et dans l'Océanie’ in 1838, commented
on by LACROIX (14, p. 64), no further observations have, as far as known, been
published on the geology or petrography of the Juan Fernandez Islands. On the
other hand questions relating to their lithological connection with other intra-
pacific islands and their geophysical position in relation to eastern-pacific volcanic
centres has been the subject of repeated discussions during later years.

Main Geological Features.

The two islands Masatierra and Masafuera present very different aspects with
regard to their bulk configuration. Masatierra exhibits a rugged appearance of
isolated jagged cliffs (Fig. 2). The highest peak, El] Yunque, is 927 m. On the
clear-cut shore-bounded escarpments one can distinguish many hundred lava
beds overlying one another. In thickness they can vary from some few metres
up to 20m or more.

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 4I

Fig. 1. Perspective view of Masafuera from the East. ‘Reproduced from F. Johow, Estudios sobre
la flora de las Islas de Juan Fernandez. Santiago de Chile 1896.

Masafuera presents notable dissimilarities to Masatierra. Its dome-shaped,
sharpiy bounded outline with an approximately oval circumference and its impos-
ing height give an impression of a volcanic configuration deviating from that
of Masatierra. On the west side of the island the cliffs fall all but perpendicularly
for over 1000m. The shore line below is a stony and sandy reach, in part
named Loberia vieja on account of the number of fur seals found there at certain
seasons. The eastern coastline has a very different appearance. As the highest
ridge lies towards the west, the fall on this side is not so precipitous. Instead
it is traversed by numerous narrow and steep sub-parallel erosion valleys (que-
bradas). Attempts to ascend the higher parts of the island can only be made from
the eastern side by means of these quebradas, a climb which is anything but
easy going (Fig. 3). The highest peak is 1 500 m.

The sketch in Fig. 1 gives an excellent conception of the singular configura-
tion of Masafuera’s eastern escarpments.

The Juan Fernandez Islands are exclusively formed of volcanic material, in
many respects of much the same nature as in other volcanic islands of the
eastern Pacific. All indications tend to show that they are of late origin. Both
VON WOLF (15, 1929, p. 771) and J. BRUGGEN (16, p. 59) consider them to be
late tertiary or pleistocene in age. Lacroix says regarding all the non-coral-
line islands of the southern central Pacific: “leur age précis est indetermine,
mais il est certainement tertiaire, pléistocene ou meme, dans certains cas, récent”
(14, .p. 55).

No signs of recent activity have been found on the Juan Fernandez Islands.
That the immediate environs have recently been subjected to sub-marine volcanic
activity is, however, evident from a narrative, published in the Report of the
Challenger Expedition as follows: “In 1835 Masatierra appears to have been
governed by a Mr SUTCLIFFE, an Englishman in Chilean service. He was present
when the earth-quake took piace on the 20th of February of that year, of which
he gives the following account: At 11.30 a.m. the sea rose over the mole and
afterwards retired, leaving the greater part of Cumberland Bay dry, so much so
that old anchors on the bottom became clearly visible. The earth then began to
shake violently, and a tremendous explosion was heard, the sea still receding
in immense rollers, which afterwards returned, violently rising to such a height
that the settlement was literally covered and washed away, when the sea again
receded. The phenomenon occurred four times, causing much destruction, uproot-
ing trees and drowning cattle. Shortly after the explosion, a large column, some-

42 PERCY QUENSEL

Fig. 2. The south slope of the high ridge along the western half of Masatierra as seen from
Portozuelo. Photo C. Skottsberg.

what resembling a water-spout, was seen ascending from the sea off point Bacalao,
which proved to be smoke, but at 7 p.m. volcanic flames were visible through
the smoke, which lasted till 2 a.m. on the 21st. The depth of the water on the
spot, where the eruption took place, was from 50 to 80 fathoms; no alteration
in the depth was detected after the eruption had subsided” (17, p. 818). Sutcliffe
has published an account of the ‘earthquake’ in a separate publication (18) and
reproduced a sketch of the sub-marine eruption (19, p. 387).

It is obvious that these narratives must refer to a sub-marine volcanic explo-
sion. CHARLES DARWIN also mentions the phenomenon in his ‘Geological Observa-
tions on the Volcanic Islands’ (20, p. 149). Renard gives the position of the
explosion as 1 English mile from the island and remarks “‘that the close prox-
imity of a volcanic centre seems therefore to be implied” (8, p. 176).

The sub-marine eruption must be taken as conclusive evidence that the
immediate neighbourhood of the Juan Fernandez Islands has been the seat of
volcanic action within the last 115 years. A point of further interest is that the
explosion was simultaneous with violent earthquakes on the Chilean coast, as
Darwin already observed (see p. 75).

Briggen refers to some further observations of sub-marine eruptions in
the vicinity of the Juan Fernandez Islands, recorded by FR. GOLL in his paper
‘Die Erdbeben Chiles’ (Miinchener Geogr. Studien 1904, Nr. 14). The following
denotements by GOLL are taken from Briiggen (16, p. 332):

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 43

Bi itil.

Fig. 3. Quebrada del Varadero on the east coast of Masafuera. Photo C. Skottsberg.

44 PERCY QUENSEL

“Segun Goll se produjo, el 12 de febrero de 1839 una erupcion submarina
y maremoto a unas 120 km al este de la Isla de Mas a Tierra. El mismo autor
cita la observacion siguiente hecha en un punto un poco mas austral: “En octubre
de 1867, se sinti6 un temblor submarino en 34°55 Siy 77 38° W (umasmnae
millas SE de Juan Fernandez); después el buque navego durante dos horas por
agua de color blanco lechoso, habiendo mucho pescado muerto en la superficie.’
Se trata probablemente del mismo fendmeno que describe JOSE M. POMAR en
la forma siguiente, aunque dice que el punto se halla a 100 millas al SW en
vez de SE de Juan Fernandez: ‘En 1867, el capitan SIMPSON de la barca britanica
Coronella navegaba en el Pacifico con mucha calma y vientos contrarios, con
excepcion de un fuerte viento acompafiado de siete temblores que se produjeron
como a 100 millas al SW de la Isla de Juan Fernandez; durante dos horas navego
por agua tan blanca como leche; sondeé, pero no tocd fondo en 100 pies de
profundad, vid muchos pescados muertos y una gran cantidad de pajaros por
todas partes. Agregaba el capitan Simpson que si hubiera estado 10 millas mas
adelante, el choque hubiera sido peor y hubiera causado averias al buque.’”’

Concerning the earthquake of Vallenar in the province of Atacama on the
1oth of Nov. 1922 Briiggen cites BAILEY WILLIS’ observations on the contem-
poraneous volcanic activity on San Felix.

Briiggen concludes his opinion on the submarine explosions, given above,
as follows: ‘A la teoria del origin de los tsunamis por erupciones submarinas podria
objetarse que serian erupciones muy excepcionales, ya que consisterian en una sola
o muy pocas explosiones que causan las pocas olas sismicas, apagandose luego
la actividad. En realidad se tratara solamente de las primeras explosiones que
abren la chimenea para la salida de la lava o de los gases y que tienen la fuerza
suficiente para causar el tsunami. También en otro sentido, las erupciones sub-
marinas se distinguen de las de los volcanes de los Andes, que solo excepcional-
mente entran en actividad durante los terremotos. Los volcanos submarinos
parecen estar en relacién mas estrecha con los focos sismicos de los grandes
terremotos chilenos’ (16, p. 332).

Petrology of the Volcanic Formations,

The following description of the rocks of Masatierra and Masafuera is based
on specimens collected by myself in 1908 and by SKOTTSBERG in 1917. In many
cases the two collections supplement one another and help to elucidate to a
certain degree the distribution of the somewhat varying types of the lava flows.

The specimens from Masatierra in both collections give conclusive evidence
that this island in the main is formed of a rather uniform series of basaltic lava
beds, only diverging in respect of coarser or finer grain or of a higher or lower
content of olivine. The specimens from Masafuera on the other hand indicate
more obvious dissimilarities in the composition of the rocks at different levels.

In the following the principal petrological features of the two islands will
first be treated. Under a later heading references will be made to resemblances
in various respects to other volcanic islands of the eastern Pacific.

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 45

Masatierra.
Lat..-33° 374° S., Long... 78° 50° W,,

The two collections of specimens from Masatierra by Skottsberg and my-
self have partly been taken from different localities. The central parts of the
island are however sparsely represented. Somewhat more complete series of rock
samples originate from the heights around Bahia Cumberland (Ensenada de San
Juan Bautista), Puerto Frances and Portezuelo as well as from Bahia del Padre.
From the adjacent small island Santa Clara there is only one specimen. It is
not easy to single out the local distribution of the different lava beds on account
of the inconnected localities from which specimens have been collected. Some
characteristic features may however be found which indicate that certain types
of lavas are restricted to localised areas. To some extent one may then draw
conclusions regarding the sequence of the volcanic eruptions.

The predominating rocks of Masatierra are olivine basalts, differing only in
their content of olivine. Lava beds from around Puerto Frances are exceptionally
rich in this mineral, the content of which can reach 40 vol.% of the rock (Fig. 6). Such
rocks, with an extreme content of olivine, I named picrite basalts in my earlier
paper (12, p. 265). Lacroix originally named such basalts ‘picrite feldspatique’
but later discarded this name, substituting for it the name oceanite‘, under which
name he includes the basalts of the Juan Fernandez Islands with an exceptionally
high content of olivine (14, p. 65). Since the name picrite basalt, as originally
defined in my former paper, has later been adopted’, I will retain this name for
the rocks in question, with the name oceanite as synonym.

From the extremely olivine-rich basalts around Puerto Frances there is every
transition over intermediate types to feldspar basalts without any olivine at all
or with only a very insignificant amount thereof. Such rocks, however, have a
relatively limited distribution on Masatierra. In general one may say that types
with a very high content of olivine are restricted to lower elevations, whereas
higher up more normal olivine basalts predominate.

A second feature of dissimilarity in the basalts is found in their texture.
Many of the lava beds show a coarse-grained ophitic texture and may be classed
as dolerites, and have been so named by Hagerman and myself in our previous
papers. Such rocks are usual around Puerto Frances and Bahia Cumberland as
well as at Vaqueria and Tres Puntas. Specimens from all these localities are
in outward appearance very much alike. In general these rocks, in contrast to
other lavas, are singularly fresh. Only the olivine often shows a dark brown
rim, indicating an incipient alteration to iddingsite (cf. 12, p. 260).

The doleritic basalts seem only to occur at lower horizons, where the lava
beds generally attain their greatest thickness. All the specimens thereof at hand
are from between sea level and 200 m. A sample in Skottsberg’s collection from
Bahia Cumberland (Fig. 7) and another from Vaqueria are both from 150 m above sea

* The name océanite was first proposed by Lacroix in 1923 (in Minéralogie de Madagascar,
Vol. Ill, p.49). The name is given ‘‘A cause de leur abondance dans les iles du Pacifique”.
? Holmes, Q. J. Geol. Soc. 172, 1916, p. 231; Washington, Am. J. of Sc. V, 1923, p. 471.

UENSEL

PERCY

‘oL6 £O1 : I afeoG “1Z6I JO JAVYD “AO ULITIYDO oY) WIOIJ SUOLIITIOD ILA C6Z1 jo vAvWLY op Jopruly Opuvuoy Aq dey ‘vitonseq Psy

( PUATMIPSOND df /

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 47

Basaltic lava beds. Western head of Bahia Cumberland, Masatierra. Photo C. Skottsberg.

Fig.

utr

level. The two specimens are in texture and mineral composition identical. A
third sample from Tres Puntas is from 200 m. Specimens in my earlier collection
were taken from the lower lava beds in the quebradas around Puerto Frances.

As no scoriaceous or slaggy development is to be seen in the upper or lower
parts of the doleritic beds, which have a very coarse-grained texture, they may
well represent intrusions between previously consolidated lava flows, in accordance
with what DALY has assumed to be the case with similar doleritic rocks in Hawaii
{R. Daly, Differentiation in Hawaii. Journ. of Geology, Vol. 9, 1911, p. 291).

The doleritic basalts and the picrite basalts have much the same mineralogical
composition. The only essential difference is the higher content of olivine in the
picrite basalts. The other rock-forming constituents in both rocks are labradorite,
a pleochroic Ti-augite and magnetite.

In the picrite basalts (oceanites) from around Puerto Frances as well as in
the olivine basalts in general the phenocrysts of olivine are often more or less
altered to iddingsite. In some cases only insignificant rests of olivine are left;
the iddingsite pseudomorphs, however, still retain the crystal habit of the olivine.
To the petrogenetic problem of iddingsite as representing a deuteric mineral I
will return later in connection with equivalent alterations in certain lava beds of
Masafuera, where the ‘iddingsitisation’ has gone further and there gives the rocks
a very characteristic aspect (see p. 60).

In the picrite basalts from Puerto Frances inclusions of dunite occur. The
large olivine crystals of this rock are singularly fresh, without any signs of even

48 PERCY QUENSEL

oO <

Fig. 6. Picrite basalt (oceanite). x g. Puerto Frances, Masatierra.

Fig. 7. Doleritic basalt. x 37. Bahia Cumberland, Masatierra.

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 49

a periferic alteration to iddingsite which is otherwise usual in the olivine of the
surrounding basaltic lavas.

At higher levels the basalts have a more normal character. Large phenocrysts
of olivine, or olivine and feldspar, lie in a more or less glassy groundmass of
augite, olivine and labradorite with abundant small grains of magnetite. Ilmenite
in tabular form or in skeleton crystals is now often present. Most of these lavas
are vesicular, scoriaceous or slaggy. The cavities are in many cases rimmed or
filled with opal.

The occurrence of basalts of this type is widespread up to the highest parts
of the island, and they are without doubt the dominant lavas of Masatierra.

Somewhat divergent lava beds seem, however, to predominate at intermediate
horizons. All the specimens at hand of this type are holocrystalline rocks of an
ash-grey colour, aphanitic and aphyric in texture (Fig. 8). They generally show a
characteristic light zone of weathering and a tendency to develope a columnar struc-
ture (Fig. 9). The fine-grained mineral assemblage consists of augite, labradorite and
very abundant magnetite in small euhedral crystals, evenly dispersed throughout
the rock. Ilmenite is also generally present in skeleton crystals. Stray small
grains of olivine may in some cases be observed; in other specimens olivine is
completely absent. In vesicular lavas of this type the cavities are again more or
less filled with opal (Fig. 8 b).

These lava beds, which represent the only specific feldspar basalts of Masa-
tierra, are found at heights between 400 and 500 m (Cordon Chifladores 400 m,
Portezuelo 500m). At an elevation below 100 m at Punta Larga in the more
western part of the island the same type has been found, but then in the form of
a dike, which may signify a channel for the analogous lavas at higher levels.

The very characteristic aspect of these rocks seems to indicate that they
represent a definite epoch of intrusion, intermediate between the doleritic basalts
and picrite basalts of the lower parts and the scoriaceous olivine basalts of the
higher horizons. An analysis has been made of a very similar rock from Masafuera,
which confirms its classification there as a feldspar basalt (see p. 66).

Lacroix’s four new analyses of basaltic lavas from Masatierra indicate that
the analysed rocks are similar in composition. It is regrettable that the specimens
all originate from Bahia Cumberland. Probably the members of the d’Urville
expedition only brought back samples from that locality and these were there-
fore the only specimens available in Paris for Lacroix’s analyses. On the other
hand the insignificant dip of the lava flows on Masatierra (15—20°) may infer
that the analysed rocks can be taken as representative in chemical composition
for the basal basalts of the island. Microscopic determinations of corresponding
specimens from the other localities point in the same direction.

With the exception of the ultra-femic picrite basalts from Puerto Frances
and the inclusions therein of dunitic rocks, and the light grey feldspar basalts
of intermediate horizons, we may conclude that the dominant rocks of Masatierra
consist of rather normal olivine basalts deviating principally in texture and ina
varying content of olivine.

ae) PERCY QUENSEL

Fig. 8a. Vesicular aphyric feldspar basalt (columnar structure). » 30. Portezuelo, Masatierra
see Fig. 9).

Fig. 8 b. Same specimen as Fig. 8a, magnified x 65. Vesicle in upper right corner filled with
z 5 5 bp) : 5
opal. Portezuelo, Masatierra.

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS SI

te x
» % ; ete
A ¥ . eck
gi ? 2 “ ee ow
is s \, i eae oie ee Fas ee a ed i Sa Ri:

Fig. 9. Northern wall of the Portezuelo Pass, showing columnar structure of feldspar basalt
Fig. 8 a). Photo C. Skottsberg.

52 PERCY QUENSEL

Fig. 10. Spheroidal weathering of basalt. Bahia Cumberland, Masatierra. Photo C. Skottsberg.

Quantities of pyroclastic material are found interbedded between the lava
flows all over the island. Pohlmann has described such products from Bahia
del Padre as follows: ‘‘Uberall sieht man Decken von basaltischem, meist sdulen-
formig abgesondertem Eruptivgestein mit Schichten von vulkanischem Auswurfs-
material wie Asche, Lapilli, Bomben u.s. w. wechsellagernd.

Eine solche aus Lapilli und Bomben zusammengesetzte Schicht von mehreren
Metern Machtigkeit an der rechten Seite der Bucht in der Nahe der sog. Kapelle
gelegen, liefert das Material der sog. ‘Glockensteine’. Der Vorgang ihrer Bildung
ist, kurz gesagt, folgender: zunachst entstehen traubige und nierformige Con-
cretionen zwischen den losen zusammengefiigten vulkanischen Massen; diese weissen
Knollen (Fig. 12) gelangen beim Abstiirzen der Schichten an den Strand und erhalten
als Rollsteine durch die Thatigkeit des Wassers ihre gerundete Form” (10, p. 321).

I have previously noted that the lava beds of the eastern parts of the island
are also often interbedded with agglomeratic layers (12, p. 257).

Some stray occurrences of superficial tuffs seem still to have evaded destruction
by erosion. When Renard says that “amongst the specimens collected at Juan
Fernandez (Masatierra) by the Challenger Expedition in 1875, we have not found
any specimen which might belong to any recent eruption, no tuffs, no volcanic
ash are to be found and everything seems to prove that they have been washed
away by the waves and the atmospheric agencies” (8, p. 176), this last conclusion
seems questionable. Hagerman refers to two specimens in Skottsberg’s collections

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 53

as representing recent tuffacious material. The one he describes as “ein pordses,
dichtes Gestein von El Puente, dem Istmus zwischen der Padrebucht und Carbajal
und ist ein Palagonittuff mit einigen sporadischen Augit- und Magnetitkérnern.
Der andere Tuff stammt von dem nordlichen Ufer der Padrebucht, von wo einige
umgewandelte Olivinbasalte herriihren’” (13, p. 26). As these specimens have
been collected without any observations regarding their petrological position, one
cannot draw any conclusive evidence as to their age, but the specimens from both
the localities have every appearance of being recent pyroclastic sediments.

I have previously described rocks from Bahia Cumberland, filling out the
greater part of the bay, which I assumed to be of recent tuffaceous origin. The
description was given as follows: “In den zentralen Teilen von Cumberland Bay
liegen noch ziemlich machtige, meist lebhaft rot gefarbte Tufflager, die sehr stark
umgewandelt sind. Bruchstiicke von Olivinkristallen, lapilliartige Lavabruchstiicke,
Erzk6rner und Glas liegen in einer Grundmasse, die aus einer weichen, mit Messer
schneidbaren, roten lateritahnlichen Lehmsubstanz besteht, die durch Verwitterung
aus dem urspriinglichen Tuffmaterial hervorgegangen sein diirfte. Uberall in den
Tuffsedimenten ist eine deutliche Lagerung sichtbar’ (12, p. 266). It is over 40
years since I visited the locality and naturally I cannot now rely on any personal
recollection. The inundation in connection with the volcanic disturbance of 1835,
referred to above (p. 41), may have wrought such havoc, that superficial deposits
could have been re-formed. But the composition of the formation, as well as my
notes from the field, offer indications that the tuffaceous material of Bahia Cumber-
land also represents pyroclastic sediments of recent volcanic origin.

Even if only trivial remains of tuffaceous formations, indicative of late volcanic
activity on the island, are left, the submarine explosion of 1835 confirms without
doubt that the area in the immediate vicinity has at that time been subjected
to disturbances of volcanic nature.

Some rocks from around Bahia del Padre deserve special notice. Schulze
and Pohlmann have observed the deviating nature of the rock assemblage and
the latter has commented thereon as follows: ‘‘De suma importancia para esplicar
la formacion jeoldjica de Masatierra es la entrada a Bahia del Padre, situada en
la parte suroeste de la isla. Aqui se observa debajo de las capas basalticas ya
descritas un grand macizo de roca compacta verdosa, que es andesitica. Segun
la opinion de Schulze’ que, a me parecer, es correcta, esta roca verdosa es la
mas antigua de la isla. En ningun otro punto, ni en Masatierra, ni en Masafuera
se ha observado una configuracién jeoldéjica analoga a la mencionada”™ (rasp).

The rocks referred to by Péhlmann certainly show a divergent aspect. But
I do not think they can be taken as representing rocks belonging to a more
ancient formation than the lavas around, nor that they should be classified as
andesites, as Pohlmann has assumed. I have as the result of microscopical
examination of the rocks come to the conclusion that the deviating character is
the result of post-volcanic alterations through thermal processes. In my earlier
paper I have given in some detail the reasons for this conclusion (12, p. 266), and

* Dr. Schulze died before publishing his observations.

4 — 516795 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I.

54 PERCY QUENSEL

Fig. 11. Volcanic ‘1 merate. Bahia Cumberland, Masatierra. Photo C. Skottsberg.

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS

wal
On

Fig. 12. Lumps of magnesite (the so-called Glockenstein). Nat. size. Bahia del Padre, Masatierra.
to) c

Reproduced from P. Quensel, 12, p. 269.

Fig. 13. Scoriaceous lava with newtonite and bole. Nat. size. Behia del Padre, Masatierra.

Reproduced from P. Quensel, 12, p. 269.

will in this connection only summarize the facts. The
fine-grained and of a greenish colour. The abundant cle

with epidote and calcite. Pyrite is finely dispersed thru”:

the microscope one can see that original phenocrysts of

s in question are very
age plains are coated
out the rock. Under
th olivine and augite

are more or less completely changed to serpentine and c_ orite. The matrix con-

sists of olivine, augite, plagioclase, magnetite and ilmenite — 1e., the usual
mineral assemblage of the basaltic lavas. The greater part of the silicates are,

56 PERCY QUENSEL

however, very much altered to serpentine, chlorite and prehnite. Secondary albite
and some scapolite is also present together with abundant pyrite. Small cavities
are filled with epidote and calcite. In the slaggy lavas in the immediate vicinity
larger cavities are filled with a purely white clay-like substance which was found
to be newtonite. In intimate association with this mineral is nearly always found
a yellowish waxy substance which shows every resemblance to what mineralogi-
cally may be signified as bole (Fig. 13). It is very brittle with a conchoidal
fracture. In water it readily disintegrates into small angular fragments. This bole
mineral is also found filling cracks, or occurs in smaller masses in the cavities
of the basalts around the bay.

In the agglomeritic lavas of the vicinity large cavities are filled with hard
compact magnesite. As early as 1886 Darapsky described this mineral by the
name ‘“‘Glockenstein’ and gave an analysis thereof, which shows it to be an
exceptionally pure magnesite (9, p. 113). Without doubt the magnesite is primarily
derived from olivine, the decomposition being caused by the same processes as
have changed the basaltic lavas nearby.

The whole aspect of the rocks from Bahia del] Padre, with magnesite, calcite,
serpentine, chlorite, scapolite and pyrite as secondary minerals, seems without
doubt to indicate that the lava beds in quéstion have been subjected to alterations
in connection with thermal processes during some intermediate phase of volcanic
activity on the island. There seems no reason to classify them as andesites of
an older formation, as is done by Schulze and Poéhlmann.

In a specimen collected by Skottsberg from the shore south of El Yunque,
Hagerman also found evidence of a far-reaching decomposition. Under the heading
“Hydrothermale Bildungen” he gives the following description: “Das Handstuck
ist ein von weissen Streifen durchzogenes scharfgriines Gestein, das u. d. M.
grosse Augitkristalle in einer vollig zerflossenen Serpentinmasse zeigt. Das Pra-
parat ist von Aragonitbandern durchzogen. Dieses Gestein muss als ein stark
umgebildetes Olivinfels bezeichnet werden” (13, p. 27). The large olivine crystals of
the dunite, found as inclusions in the picrite basalt at Puerto Frances, show, on the
contrary, no signs of secondary alteration (see p. 47). There is therefore no doubt
that the highly decomposed “Olivinfels’ described by Hagerman has succumbed
to a later decomposition of much the same nature as has been active around
Bahia del Padre.

Masafuera.
Mate 33 526 S.. Lone.So. 541. WwW:

The lavas of Masafuera present a more varying aspect than those of Masa-
tierra and contain several types of petrographic interest. They have, however,
hitherto only been summarily described in the papers by myself (12, p. 274) and
by Hagerman (13, p. 28).

The rocks which predominate at lower levels are mostly vesicular to slaggy
basalts. They are well represented around the Quebrada de las Casas. At higher
levels the basalts consist of more compact lava beds. In contrast to the basalts

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 5

—I

ii, Pa Ps

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Fig. 14. Masafuera. From the Chilean Gov. chart of 1921. Scale 1 : 70000,

PERCY QUENSEL

ut
Co

Fig. 15. View of the interior of Quebrada de la Loberia vieja towards the shore.
Photo C. Skottsberg.

of Masatierra, olivine is more scarce in the dominant basalts of Masafuera, but
is singularly rich in the basaltic dikes which in hundredfold vertically traverse
the island.

The rocks prevailing at heights up to c. 1000 m may for the greater part
be classified as dark vesicular feldspar basalts, with large phenocrysts of labradorite
(Ab 45, An 55). Olivine is scarce as phenocrysts but occurs in varying amount
in the groundmass together with augite, labradorite and magnetite in a dark,
glassy matrix (Fig. 16).

In many specimens of the vesicular lavas the vesicles are filled or lined with
zeolite minerals. In some samples the vesicles present, from the outward rim,
first a coating of glass, followed by chalcedon and chabasite and a central
replenishment of well developed natrolite spherolites.

At an elevation of about 1000 m a lava bed of a very different aspect is
met with. In contrast to the dark basalts of lower horizons, the rock now in
question is light grey in colour. Large phenocrysts of olivine (up to 5 mm in
diameter) and labradorite are uniformly distributed in a very fine-grained ground-
mass, consisting of augite, slender laths of labradorite, abundant small grains of
magnetite and a small amount of a nearly colourless glass (Fig. 17).

In outward appearance this lava has a very singular aspect. The surface
feels rough and grainy which, together with the light grey colour, at first gives
the impression that the rock would have a trachytic or trachy-andesitic compo-

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 59

—e

Fig. 16. Vesicular feldspar basalt. x 8. Normal development under 1 ooo m elevation. Masafuera.

sition. The analysis (Table I, No. 7), however, shows that the lava must be
assigned to the basaltic group, though in many respects it obviously differs from
all the dark basaltic rocks of lower elevations on the island. With some hesitation,
in my former publication I referred the rock to the basanite group, assuming
that a content of virtual nepheline might be concealed in the groundmass, which
was readily affected when treated with HCl. This circumstance Lacroix assumes
to be characteristic for the basanitoid lavas of the Pacific, where he says the
nepheline often ‘‘n’est pas exprimée; elle est restée a l'état potential dans la verre.
La poudre des ces roches, traitée par HCl s’attaque plus au moins facilement”
(C. R. 169, 1919, p. 402). BARTH has, in his paper Pacificite, an anemousite
basalt’, suggested that the virtual nepheline may be concealed in the plagioclase
in the form of carnegieite. For such basalts he proposes the name pacificite, or
if olivine is present, olivine pacificite and on account of my former description,
refers the lava in question from Masafuera to the pacificite group (21, p. 60;
B24 p. 380,510).

In publishing the description of the rocks from the Juan Fernandez Islands
in my earlier publication the analyses were not recalculated according to modern
methods. This has been done now and the results recorded in the appendixed
table of analyses. With regard to the rock, then tentatively named basanite, the
calculated norm shows no nepheline and an excess of 1.99 % of aluminium. This,
together with a content of 3.36% H,O, gives the impression that the rock is de-
composed. On this account Lacroix, in reproducing my analyses of the Juan
Fernandez rocks, omitted this analysis, ‘‘car les résultats ne correspondent pas

60 PERCY QUENSEL

Fig. 17. Olivine basalt. Nat. size. Elevation c. 1000 m. Masafuera. The olivine phenocrysts more
or less completely altered to deuteric iddingsite.

a la composition minéralogique décrite; le calcul fait voir qu/il s'agit d’une roche
alterée’’ (14, p. 65).

A renewed scrutiny of the rock has offered new aspects regarding its petro-
logic and petrographic position. The essential point has been that the large olivine
phenocrysts are altered to such a great extent to iddingsite. We have every inter-
mediate phase from a more or less broad rim of iddingsite around a kernel of per-
fectly fresh olivine to complete pseudomorphs of iddingsite, retaining the crystal
habit of the olivine. The circumstance that the kernels of olivine are absolutely un-
altered, leads to the conclusion that the iddingsite is not a product of normal
weathering but a deuteric mineral, derived during a final stage of cooling of the
lava in which it occurs.

During later years several papers have been published, in which the origin
of iddingsite has been discussed in detail. The conceptions, there put forth, to
all intent conform with the characteristic features of the lava from Masafuera.
As the petrogenesis of the rock in question is of a certain interest, some signi-
ficant quotations may be given, relating to the formation of iddingsite from
different localities.

Ross and SHANNON summarize their conclusions as follows: “‘Iddingsite is
not confined to weathered surfaces; its development shows no proximity to joint
cracks and evidences of weathering in associated minerals is entirely absent. Thus
it is concluded that iddingsite is not a product of ordinary weathering but
a deuteric mineral, that is to say the result of metasomatic processes associated
with the later stages of a cooling magma.’ They also emphasize that the
magma must have come to rest before iddingsite formed, for though it is a
very brittle mineral, it is never fractured or distorted by flow (U. S. National
Museum, Proc. 67, 1925, Art. 7, p. 18).

AUROUSSEAU comes to the conclusion that “iddingsite is the result of oxida-
tion processes that acted rapidly on the olivine during the liberation of copious

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 61

active volatile phases at the time of eruption” (Linnean Soc. of N.S. W., Proc. 51,
1926, p. 617).

Finally EbWARDs in his paper “The formation of iddingsite’ says: “‘It is
concluded, therefore, that iddingsite forms during or after extrusion, according
to the temperature of the magma at the time of extrusion, and that, if the magma
has cooled sufficiently before extrusion, not enough iron oxides are left in the
residual volatiles for iddingsite to be formed’ (Am. Min. 23, 1938, p. 281).

The formula for iddingsite is given by Ross and Shannon as MgO- Fe,O3-
3 SiOg-4 H,O with magnesia replaced in part by CaO, approximately in the ratio
1:4. The calculated composition according to this formula includes 7.g0 % H,O*
and 7.90 % H,.O7 (ibid. p. 17).

All the authors, now quoted, come to the same conclusion that the forma-
tion of iddingsite cannot be regarded as the result of normal weathering processes
but that the mineral must be of deuteric origin, formed during the later stages
of a cooling magma. The development of abundant iddingsite in the lava flow
from Masafuera leads to the same conclusion. I may also remark that most of
the photomicrographs, representing different stages of alteration from olivine to
iddingsite, published in Pl. I—II in Ross and Shannon's paper, correspond in
nearly every detail with equivalent alterations in the lava from Masafuera (Fig.
18). As iddingsite according to the formula of Ross and Shannon contains
15.80 % HO, we may conclude that an appreciable amount of the 3.46 % H,O,
according to the analysis of the lava in question (Table I, No. 7), enters the
iddingsite molecule.

Another singular alteration in the iddingsite basalt in question is also worthy
of notice. The slender labradorite laths in the groundmass as well as some
phenocrysts of the same mineral are found to be more or less completely
altered to an isotropic substance, though retaining their crystal habit perfectly
intact. I had observed this circumstance in my former publication on the Geology
of the Juan Fernandez Islands (12, p. 278) and then came to the conclusion that
the invading material was glass, formed through fusion of the feldspars in the
volcanic vent (Fig. 19).

Dr OrTo MELLIs has kindly undertaken a renewed investigation of this
isotropic alteration product. At first he also came to the conclusion that it con-
sisted of glass, formed by fusion of the feldspars during a period of high-temperature
autometamorphism. A more detailed study, however, led to different conclusions.
Dr Mellis gives the result as follows: “Repeated determinations showed that the
refractive index of the isotropic material lies between 1.49 and 1.50. These values
must exclude the presence of glass, formed by fusion of a labradoritic feldspar.
The index of refraction should according to Franco and Schairer in such a case
lie between 1.53 and 1.54 (Journ. of Geology, Vol. 59, 1951, p. 266). A further
evidence in this respect is that, when treated with HCl, the isotropic matter
readily gelatinized, which is not in accordance with what one can assume to be
the case with glass, formed by fusion of a basic plagioclase. Opal must for the
same reason be excluded as a possible component. Continued investigations
resulted in the conclusion that the alteration of the feldspars in the iddingsite

62 PERCY QUENSEL

Fig. 18. Iddingsite forming sharp outer borders around unaltered olivine. 100. From specimen
Fig. 17. Elevation 1000 m. Masafuera.

Fig. 19. Phenocryst of labradorite, traversed by a zeolitic matter. x 170. From specimen
Fig. 17. Masafuera.

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 63

basalt must be attributed to an invasion of zeolitic matter. As there seems to
be a slight difference in refractive indices in the alteration product, this is prob-
ably a mixture of different zeolites or related material. It has, however, not
been possible to determine the definite nature of the zeolites. The fact that they
show no sign of birefringence indicates that they must belong to the isotropic
group or to those of very near zero birefringence.’

It is of interest to note that Renard evidently has observed a very similar
circumstance in a lava flow from Masatierra, when he says that sections of a
basic feldspar from a greyish very scoriaceous lava under the microscope are
seen to be “cracked and pervaded with zeolitic matter, which forms an irregular
network. This matter, which looks slightly grey, when seen in ordinary light,
remains obscured between crossed nicols.’’ This description might as well refer
to the zeolitisation of the lava bed from Masafuera, especially as Renard in the
same lava mentions olivine, uniformly changed to iddingsite (Renard says hematite,
8, p. 176).

It therefore seems probable that lava beds, in which the content of olivine has
been more or less changed to deuteric iddingsite, also have succumbed to a high
degree of zeolitisation. The fact that the large phenocrysts of olivine, only
rimmed with iddingsite, show no signs of alteration, whereas the feldspars in the
same section may be completely altered to an isotropic substance of zeolitic
composition, must indicate that also the zeolitisation of the rock has taken place
at an early stage and cannot be attributed to periods of normal weathering or
later thermal activity.

In consequence of the changes in chemical composition which must have
taken place in connection with the formation of iddingsite and the zeolitisation
of the lava, it is no longer possible to establish the primary composition of the
rock in question. Though the whole aspect of the lava bed so obviously differs
from the normal olivine basalts on Masafuera, there is evidently not reason enough
with any degree of certitude to classify the rock as an olivine basanitoid. The
presence of alkaline rocks on Masafuera must for the present, therefore, be
restricted to the occurrence of the soda-trachytic lava flows, now to be described.

At an elevation between I 000 and 1 100 m there occur rocks of quite another
type. In outward appearance they are dense, aphanitic and aphyric in texture
and of a light yellowish green colour. The dominant minerals are an acid plagio-
clase and, subordinate, orthoclase. Olivine, a light green diopsidic pyroxene, and
ore minerals are sparingly found in small individuals between the feldspar laths,
which have a well-defined trachytoid orientation (Fig. 20). An analysis of this
rock was made for my former description and is reproduced in the annexed table
of the analyses of the Juan Fernandez rocks (Table I, No. 8). I named the rock
soda-trachyte.

On my survey I had assumed that rocks of this composition only existed
on and formed the highest parts of the island, as at that time no higher elevations
‘were reached. New specimens in Skottsberg’s later collection disprove my earlier
‘supposition in two essential respects. In the first place, rocks of much the same

04 PERCY QUENSEL

Fig. 20. Soda trachyte. x 50. Elevation c. 1050 m. Masafuera.

Fig. 21. Soda trachyte (boulder). x 30. Nic. +. Elevation about 400 m.
Tierras Blancas, Masafuera.

character as the soda-trachyte have now been found at lower levels. Secondly,
Skottsberg reached the summit of the island and was there able to establish that
basaltic lavas occupy the whole highest part of the island.

The new find of a rock, closely related to the soda-trachyte from higher
levels, was a loose boulder at the foot of Tierras Blancas. According to Skotts-

ISLANDS

FERNANDEZ

THE JU

GEOLOGY

THI

COMMENTS ON

ADDITIONAL

iy phenocrysts of olivine.

20 m elevation.

basalt with strz

ar phyric feld

. Vesicul

Masafuera.

cimen from

150m.

ration C.

ley

x 50. I

structure.

Masafuera.

Aphyric feldspar basalt. Column

2B.

Fig.

EI! Ovalo,

66 PERCY QUENSEL

berg’s information, boulders of this character were very numerous in the talus
under a steep escarpment and can here hardly have originated from higher eleva-
tions than about 400 m.

In outward appearance this rock shows an aspect different to the compact
soda-trachyte between 1000 and 1 100 m, which on account of its dense texture
at first sight gives the impression of a quartzitic rock. The new specimen has
on the other hand a rough, loose-grained structure. Minute intergranular cavities
in the rock are seen under the microscope to be filled with limonitic matter. But
allowing for this structural dissimilarity the rocks from the talus below Tierras
Blancas seem to be closely related to the soda-trachyte as described above (Fig. 21).
Taking into consideration the seemingly horizontal position of the lava beds of
Masafuera, the conclusion must be that lavas of soda-trachytic composition have
been emitted at different times and that they have alternated with lavas of more
normal basaltic composition (see p. 71).

Specimens of the rocks from between I 100 and I 400m present again a
different aspect. Five samples from 1100, I 200, I 300 and 1420m all have
very much the same appearance. They are all pbyric lavas of an ash-grey colour.
Feldspar and, more rarely, olivine occur as phenocrysts in a groundmass of
basic plagioclase, augite and abundant magnetite. In a somewhat vesicular specimen
from I 100 m olivine is also present as sporadic grains in the groundmass.

The groundmass has in most of the specimens a trachytic texture; the narrow
laths of feldspar circuiting the phenocrysts in a more or less well-defined
manner (Pie. 22).

In my previous paper I named the rocks of this nature trachyandesite.
Hagerman says, referring to much the same specimens in Skottsberg’s collection:
“Die Klassifizierung hiehergehGriger Gesteine ist etwas unsicher. Geniigende
Griinde sie als Trachyandesite zu bezeichnen, liegen jedoch nicht vor” (13, p. 30).
Hagerman names the rock andesite in his paper.

These rocks evidently have a rather wide distribution on Masafuera, probably
forming the whole complex of lava beds between 1 100 and 1420 m. To certify
their petrographic position an analysis has now been made of a typical specimen
(Table I, No. 5). The calculated norm and Niggli values definitely classify the
rock as a feldspar basalt, in many respects of similar composition as some of
the analysed basalts from Masatierra (Table I, No. 1—4). A difference of interest
is the still lower content of K,O; the Niggli value £ is now 0.08 against 0.15—-0.20
in the basalts from Masatierra.

These phyric feldspar basalts from Masafuera in general aspect also very
much resemble the aphyric feldspar basalts from between 400—600 m elevation
on Masatierra. Occasionally at lower levels they also on Masafuera may be aphyric
in texture (Fig. 23). They seem also in several respects to correspond to certain
phyric and aphyric basalts from the Hawaiian Islands, as described by WASHINGTON
(Petrology of the Hawaiian Islands. I. Kohala and Mauna Kea. Am. J. of Sc. V,
1923, p. 487). To this question I will return under a concluding heading dealing
with the chemical and petrological connections between the rock of the Juan
Fernandez Islands and other volcanic islands of the eastern Pacific.

ADDITIONAL COMMENTS ON THE GEOLO¢

THE JUAN FERNANDEZ ISLANDS 07

Fig. 24. Olivine basalt, supersaturated with iron oxide. 35. Vesicular flow-breccia. Los Inocentes
§ : 35
elevation c. I 50om), Masafuera.

Fig. 25. Olivine phenocryst from specimen Fig. 24. 110. Specks and streaks of unaltered
olivine as rests in a pseudomorph after olivine. Los Inocentes, Masafuera.

68 PERCY QUENSEL

Fig. 26. Disintegrated phenocryst of olivine, caused by high temperature oxidation. The rim
around the olivine crystal is hematite. Within the olivine the hight grains are mineral components,
formed by exsolution. 150. Los Inocentes, Masafuera. Photo P. Ramdohr.

Fig. 27. Part of disintegrated olivine phenocryst, enlarged 600. Against the dark background
of olivine the exsolution product is seen in the form of small composite grains consisting of
hematite (light) and a darker undefined component. Larger light grains see PI. Il, Fig. 1.
Photo P. Ramdohr.

ADDITIONAL COMMENTS ON THE GEOLO O! LHI UAN FERNANDEZ SLANDS O ¢

Fig. 28. High-temperature exsolution of spinel, enclosed in olivine phenocryst ‘Pl. IJ, Fig. 2). Light
lamellae, according to Ramdohr probably magnetite in groundmass of excess spinel. 600.
Photo P. Ramdohr.

Fig. 29. Magnetite in groundmass of the lava bed as seen in PI.], Fig. 1 partially changed to
hematite (martite 300. Photo P. Ramdohr.

5 516795 I Nat. H Juan Fernandez and Ea

TO PERCY QUENSEL

An observation of importance with regard to the distribution of different
lava beds on Masafuera is the find of a deviating type of olivine basalt from Los
Inocentes. According to Skottsberg’s observations such lavas probably occupy
the highest part of the island, representing elevations above 1420 m. The few
samples brought back are highly scoriaceous flow breccias containing a high
content of iron oxides (Fig. 24). Hagerman has in the preceding publication
of this series given a description of these lavas which he characterizes as slaggy
olivine basalts with large olivine phenocrysts, supersaturated with magnetite

(Geshe DP 2115)

The phenocrysts of olivine are under the microscope found to be almost
opaque, due to the precipitation of new-formed ore minerals. A varying amount
of residual olivine is, however, nearly always to be observed in the form of specks
or streaks (Fig. 25). No signs of alteration are to be observed in this olivine.
Optical determinations indicate that only a low content of about 8 % FeO is
present in the molecule.

The groundmass consists of slender laths of labradorite, small grains of augite,
magnetite, ilmenite and pseudobrookite in a dark brown glass matrix.

To determine the mineral composition of the pseudomorphs after olivine
Professor S. GAVELIN and Dr UYTENBOGAARDT kindly undertook to examine some
polished sections of the rock. Professor P. RAMDOHR (Heidelberg) contempora-
neously supervised a section for the same reason. It thereby became apparent
that the seemingly opaque constituent was not magnetite and that the mineral
assemblage of the pseudomorphs was of a complicated nature. Professor Kamdohr
has taken four photomicrographs thereof and kindly put them at my disposal.
They are reproduced in Fig. 26——29 with Ramdohr’'s explanatory notes. In Fig. 26
the essential components can be observed. A rim of hematite is seen to encircle
an idiomorphic crystal of olivine with specks of disintegrated minerals. In the
enlarged microphotograph Fig. 27 these minerals are seen in the form of small
lighter grains uniformly distributed against the dark background of olivine. One
can now observe that the grains consist of two constituents. The one component
is hematite. Kk peated attempts have been made to determine the second com-

ponent both 1: + lished sections and with X-ray powder photographs. No con-
clusive evidei regarding the true nature of this mineral has, however, been
attained.

The singular alteration of the olivine phenocrysts must in all probability be
connected with the same processes as have controlled the formation of the deuteric
iddingsite, though in the samples at hand this mineral is not extant. Edwards
seems to have described a very similar formation in the iddingsite-bearing basalts
from two Victorian localities in Australia. After concluding that the iddingsite
must have been formed before the ultimate consolidation of the lava flow, Edwards
says: “In some instances, however, the action has gone further, and a rim of iron
oxid is formed on the outer margin of the iddingsite. Eventually all the original
olivine vanishes, and the iddingsite, which had formed a rim about it, is completely
replaced by magnetite ... It is essential for the formation of iddingsite that the
magma should not only be rich in water vapour, but that it should have differentiated

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS ex

in such a manner as to give rise to an iron-rich final fluid’ (Am. Min. 23, 1938,
p. 280). A photomicrograph in the text shows, according to Edwards, ‘‘a pseudo-
morph of iron ore after iddingsite, itself a pseudomorph after olivine’.

These suppositions seem also referable to the very similar formation on
Masafuera. The lava bed in question contains 33.86 % FeO according to deter-
mination by LANDERGREN (13, p. 33). This is more than double the content of
FeO in any other basalts of the Juan Fernandez Islands. There seems little doubt
that the pseudomorphs in question have formed under much the same conditions
as advanced by Edwards. This is furthermore supported by the nearly identical
appearance of the photomicrographs of the pseudomorphs from the Australian
localities and from Masafuera, in both cases in connection with lavas containing
iddingsite. Edwards assumes that, via an intermediate stage of iddingsite, the ulti-
mately formed component is magnetite. As narrated above, this is not the case
in the lava from Masafuera, where the pseudomorphs are found to be of a more
complicated composition, formed without iddingsite as an intermediate phase.

All the observations given above seem to indicate that the iddingsite-bearing
basalt from elevations about I 000 m as well as the flow breccias from the highest
elevations on Masafuera have been subjected to an automorphic re-mineralisation,
prior to the final consolidation of the magma. This would suggest that the volcanic
eruptions have been interposed by periods, during which the lava in a molten
state has temporally stagnated in the volcanic vent under conditions which in
connection with active volatile phases have led to the formation of such deuteric
minerals as iddingsite and to the partial high temperature exsolution of the oli-
vine phenocrysts.

The conclusions which may be drawn regarding the distribution of the
different lavas of Masafuera would be now that rocks of more normal basaltic
composition, principally feldspar basalts, occupy the lower and intermediate eleva-
tions of the island, whereas the highest parts consist of an olivine basalt, super-
saturated with iron oxides. At some intermediate elevations lavas of a more
alkaline character are found in the form of at least two beds of soda-trachyte,
interposed between dominant flows of basaltic lavas.

Horizontal distribution of different types of lava on Masafuera.

in m X100 Ol Hh Be Be I GP OE Ho wie Ta NS TA Goo

| Olivine basalt supersaturated |

with marnetite 3) 30's. 2
Light grey phyric feldspar |

Beers lie Pins ee meee tec eae | x
| et trachyte: | 20 fe SLs x |

Iddingsite-bearing phyric oli-
mmnenbasalt!.<. i) its)

Dark phyric and aphyric
feldspar- and olivine basalt) x

eee SLO705

~I
N

PERCY QUENSEL

Fig. 30. Picrite basalt (masafuerite). Dike rock. Nat. size. Loberia vieja, Masafuera.

Hagerman has recorded in his paper in tabular form his conception of the
horizontal distribution of the different lavas of Masafuera. This is reproduced on p.
71 with some slight corrections on the base of renewed examination.

My former supposition that the basaltic lavas only occur up to an elevation
of about 1000 m, from there on being succeeded by more alkaline rocks of
soda-trachytic composition, is no longer in agreement with more recent observations,
based on Skottsberg’s new collections. Any thought of gravitative differentiation
to explain the sequence of the volcanic rocks, which I tentatively proposed in
my former publication, must in the light of later observations be discarded now.

The numerous basaltic dikes, traversing the whole island in a West-East
direction, are worth special notice as representing rocks exceptionally rich in
olivine. In this respect they exceed the most olivine-rich picrite basalts from
Puerto Frances on Masatierra. BOWEN has commented on these rocks as follows:
“One other rock may be mentioned in this connection. It is a picrite basalt from
Juan Fernandez, a dike, not a lava, but quenched so as to reveal the fact of its
origin. In it is shown the highest amount of normative olivine (53 %) of any rock
termed basalt by the author describing it. Great crystals of olivine lie in
an aphanitic ground composed mainly of plagioclase and augite (Fig. 30—32).
Some of the olivine basalts of this island group are, locally at least, about
as rich in olivine as this dike, but they have not been analyzed. Their high
olivine content is invariably due to an increased amount of phenocrysts of
olivine about I cm in diameter. Plainly these crystals were not in solution
in the dike or flow material at the time of its intrusion or extrusion. This

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS g [ee

A. a , - , < Que :
‘ eo

pf
Vw a
a

MS,

Fig. 31. Masafuerite dike. x 9. Loberia vieja, Masafuera.

gs ce et) 8
RTE ee a

Be ‘

Fig. 32. Masafuerite dike. x 9. Analogous to sample Fig. 31 but with rims of iddingsite around
all phenocrysts of olivine. Loberia vieja, Masafuera.

74 PERCY QUENSEL

fact does not prove that they were not in solution in that material at an earlier
time. But if one finds the condition shown by these basalts to be invariably
true of all rocks rich in olivine, which have suffered quenching, one must
conclude that large amounts of olivine occur in solution in magmatic liquids.
A survey of igneous provinces leaves no question that such rocks do have this
character, that is, they always contain either all of their olivine or all in excess
of a quite small amount (apparently some 12—15 %) as relatively large pheno-
crysts. They, therefore, force acceptance of the stated conclusion” (23, p. 163).
The quotation above refers to purely theoretical questions but indicates the extreme
position these rocks held in petrographic classification with regard to the abnor-
mally high content of olivine in basaltic lavas. JOHANNSEN has named these rocks
masafuerite, with the following argument: ‘‘The picrite basalt from Masafuera of
the islands of the Juan Fernandez group is a most extraordinary rock ... While
this particular rock occurs as a dike, on the adjacent island of Masatierra, for example
at Puerto Frances, there is a similar rock in the form of a lava flow with large
olivine crystals in a groundmass containing more or less the same mineral. I am
placing the rock among the hypabyssals on the basis of the occurrence on Masa-
fuera. To all olivine-melabasalt dikes which contain more olivine than any other
mineral and in addition carry basic plagioclase and augite, I should like to apply
the name masafuerite” (24, p. 334).

We already have the name picrite basalt for the lava flows of much the same
composition on Masatierra, with Lacroix’s name oceanite as synonym, given with
the following definition: “les roches basaltiques porphyriques a olivine sont parfois
extraordinairement riches en péridot; dans l’échantillon analyse, tous les grains
de ce minéral se touchent, ils sont réunis par une petite quantite de plagues de
labrador, englobant des microlites d’augite et des lames d’ilmenite’ (14, p. 44).
Johannsen has restricted the name masafuerite to aschistic dike rocks of a
picrite basaltic magma.

These dikes may represent, at least in part, transmission channels for the
upper basalt beds, supersaturated with iron oxides. A significant feature in this
respect is that they contain numerous ‘schlieren’ of darker colour, due to abundant
minute grains of magnetite. These streaks may indicate relics from a magmatic
flow, subsequently consolidated in the shape of the olivine basalt, supersaturated
in iron oxides, which now forms the highest parts of the island. At a later period
the channels may then have been filled with the melanocratic magma, which now
characterizes them as such singular rocks. Sequent intrusions of this nature might
lead to phases of crystallisation in accordance with Bowen’s conception of these
dikes as quenched rocks, referred to above.

Regional Relations.

Tectonic Connections.

In the introductory lines I already noted that during recent years as well as
in older reports speculations have been offered regarding connections in one or

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS I

other respect between the Juan Fernandez Islands and the volcanic islands of the
eastern Pacific. Some quotations may be given.

CHARLES DARWIN already expressed views regarding geotectonic connections
between the Juan Fernandez Islands and, on the one side, the South American
continent, on the other, the Galapagos Islands. In his “Observations on the
volcanic islands and parts of South America’ he says: “Some authors have
remarked that volcanic islands occur scattered, though at very unequal distances,
along the shores of the great continents, as if in some measure connected with
them. In the case of Juan Fernandez, situated 330 miles from the coast of Chile,
there was undoubtedly a connection between the volcanic forces acting under
this island and under the continent as was shown during the earthquake of 1835.
The islands, moreover, of some of the small volcanic groups, which border the
continents, are placed in lines, related to those along which the adjoining shores
of the continent trend; I may instance the lines of intersection at the Gala-
pasos) (20, p. 144).

In his paper ‘Constitution lithologique des iles volcaniques de la Polynésie
Australe’ LACROIX gives expression to much the same trend of thought when
he writes: ““Particulierement intéressantes sont les iles volcaniques qui se trouvent
aA une plus ou moins distance de l’Amérique du Sud, les iles Juan Fernandez,
San Felix et San Ambrosio, et enfin Galapagos, puis au large de l’Amerique
centrale, l’ile Clipperton. Bien que la connaissance de la lithologie de ces iles
soit loin d’étre complétement éclaircie, on peut a présent assurer que leur laves
différent de celles des Cordilléres des Andes, c’est-a-dire de la série circumpacifique
et montrer qu’elles se rattachent a la série intrapacifique’ (14, p. 64).

In ‘La Face de la Terre’ EMMANUEL DE MARGERIE refers to the same
subject as follows: “Le Relay de la marine des Etats-Unis a signalé, au large
de Valparaiso 5.651 métres. A l’ouest de ces fosses sont situées les deux iles
volcaniques anciennes de San Felix et San Ambrosio; au Sud de ces iles le
croiseur Chilien Présidente Pinto a trouvé, sur une étendue de 760 km, des
profondeurs si faibles qu'il est probable qu'une créte sous-marine, orientée a
peu pres N—S, s’allonge dans la direction de l'ile Juan Fernandez’ (ei slilh
p. 1359).

In his ‘Description and Geology’ of San Felix and San Ambrosio, BAILEY
WILLIS writes: “San Felix and San Ambrosio are volcanic islands in the South
Pacific Ocean, San Felix being situated in latitude 26°15’ south and longitude 80°7’
west of Greenwich and San Ambrosio lying about 16 km to the east-south-east.
They are about 500 miles west of Chafiaral on the east coast of Chile, and the
same distance due north of the group of Juan Fernandez and Mas-a-fuera. The
South Pacific charts show several rocks or islets and some whose existence is
recorded as doubtful, which, with the above-named islands, form an archipelago
strewn on a narrow submarine ridge that extends along the meridian of 80
degrees west from about 36 degrees south to 26 degrees south, the ridge being
defined by the 2.000-meter contour line. Knowing that all these islands and
islets are peaks of volcanoes, we may suspect that there are more of them than
we can see; but this must remain an unverified guess until detailed soundings

76 PERCY QUENSEL

can be made. The depth of the ocean in this region, which lies west of the
Richards Deep, varies from 4.000 to 5.000 meters. The islands, therefore, repre-
sent the summits of volcanoes probably sixteen to eighteen thousand feet or
more in height — that is to say, they compare with the volcanoes of the Andes,
which are situated on the other side of the deep” (26, p. 365).

In an interesting paper on the geology of Galapagos, Cocos and Easter
Island, L. J. CHUBB has published some noteworthy remarks on the regional
relations of the volcanic islands of the eastern central Pacific. He: writes as
follows: “‘Under the east central Pacific there lies a vast area, the Albatross
Plateau, under depths of less than 2.000 fathoms, though on all sides the depths
exceed this figure. No islands rise from the central part of this plateau, but at
each end is an archipelago that appears to have been built up on a set of
intersecting fissures, the Marquesas at the western end, and the Galapagos at
the eastern. On or near its southern margin too there are several volcanic islands,
including the Mangareva (Gambier) Archipelago, Pitcairn, Easter, Sala-y-Gomez
and the Juan Fernandez Islands.

“It is suggested that the plateau constitutes a resistant block which has
withstood lateral pressure that has been brought to bear on it from all sides,
that around its margin it has become cracked and fissured, and that on the
fissures volcanic islands have been erected. That these islands owe their origin toa
common cause is suggesed by the similarity of their structure and geological
history, so far it is known. ... Petrographically, too, these islands resemble each
other and differ from most of the other Pacific islands. The most striking charac-
teristics of their rocks are the almost complete absence of nepheline-bearing
types and the presence of virtual free silicia in many.

“Cocos, St. Felix and St. Ambrose islands are constituted in part of nephe-
line rocks, and for this reason they are regarded as lying, not on the resistant
block, but beyond its eastern margin. Petrographically they resemble the Society
Islands and Austral Islands which lie to the west of the plateau.

“It is thought that beyond the margins of the block the crust is more
pliable and has yielded to pressure, with the formation of anticlines and synclines.
Volcanoes that have produced nepheline-bearing rocks have been erected on the
anticlines. The folds have tended in the western area to migrate from southwest
to northeast with a wave-like motion proved by the history of their coral reefs.
There is not sufficient evidence, however, to determine whether the folds which
probably underlie Cocos, St. Felix and St. Ambrose islands have suffered a
similar movement” (27, p. 43).

JUAN BRUGGEN has recently in his book ‘Fundamentos de la Geologia de
Chile’ discussed the geotectonic position of the Juan Fernandez Islands. He
writes: ‘Esta zona (la region situada al este de Llico, en Arauco) de dislocaciones
tan extrafias a la structura de la Cordillera de los Andes, coincide con la region
donde una ancha loma submarina se desprende del continente. Encima de la
loma se levantan las islas Juan Fernandez y mas al norte las de San Ambrosio
Vesa belize

Parece que se trata de una antigua cordillera que se separo del actual conti-

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 7

~—I

nente entre Magellanes y Arauco y que se hundio posteriormente. A este cordillera
o simplemente zona continental, que llamaremos ‘Tierra de Juan Fernandez’, se
debera el término de las capas de la Quiriquina en el sur de Arauco y también
el hecho que sedimentos marinos de Koceno no se conozcan mas a sur.

Pero en el Oligoceno, cuando la costa del piso de Navidad se extendio
hasta la region de Yptin (45° L.S.), se habia hundido una gran parte de la
Tierra de Juan Fernandez, conservandose probablemente cierta extencion en la
vecindad de las islas volcanicas, cuyas rocas se formaron solo mas tarde en
erupciones posteriores. A juzgar por el grado de denudacion y en vista de las
actividades volcanicas recientes en Mas a Tierra y San Felix, la parte volcanica
de estas islas se habra formado en el Terciario superior, probablemente en el
Plioceno, cuando existia todavia un resto de la antigua Tierra de Juan Fernandez,
de la cual immigr6é la flora del Eoceno. Cuando mas tarde se hundio también
este resto, sobresalian solamente las partes volcanicas, constituyendo las islas de
Juan Fernandez, que servian de refugio para la flora’ (16, p. 59).

The references now given suffice to show how the position of the Juan
Fernandez Islands has from different points of view been geotectonically connected
with other volcanic islands or groups of islands of the eastern Pacific.

Petrographic Connections.

In many papers of recent years, petrological and petrographical connections
between the rocks of the Juan Fernandez Islands and those of other volcanic
islands of the Pacific Ocean have also been the subject of discussion. The types
of lava which in this respect have been of special interest are the extremely
melanocratic picrite basalts (oceanites of Lacroix) and their occasional combi-
nations with more alkaline rocks. A short summary of the literature on this
subject may be given first.

We may conveniently begin with the islands of San Felix and San Ambro-
sio, which geographically lie nearest. The distance is 760 km due north of Juan
Fernandez. H.S. WASHINGTON has given a petrographical description of the
rocks. I may quote some lines from his general conclusions: “It would appear
from the specimens brought back by Willis that the lavas of San Felix volcano
are, so far as known, only of two kinds — a decidedly sodic trachyte and a
somewhat variable nepheline basanite, which seems to be highly vitreous. There
is little doubt that the yellow tuff is derived from a nephelite basanite magma
closely similar to that of the flows. The prominent characteristic of these two
types of lava is their high content in alkalies, especially in soda, while high
titanium and phosphorus appear to be other constant characters of minor but
still considerable interest. This conclusion as to the generally highly sodic
character of the San Felix lavas is subject to the limitations imposed by the
absence of specimens from the lower flows and from various parts of the island.
Such basaltic lavas, especially if highly vitreous, may appear megascopically to
be very uniform and yet modally and chemically very diverse. It is, therefore,

78 PERCY QUENSEL

possible that earlier, lowermost flows are less sodic and more typically basaltic
than the upper, which were the ones examined.’’ Washington continues: “In
this predominantly highly sodic character of the lavas San Felix appears to
differ widely from other Pacific islands. At Masafuera, it is true, both soda
trachyte (and nepheline basanite)', closely like those of San Felix, occur, but
these are accompanied by basalt and picrite basalt, whereas at the neighbouring
Juan Fernandez (Masatierra) the lavas appear to be, to judge from Quensel’s
description, only olivine basalt with neither trachyte nor basanite. Trachyte, also
highly sodic, occurs at several other Pacific volcanic islands, as do also nephelite
basanite and similar rocks high in soda; but at all of them the predominant
lavas are more or less normal basalts or andesites; so that the general magmatic
character is basaltic — that is to say, sodi-calcic, somewhat modified by distinctly
sodic facies” (26, p. 382).

Referring to the trachyte of San Felix Washington says that “‘in thin section
the rock shows a somewhat peculiar texture, which resembles that of the trachyte
of Masafuera described by Quensel, that of Puu Anahulu on the island of Hawaii
and of the trachyte of Lahaina on Maui. The texture seems to be rather usual
in the trachytes of the Intra-Pacific volcanic islands. Ill-defined laths of alkali
feldspar make up most of the rock. Most of these are arranged irregularly, but
here and there flow texture is evident’ (26, p. 375).

In several places in his paper ‘La constitution lithologique des iles vol-
caniques de la Polynésie Australe’ Lacroix compares the petrographic character
of the lavas of the Juan Fernandez Islands with those of other volcanic islands
of the Pacific. Concerning the basalts of the Galapagos Islands he observes that
they “offrent l’analogie la plus grande avec les basaltes de Masatierra et avec
ceux des iles Gambier, c’est-a-dire avec les plus calciques des basaltes du
Pacifique et les plus pauvres en potasse’”’ (14, p. 68). And further on in the same
paper: “Les iles Juan Fernandez se groupent au voisinage des iles Marquises et
l'on a vu quil faut comparer ce que l'on sait des roches des iles Galapagos aux
données concernant ceux des iles Gambier” (14, p. 77).

These conclusions of Lacroix refer to the normal basalts of Masatierra and
are founded on the four new analyses of such basalts. The similarity in respect
of the Gambier Islands, however, goes a step further, as Lacroix describes from
there, associated with more normal basalts, typical oceanites (picrite basalts),
which, as we have seen, also occur on Masatierra.

A third author who has brought the Juan Fernandez rocks under discussion
with reference to chemical similarities with other volcanic islands of the Pacific is
CONRAD BurRRI. Under the title ‘Chemismus und provinziale Verhaltnisse der
jungeruptiven Gesteine des pazifischen Oceans und seine Umrandung’ he coor-
dinates under the heading “Typus Hawaii” (in contrast to ““Typus Tahiti’’), the
rocks of Hawaii, the Leeward group, Juan Fernandez and Samoa, remarking that
the Juan Fernandez, San Felix and San Ambrosio lavas are good representatives
of the group, the basanite from Masafuera, however, showing a small deficiency

* This name now discarded (see p. 63).

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 79

in al — alk (28, p.177). — This deficiency is now explained by the fact that
the analysed specimen has attained an abnormal chemical composition through
the formation of deuteric minerals (see p. 60).

A second part of Chubb’s paper on the geology of Galapagos, Cocos and
Easter islands contains the ‘Petrology of the Galapagos Islands” by C. RICHARD-
SON. Under a concluding heading he says: “The Juan Fernandez Islands are the
only islands on which both types of basalt (porphyric with dominant phenocrysts
of olivine or with basic plagioclase) are found in addition to soda trachyte similar
to that occurring in the Galapagos Archipelago. Although oceanites (and basanitic
lavas) are also present, the Juan Fernandez Islands are petrologically closer to
the Galapagos than any other islands” (27, p. 64).

General Conclusions.

A characteristic feature of the volcanic rocks of the Pacific is the universal
predominance of sodium over potassium. LACROIX says that in this respect
“toutes les roches étudiées présentent la commune caractéristique d’étre plus riche
en soude qu’en potasse”’ (14, p. 55). BURRI comes to the same conclusion, based
on recalculations of all available analyses. He states that the Niggli value & is
always under o.4 and for the most typical regions of volcanic rocks of the intra-
Pacific Ocean under 0.25 (28, p. 173).

The value & in the four new analyses from Masatierra in no case exceeds
0.20 (average 0.17). And the light grey basalt from Masafuera has a still lower
content of K,O (Niggli value £& 0.08). Therefore, with regard to low percentage
of potassium the basaltic lavas of both Masatierra and Masafuera must be con-
sidered in this respect as representative for the basalts of the volcanic islands
of the Pacific.

In other respects the petrographic relationship between Juan Fernandez
and other intra-pacific islands has been interpreted somewhat differently. The
reason is, however, easy to explain. In some cases only the basaltic rocks
of Masatierra have been taken into consideration, in other cases special notice
has been given to the soda-trachytic lavas of Masafuera as indicating a casual
presence of more alkaline rocks. Since no rocks of alkaline character occur on
Masatierra, this island has petrographically been connected most closely with the
Gambier and Marquesas Islands as, according to Lacroix, representing “les plus
calciques des basaltes du Pacifique et les plus pauvres en potasse”’ (14, p. 68 and 77).

In chemical composition the basaltic lavas of the Juan Fernandez Islands also
show similarities with some of the basalts from the Hawaiian Islands. Several
analyses of phyric and aphyric feldspar basalts from Kohala and Maunakea as
well as from Kilauea, published by Washington are very similar to those of
the basalts from Juan Fernandez (Am. J. of Sc. 5, 1923, p. 482—87 and 6,
p. 341). It can be suggestive that together with both ancient and recent lavas
of Kilauea, Washington also describes chrysophyric picrite basalts of much the
same character as those from Masatierra and Masafuera. Another similarity can
be given. WHITMAN Cross has described, together with the normal basalts,

80 PERCY QUENSEL

soda-trachyte from Maui and Anahulu, on the first island in connection with a
‘picritic basalt’ (U.S. Geol. Survey, Prof. Paper 88, 1915, p. 26—28). This corre-
sponds exactly to the rock assemblage of Masafuera.

Petrographic description of the Hawaiian basalts from the mentioned localities
is, also in other respects, found to agree with both megascopic and microscopic
features of the basalts of Juan Fernandez. Washington describes an aphyric basalt
from Kohala as follows: “The type is a light grey, almost aphanitic lava, except
that some rare, very small feldspar phenocrysts may be present, and a few pheno-
crysts of olivine are seen in most specimens ... The texture is rough and trachytic,
so that the rock would probably be considered an andesite or trachyte in the
field. Most specimens are dense and very fine-grained or aphanitic, but vesicular
forms may occur” (I. c., p. 485). This description might as well refer to the light
grey basalts of Masafuera at elevations between I 100 and 1 400 m, which I also,
before an analysis was made, tentatively denoted as a trachy-andesite (12, p. 282).

Although certain lavas of Hawaii evidently present similarities with the basalts
from Juan Fernandez, the general assemblage has, however, a different character.
According to Washington “‘olivine-free labradorite basalts constitutes the most
abundant type, followed in abundance by andesine basalt and then by oligoclase
andesite’ (Am. J. of Sc. 6, 1923, p. 355). The high percentage of andesine basalt
and andesite denote a magmatic sequence differing from that of the non-alkaline
lavas of more southern latitudes of the Pacific. It may, therefore, be advisible for
the present to comply with Lacroix when he says: “Les roches de cette ile
(Hawaii) constitueraient une division spéciale, ayant une originalité propre”
zp 27/0).

If we take into consideration the assemblage of olivine and feldspar basalts
and soda-trachytic lavas on Masafuera as a characteristic feature for this island
it seems evident that, as Richardson already has assumed, the rocks of the
Galapagos Archipelago display the closest similarities. According to the analyses,
published by Richardson, both the basalts and the soda-trachyte are in chemical
composition very similar to equivalent rocks of Masafuera. Also soda-trachyte is
of the same scarce occurrence in the Galapagos Islands as on Masafuera, the
only sample being collected by Darwin on the Beagle voyage of 1835. Richardson
says: “Juan Fernandez are the only islands on which both types of basalt are
found in addition to soda trachyte similar to that occurring in the Galapagos
archipelago” (27, p. 64). The low content of potassium is in common for the
basalts from both island groups.

On the other hand we must evidently exclude any petrographical relationship
between Juan Fernandez and San Felix—San Ambrosio where the lavas have, as
far as is known, a more pronounced alkaline composition, classified by Washington
as soda-trachytes and nepheline basanites (l.c., p. 382). Richardson says: ‘The
Juan Fernandez are petrologically closer to the Galapagos than are any other
islands ... Both are situated comparatively near the American coast of the Pacific,
but their similarity is not shared by San Felix and San Ambrosio islands, or
any other islands on that side of the Pacific’ (27, p.64). Lacroix comes to the
same conclusion. He finds the closest connection with San Felix—-San Ambrosio

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 81

to be the highly alkaline rocks of Tahiti in the mid-Pacific (14, p. 77). Though
lying geographically nearest Juan Fernandez, the rocks of San Felix—San Ambrosio
evidently represent lavas of a more alkaline composition, with the exception of
Cocos Island, not otherwise met with among the volcanic islands of the eastern
Pacific.

We may conclude, therefore, that the lavas of the Juan Fernandez Islands
in chemical composition, and to a certain degree also with regard to their general
aspect and fluctuations in petrographic character, must be regarded as most closely
associated with the rocks of Galapagos and in some respects also with some
rock assemblages of the Hawaiian Islands, and in the more central parts of the
Pacific Ocean, with the Gambier and Marquesas island groups.

These connections are purely based on petrographical and chemical similari-
ties. It is of interest to find how these connections coincide with Chubb’s more
theoretical ideas regarding the regional distribution of the volcanic islands of the
Pacific. I refer to the quotation above (p. 76) and will here only recapitulate the
following: “No islands rise from the central part of this plateau (the Albatross
plateau), but at each end is an archipelago that appears to have been built up
on a set of intersecting fissures, the Marquesas at the western end, and the Gala-
pagos at the eastern. On or near its southern margin too there are several vol-
canic islands, including the Mangareva (Gambier) archipelago, Pitcairn, Easter,
Sala y Gomez and the Juan Fernandez islands.”

An approximately symmetrical distribution of alkaline and non-alkaline rocks
of the volcanic islands in the Pacific, which are situated between Lat. o° and
35. S. may, therefore, be assumed to encircle the Albatross plateau, itself so
obviously unendowed with any islands. The outward lying connections would
then include San Felix-Ambrosio and Cocos on an eastern marginal zone and
the Austral and Society Islands on the western margin. The rocks of these
islands all have a pronounced alkaline character and have been referred by Lacroix
to “la série néphélinique”’ of the Pacific islands. Inward zones of connection would
include Juan Fernandez—Galapagos and the Gambier (Mangareva) and Marquesas
island groups, characterized by absence of nepheline-bearing types and the presence
of virtual free silica in many. Lacroix unites them as belonging to “la série
sans néphéline” (14, p. 59).

A plausible explanation of these circumstances might be that tectonic dis-
turbances at different intervals have located cracks and fissures around a resistant
block, represented by the Albatross plateau. Volcanic eruptions of different
magmatic composition have then been localised to different areas, the alkaline
lavas being restricted to peripherical dislocations in contrast to the non-alkaline
lavas, located along inner lines of connection. Easter Island and Sala y Gomez,
composed of lavas of a deviating type and composition, may have been orientated
by intersecting fissures along other trends.

The climax of volcanic activity on all these islands belongs to past periods
of probably late tertiary or pleistocene age. However, several of the islands
bordering the American continent still manifest obvious indications of volcanic
nature. On San Felix volcanic gases were issuing from a crevice on the southern

82 PERCY QUENSEL

rim in May 1923. Bailey Willis says: “In this sense and to this extent we
may consider San Felix an active volcano” (26, p. 370). The Galapagos Islands
are still the seat of volcanic eruptions. As late as 1925 lava flows reached the
sea, pouring over the 100-foot cliffs (27, p. 9). The Juan Fernandez Islands
on the other hand show no signs of recent volcanic activity but, as recorded
above, one, and possibly, several sub-marine explosions have taken place in their
immediate vicinity during the past century.

The formation of fissures in locating the position of the volcanic islands
bordering the South American continent may be conceived to be connected with
dislocations of the oceanic sub-stratum as an after-sway of displacements during
the formation of the continental mountain ranges. Time connections between
violent earthquakes on the continent and volcanic activity on some of the adjacent
islands are significant in this respect.

However, much of what has been said regarding connections of the volcanic
islands of the Pacific, whether founded on petrological or petrographical similarities
or on geotectonic orientation, must still be considered as conjectural. For the
present, we must agree with Daly that “‘a glance at the larger aspects of Pacific
petrology shows how pitifully slight is our knowledge of the island petrography.
Now is not the time for settled convictions. Now is the time for concerted,
persistant effort, leading to a thorough exploration of the Pacific archipelagos,
under the auspices of a single institution with a staff of cooperating observers”
(R. Daly, Petrography of the Pacific Islands. Bull. Geol. Soc. of America, 27, 1916,

P2331):
Acknowledgements.

The writer is indebted to Professor P. RAMDOHR, Heidelberg, and to Pro-
fessor S. GAVELIN and Dr W. UYTENBOGAARDT for friendly cooperation in deter-
mining the ore minerals. Professor Ramdohr has taken some photomicrographs
of the same and kindly put them at my disposal for reproduction.

Dr O. MELLIS has helpfully co-operated in taking most of the other photo-
micrographs. The landscape photographs have kindly been put at my disposal by
Professor C. SKOTTSBERG. They are taken by him on his visits to the Juan
Fernandez Islands in 1908 and 1917.

A grant from the foundation LARS HIERTAS MINNE 1s gratefully acknowledged.

Mineralogical Department, University of Stockholm,
March 1952.

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 83

Table J.

Analyses of the rocks from the Juan Fernandez Islands.

Masatierra (I—IV) Masafuera (V—VIII)
I Il Ne ave V VI VII VIII
SiO}, ‘eeeidieu, sarees 46.08 46.10 46.50 47.62 AG OL 43237 43.47 63.43
LOE See ee 3.48 3.80 3.04 3.42 3.25 1.03 2.68 0.28
[OK cee aera ee eer 15.98 16.54 13.34 16.24 | 16.68 8.48 17.30 18.64
1? (0), Biaeeneeereree DGG 5.48 5.34 4.74 4.30 2.91 6.87 2.78
ISOM SA gin ks oe 8.83 725 6.71 made | Siz. ||, aaeS 7.09 1.02
INilia\(O) gate roererrneeeee 0.2 0.2 O.11 0.18 0.18 0.13 0.07 0.09
Cal) “Sisiadeia Geers 10.54 10.64 10.04 10.60 8.85 5.03 6.09 1.68
ISAO) oi a ots RCneE eee = = = = 0.01 = — —
IM (©) ts Gianna aaa 6.53 4.58 8.89 4.41 6.40 25.93 8.60 1.38
Nig OO} Seer eee 3.61 2.94 232 2.60 2.80 1.33 253 6.77
REO Matic fs ok ae 1.36 0.88 0.63 0.68 0.36 0.58 0.74 3.82
IP-(O}S ty oscectere ac roms 0.32 0.30 0.2 0.30 0.46 0.19 0.2 0.18
Or OR ae sisi 5's = = = == — = 0.13 —
1D RG eROo cna = = == = 0.15 — — =
CQMhodé och Renee = = = = 0.08 0.08 0.18 0.04
S\ o Bole Coe ope = = = == = trace 0.12 0.01
HeOm > socom 0.34 0.96 1.63 1.24 1.09 — = =
Cr ee 0.05 0.37 1.39 1.61 1.06 0.19 3.36 0.24 |
| 100.10 | 100.08 | 100.23 99.84 | 100.35 | 100.25 | 99.48 | 100.35

I. Olivine basalt (dolerite), Masatierra, Bahia Cumberland. Raoult anal.

II. Olivine basalt. Masatierra, Bahia Cumberland. Raoult anal.

III. Olivine basalt. Masatierra, Bahia Cumberland. Raoult anal.

IV. Aphyric basalt. Masatierra, Bahia Cumberland. Raoult anal.

V. Feldspar phyric basalt. Elevation 1 420 m. Masafuera. A.-M. Bystrom anal.
VI. Picrite basalt (Masafuerite, Oceanite) dike rock. Masafuera. N.Sahlbom anal.
VU. Iddingsite-basalt. Elevation 1000 m. Masafuera. N. Sahlbom anal.

VIII. Soda trachyte. Elevation 1200 m. Masafuera. N. Sahlbom anal.

Analyses I—IV have been made for A. Lacroix from specimens collec ed at Bahia Cumberland
by members of Dumont d’Urville’s expedition in 1854. These analyses were first published in
Lacroix’ paper: Constitution lithologique des iles volcaniques de la Polynésie Australe in 1927.

Analysis No. V is new, representing a sample of the abundant light grey feldspar basalts at
higher elevations on Masafuera.

Analyses VI—VIII are produced from my earlier paper on the geology of the Juan Fernandez
Islands (No. 12).

ee ee eee EEE
I II Ill 1V V VI Vil Vill |
SiON eens Aaa 46.21 46.68 47.83 48.60 47.36 43.34 45.27 63.36
iO eae a c 3.49 3.85 212 3.49 3.31 1.03 2.79 0.28
ATEO) tere, & aerate 16.03 16.75 1372 16.57 16.99 8.48 18.02 18.62
Hesse Severe 2.76 5.55 5-49 4.84 4.38 2.91 705 2.78
PEO ecw ate 8.86 7.34 6.90 7.29 8.32 10.99 7.38 1.02
Min@ie kero oe 0.23 O 24 O.11 0.18 0.18 0.13 0.07 0.09
CaO tee erect 10.53 10.78 10.33 10.82 g.OI 5.03 6.34 1.68
BaQOs x 2digasse = = = — 0.01 = — —
WigOm hi pe ere e 6.55 4.64 9.15 4.50 6.62 25.91 8.96 1.38
NiatOR es: Sac. Sac & 3.62 2.98 2.39 2.65 2.85 1633 2.64 6.76
KEO We ecersneycle 1.36 0.89 0.65 0.69 0:37 0.58 0.77 3.81
PLO eas ae aan oc 0.32 0.30 0.30 0.37 0.47 0.19 0.28 0.18
Cr On ecient = = = = = = 0.14 =
| Eons Cece Ben ER RES = = = = 0.15 — = ==
(CIE ees Oca ore = = = = 0.08 0.08 0.19 0.04
100.00 100.00 | 100.00 | 100.00 100.00 | 100.00 | 100.00 | 100.00

Analyses calculated as water free.

PERCY

QUENSEL

Molecular proportions (* 100).

————
I | II Ill 1V Vv VI VAL eevee |

| ‘S(O RS an oo oaaaoeT 76.94 FT TD 79.64 80.92 78.85 72.16 75-37 105.49

IO cospacn Gs obe 4.36 4.81 3.91 4.36 4.13 1.29 3.48 0.35

AUG Prana eeaaco wee W572 16.43 13572 16.25 16.67 8.32 17.68 18.27

Hes Ore circ chess 7s 3.47 3.44 BIOs 2.74 1.82 4.48 1.74
| © \arcrc cratic SOE 12.33 10.22 9.60 10.15 11.58 15.30 10.27 1.42 |
MiO™ .tieoe eee: 0.32 0.34 0.15 0.25 0.25 0.18 0.10 0.13 |

CaO Pe aanmuye eels 18.84 19.22 18.42 19.29 16.06 8.97 11.30 2.99
BAG) Bite e cre sie oness 6 = = = = 0.01 = = = |
IVI @) We wets ayene crenccens 16.24 11.51 22.69 11.16 16.17 64.26 2222 3:42)
Nias O Prac ota oie 5.84 4.81 3.85 4.27 4.60 2.14 4.26 10.90 |

aS Oe Saino aera 1.44 0.94 0.69 0.73 0.39 0.62 0.82 4.04
12 Oe pate cackencericnicka 0.22 0.21 ZI 0.26 0.33 0.13 0.28 (Ls) 4|
GH OSs Se ea ncanare = = — = — == 0.09 —_ |
Hitt AS eggs craves, «!s/2 — = = a 0.79 = z= — |
CNP ek oe cess sa = = == = 22 0.23 0.54 onan ||

ADDITIONAL

COMMENTS ON

THE

GEOLOGY OF

THE JUAN FERNANDEZ ISLANDS

Norms.
| | Il IT] LV V VI VII VIII
0) -«/teclernineerer oleae eee 0.14 0.40 4.25 0.66 | — — 4.49
C8 eC One — = | — | aon le 0.70
ONG. C’QpoctO TIC eee | 8.08 5.23 | 3.84 4.06 Ze 3.45 4.56 22.48
Di Oe eee | 18.14 25.22 | 20.19 22.39 24.12 Tie 52,33 vee)
Dil. one eee eae | 23.48 | 29.71 | 24.81 31.29 32.49 [547° | zoe Ife
NS oh Oe Oe | 6.76 — — = | = ==
ali notte Wo Gear 21.78 W7A39 19.32 16.00 7.43 6.65 siek |
IDS oils B HO ae rea ceER — 6.28 16.84 7.50 19.20 Buel 2OAOW Ne Seas
GS ee 10.46 ns == = == 53-07 | 1.55 | —
Dio) Soo ep CROre 0.74 0.71 0.71 0.87 I0r |= "0:44 7 /S.o:67, 0.45
‘l (2 ere 6.62 30 5.93 6.62 627 | 1.96) | =e oer lmmorca
Th) 2 re 4.01 8.03 7.97 7.02 634 N) A2T \etees7 | 2.78
OT tool eee } — _ —- — ey We Mes = 0.86
LN. ae ae ly bea Ave (hes hE -- Sa ei | 0.20 —
Gallo ctctecn ieee 56.39 | 60.30 | 49.24 | 61.99 59.44 | 30.14 | 58.50 91.93
WEINs. “oS eee AZ.On. || 30571 50.77 38.01 40.35 69.84 41.50 8.05
Quantitative system.
PUesse= 3): 4. -Camptonose OL Aabiane lO nARE ONG SirA 7-31
Pie G24 2 A dosodic or:ab:an 8.69: 41.92: 49.39
UTS 4 4 dosodic or:ab:an 7.86: 41.34: 50.80
HWA. ANTS ne Del dosodic Orsab any 7.03)2 38.78 2 54519
We MUGS O(a. persodic OG ab) san 3160) 7AT.03)455-28
Wile er 2 Al doferrous OR Mabiaanwy eA Aes37-2 302 5 e338
Wi 54) 4 dosodic OLA b any oO 7s Or5 La 52242
Wie leis: 2:4 Laurvikose OE Jabeanwe25-ORsO52o9 5 o-21

I II Ill IV V Vie Nant Vil
103.7 LIO/4,) || eTOuot lidices7/ |) ICL 69.8 99.7 236.3
—35.5 |—22.4 [=a18!o8 l=te5, |=17:3, ||—4toses gen ie
2.2 DI” Mimwetze8 DEG 23.4 8.0 23.4 40.9
43.6 41.2 | 51.9 40.5 47.0 80.6 55.0 18.9
25.4 Dg a | 24.3 28.3 22.6 8.7 14.9 6.7
9.8 8.2, | 6.0 WB 7.0 2 Gu i) Bie
0.58 0.66 0.47 0.70 0.48 O.11 0.2 0.35
5.88 6.83 5.16 6.40 | 5.80 12s 4.60 0.78
0.30 0.30 | 0.28 0.38 0.66 0.13 0.26 0.2
0.20 e316, |) o.15 0.15 0.08 0.22 0.16 0.27
0.50 0.40 0.58 0.40 0.48 0.77 0.53 0.40
O.1I 0.24 0.17 0.22 0.16 0.04 0.22 0.41
0.21 0.40 0.41 0.37 0.32 0.19 0.46 0.69

ios)

Io.

II.

22.

PERCY QUENSEL

Bibliography.

Tu. SuTcLiFFE. Crusoniana. Manchester 1843.

C. Bertero. Notice sur |’Histoire naturelle de l’ile Juan Fernandez. Annales des
Sciences Naturelles, Paris 1830, Tome XXI, p. 345.

A. CaLpcLEuGH. On the Geology of the Island of Juan Fernandez. Geol. Soc. of
London, Proceedings, Vol. 1, 1826—1833, p. 256. (Also published in Phil.
Mag. and Annals of Philosophy, Vol. IX, 1831, p. 220.)

P. PARKER Kinc. Narrative of the surveying voyages of H. M.S. Adventure and
Beagle. Vol. 1. Proceedings of the first expedition 1826—1430, p. 304.

J. Dumont p’URvILLE. Voyage au Pole Sud et dans 1’Océanie. Histoire du Voyage,
MS S412, 0p. 1rd).

J. Grance. Géologie, Minéralogie et Géographie physique du Voyage. Dumont
d’Urville, Voyage au Pole Sud et dans l’Océanie, 2° Partie, 1854, p.39.

L. Prater. Zur Kenntnis der Insel Juan Fernandez. Verh. der Gesellschaft ftir
Erdkunde zu Berlin. Band XXIII, 1896, p. 221.

A. Renarp. Report on the rock specimens collected on the Oceanic Islands
during the voyage of H. M.S. Challenger. Rocks of Juan Fernandez. Report
of the Challenger Expedition, Vol. II, 1889, No. 15. (Also published in
French under the title ‘Notice sur les roches de l’ile de Juan Fernandez’.
Bull. Acad. Belgique, tome 10, 1885, p. 569.)

L. Darapsky. Uber den Glockenstein von Juan Fernandez. Verh. des deutschen
wissenschaftlichen Vereins zu Santiago, 1886, Heft 3, p. 113.

R. Poutmann. Das Vorkommen und Bildung des sog. Glockensteins (Magnesit).
ibid: BandelL 13935 Heft 56.

—— Noticias preliminares sobre las condiciones jeogrdficas 1 jeoldjicas del Archi-
piélago. Publ. in F. Johow: Estudios sobre la Flora de las islas de Juan
Fernandez. Santiago de Chile 1896, p.1.

P. QuenseLt. Die Geologie der Juan Fernandezinseln. Bull. Geol. Inst. of Upsala,
Volaex, 1ron2(p..-2'52:.

T. HaGerMAN. Beitrige zur Geologie der Juan Fernandezinseln. The Natural
History of Juan Fernandez and Easter Island. Vol. I, 1924.

A. Lacrorx. La constitution lithologique des iles volcaniques de la Polynésie
Australe. Mém. Acad. des Sciences. Paris. Tome 59, 1927.

F. von Wo tr. Vulkanismus. Stuttgart. Teil I, 1913, p.290; Teil II, 1929, p.771, 805.

J. Brtccen. Fundamentos de la Geologia de Chile. Santiago de Chile 1950, p. 59,
B20. 382.

T. H. Tizarp, H. N. Mosetey, J..G. BucHANAN and J. Murray. Narrative of the
eruise ol HiaM: Ss. «Challenger: Vol; J, 1885, p. 818:

T. Surcuirre. The earthquake of Juan Fernandez as it occurred in the year
1835. London 18309.

—w— Sixteen years in Chile and Peru from 1822 to 1839. London 1841, p. 387
(with a sketch of the sub-marine explosion).

Cu. Darwin. Geological observations on the volcanic islands of South America,
visited during the voyage of H. M.S. Beagle. London 1876, p. 144.

T. Barru. Pacificite, an anemousite basalt. Journ. Washington Acad. of Sc. Vol.
ROS 1103.0, aps OO:

—— Mineralogical Petrography of the Pacific lavas. Am. J. of Sc. Vol. XXI,
NOS OO. LO:

L. Bowen. The evolution of igneous rocks. Princeton 1928, p. 163.

ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 87

A. JOHANNSEN. Petrography, III, p. 334.

E. DE MarGerig. La face de la Terre. III, p. 1359.

Battey Wittis and H.S. Wasuincron. San Felix and San Ambrosio, their geo-
logy and petrography. Bull. Geol. Soc. of America. Vol. 35, 1924, p. 365.

L. CuHuse and C. RicHarpson. Geology of Galapagos, Cocos and Faster Islands.
Bernice P. Bishop Museum, Honolulu. Bull. 100, 1933, p. 43, 47, 64.

C. Burret. Chemismus und provinciale Verhiiltnisse der jungeruptiven Gesteine
des pacifischen Oceans und seiner Umrandung. Schw. Min.-petrogr. Mitt.
Band 6," 10926, -p. 17-7.

Unfortunately an interesting paper by GorpEN A. Macponatp on the ‘Hawaiian

Petrographic Province’, published in the Bull. Geol. Soc. of America (60:2, 1949,

1588), in which comparisons with Juan Fernandez and other Central Pacific Islands

are discussed, has evaded my attention until this paper was already in print.

4. A Geographical Sketch of the Juan Fernandez Islands.

GXSKOTISBERG:

The Juan Fernandez Islands were discovered on the 22nd November, 1574,
by the Spanish navigator JUAN FERNANDEZ who called them Las Islas de Santa
Cecilia. They consist of two islands, distant from each other, Masatierra with its
satellite Santa Clara, and Masafuera. Masatierra lies 360 miles W of Valparaiso,
Masafuera 92 miles W of Masatierra. According to the charts the position of the
light in Cumberland Bay on Masatierra is 33°37'15” S. and 78°49'50” W., and
of the summit of Masafuera, 33°46’ S. and 80°46’ W.

The islands are of volcanic origin and considered to be late Tertiary. They
show no signs of recent activity, but a submarine eruption near Pta Bacalao in
Masatierra is reported by Sutcliffe to have occurred in 1835, and another E of
this island by Goll in 1839 (BRUGGEN pp. 326, 332). Sutcliffe (1, Plate p. 387)
published a drawing of the eruption; the landscape is a pure flight of fancy.
Some visitors have wanted to recognize a number of extinct craters. To this ques-
tion I shall return later. When Ulloa thought that he saw flames bursting from
the summit of Mt. Yunque, he certainly made a mistake.

No geographer has, as far as I know, visited the islands, but many notes on
their configuration and topography are found in the narratives of early naviga-
tors as well as in the official reports to the Oficina Hidrografica in Valparaiso
by the Commanders of surveying ships. Certain observations on the former dis-
tribution of the forests were referred to in an earlier paper (Skottsberg 3). Many
popular descriptions of the nature and life on Masatierra have appeared (see Bib-
liography), some also paying attention to Masafuera. The latest, by JORGE GUZ-
MAN PARADA, contains much useful material and will often be referred to here.

Comments on some earlier descriptions and maps of Masatierra.

The most interesting account of this island from the 18th century is found
in WALTER’s narrative of Captain (later Lord) ANSON’s voyage. The illustrations
are, even if not quite so accurate as the author thinks, vastly superior to the
contemporaneous ones in Ulloa’s work. Plate XIV is a prospect from E, including
Santa Clara (called Goat I.), the rock El Verdugo (Monkey Key) and part of
the north coast of Masatierra, seen under almost right angle and with the con-
spicuous mountains in correct position. Plate XV is a map, not bad in its main

6 — 537351 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. 1.

go C. SKOTTSBERG

features; of the mountains only Mt. Yunque appears. The names on the map are
Monkey Key, East Bay (Pto’ Frances), the Spout (a cascade not far from Pta
Bacalao), West Bay (Pto Ingles) and Sugarloaf Bay (Vaqueria). Woods cover the
east half; the treeless west half erroneously includes the still well wooded Villa-
gra valleys. Plate XVI is a Special of Cumberland Bay, of which Pl. XVII gives
a good view, and Pl. XVIII shows the Commodore's camp in the valley later
named in commemoration of his visit. Masatierra was the rendez-vous of Anson's
squadron and brought salvation to the remnants of the crews, of which the greater
part had fallen a victim to scorbut. The winter months of 1741 were spent here
and the ships refitted.

ANTONIO DE ULLOA’s narrative 1s accompanied by a panorama of the south
side of Masatierra showing Mt. Yunque, Mt. Piramide, Co Negro and Damajuana,
but other details cannot be identified. The map (Plate IV) is a rough sketch.
Three bays have names, Puerto del Ingles, Englishman’s harbour, very likely
named to commemorate Selkirk as the cave called ‘‘Robinson’s grotto” is found
here, Puerto Grande de Juan Fernandez (Cumberland Bay) and Puerto de Juan
Fernandez (Pto Frances). Some other (nameless) coves are indicated, e.g. Pangal.
Three rivers empty in the harbours.

I do not know the circumstances under which the survey by FRANCISCO
AMADOR DE AMAYA was made. It resulted in a map published in 1795 which
has formed the basis of the charts still in use, but it may not have been known
to THOMAS SUTCLIFFE, whose book “Crusoniana’”’ (1843) is accompanied by a
map with more details than the older ones; with regard to the coast line it
is inferior to Anson’s. Sutcliffe was Governor of the islands in the 1830's. There
are many names, but as I have not seen Amaya’s original map I do not know
which are new. Cumberland Bay is called Port of Juan Fernandez; West Bay,
Ulloa’s Puerto del Ingles, Selkirk Bay, and East Bay French Bay. Sugar-loaf Bay
(Vaquerfa) is called Sandal Bay, an interesting name; perhaps most of the sandal-
wood was obtained here in Sutcliffe’s time. West of this place we find Desola-
tion Bay, a well chosen name; now called Bahia Juanango. Herradura, undoubt-
edly an old Spanish name, is known now as Bahia del Padre; La Punta is Pta
de la Isla. The east cape, now Pta or Cabo Hueso de Ballena, is called Pta de
Juanango. On the south coast we find Caravajal (Carvajal), Loberia, Villagra,
Chamelo and Monkey I. These names are, however, misplaced. Sutcliffe’s Carva-
jal is Bahia Tierra Blanca, a name placed by him inland at the foot of the hills
(where it belongs), the two bights on both sides of “Loberia’’, B. Chupones and
B. Villagra, are nameless; the former is also called Tierras Amarillas on some
charts, a name used by Sutcliffe for a tract of land back of his Tierras Blancas.
Villagra is located east instead of west of Mt. Yunque, and Chamelo used for
the coast now called Playa Larga. The interior shows some topographical features,
a mountain range can be followed from east cape to beyond the misplaced Yunque,
and north of this is a short row of hills, corresponding to Cordon Central, which

* Abbreviations. B.=Bahia (bay), C.=Cordén (range, ridge), Co=Cerro (mountain), L.=
Loberia (sealing grounds), M.=Morro (small islet, rock), Pta= Punta (point, cape), Pto= Puerto
(port, harbour), Q. =Quebrada (narrow valley, gorge), V.= Valle (valley).

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS gI

separates “Anson's vale’ from “Lonsdale’’ (now Valle Colonial). A name not
found on any other map is “Kay's town’, the settlement in Cumberland Bay.
This name was given by Sutcliffe in commemoration of one JOHN Kay who,
through his technical skill, greatly furthered the textile industry in England and
whose biography appears in “Crusoniana’’. The Salsipuedes ridge and the ridge
between Pto Ingles and Vaqueria are marked, while the conspicuous crest unit-
ing Yunque and Salsipuedes has disappeared altogether. The topography of the
western section is poor, only Tres Puntas placed in correct position. The name
‘Puente’ is misplaced, but certainly refers to the elevated isthmus between Car-
vajal and Herradura.

Some later surveys and maps.

From time to time the Chilean Hydrographic Office despatched a vessel to
the islands as part of the work on a “‘derrotero’” for the entire coast of the Re-
public. The reports were published in the Anuario Hidrografico de la Marina de
Chile. Lopez (1876) mainly repeats older statements with regard to distances,
size of the islands, altitudes etc., VIEL (1878) concentrated his attention on
the possibilities of making Masatierra productive, VIDAL GORMAZ (1881) little
more than copied Lopez. The chart was not much improved. More information
on the nature of the coast, the serviceableness of the harbours and anchorages,
landmarks etc. are found in the compiled “‘Instrucciones nauticas of 1896. For
Cumberland Bay the original Spanish name Bahia San Juan Bautista is used,
and some other early names are preferred, Bahia del Este, B. del Oeste, Pan
de Azucar (Sugar-loaf, also Cerro Alto) and B. Pan de Azucar (Vaqueria), etc.
GUNTHER’s report of 1920 has little to add to the Instrucciones. A new chart
had now been published and is reproduced in a very small scale. The distances
between certain points indicated by Gunther agree rather well, with regard to
the east section of Masatierra, with those on my map, while considerable dif-
ference is noted in the length of the long, narrow western section, 12.96 km ac-
cording to Ginther, 10.25 on my map, so that the total length between Pta de
la Isla and Pta Hueso Ballena becomes 22.2 and 18.5 km, respectively.

From American, French and English sources the well-known editor of geo-
graphical and nautical works L. FRIEDERICHSEN of Hamburg compiled a new map
to accompany Ermel’s popular account of his visit to Masatierra (1889). The
central portion is much disfigured, but the general trend of the mountain ranges
more or less correct, the details, however, erroneous in many cases. Most of the
names used are Spanish. Some are still in use on the British and Chilean charts,
where, however, Punta is used for Cabo: C. del Padre, C. Tunguillar (Tinquillar),
C. Lemos, Morro Juanango, C. de los Negros (now also called Pta Suroeste),
B. de la Vaqueria, C. Salinas, Sal si puedes, C. San Carlos, C. Loberia, C. Bac-
alao, C. Pescadores, C. Frances, Corrales de Molina (a series of hanging gorges
E of Mt. Yunque), Morro Vifillo, Bahia Chupones, C. O'Higgins. Some of the
names on Friederichsen’s map are now forgotten: Bahia de la Fé (=B. Juanango),
El Palillo (west head of Pangal), C. Madurgo (W of the east cape, here called

g2 C. SKOTTSBERG

C. Guasabullena, a corruption of Hueso de Ballena), Morro Caletas (= El
Verdugo), C. Chupones (now Pta Larga), Bahia Aguabuena [now Tierra Blanca,
but modern charts have Pta Aguabuena between T. Blanca and Carvajal (Coq-
bajal of Friederichsen)|. The topography is much clearer and more correct than
in any of the earlier maps. With the exception of Cerro Alto and Yunque no
names of mountains have been put in.

R. POEHLMANN’s short description of the islands, with special reference to the
geology, serves as an introduction to Johow’s well-known work on the natural
history. Johow’s map of Masatierra, based on “recientes trabajos recopilados por
la Oficina Hidrografica en 1895”, gives a very unsatisfactory idea of the topo-
graphy.

Amador de Amaya’s map of 1795, with additions and corrections by the
British (no. 1383) and Chilean navies, remained the basis of all charts until 1917,
when I handed over my notes and sketches to the Oficina Hidrografica. From
1918 on several editions have appeared. Pta Suroeste replaces Friederichsen’s
Pta de los Negros, but the latter should be preferred because not this point but
Pta de la Isla is the south-west point of Masatierra. For Monkey Key El] Verdugo
is sanctioned, Cabo Chamelo is replaced by Los Chamelos, referring to the rocks
outside, Cabo Viudo by C. Norte, with the rock in front called Morro Viudo.
E of Co Tres Puntas Co Chumacera appears. More important is that, for the
first time, the valleys between Pto Frances and Pta Pescadores have been ind-
icated and named. On the Special of Cumberland Bay is the new name Cordon
de las Cabras for the ridge generally called C. Central. The British chart “with
corrections from the Chilean Gov. chart of 1921” reproduced by Quensel (2 p. 46)
shows the topography more distinctly than the former except of the east sec-
tion, where all the improvements have been omitted. A new name is Pta Mere-
daxia for Pta del Padre. The latest edition, revised up to March 1953, is iden-
tical, but for the topography a different technique has been used.

The names used by Guzman in his text do not always agree with those on
the map. He has taken up Herradura for Bahia del Padre, which is all right, but
when he called Bahia Juanango “Ensenada Pan de Azucar, cuyo nombre lo debe
a su islote Juanango’’ — the conical Morro — he made a mistake, because the
name Pan de Azucar belongs to Vaqueria and refers to Co Alto.

During our expedition I tried to sketch the distribution of the forest, using
the chart as a basis. The position of the boundaries was determined with help
of simultaneous aneroid and temperature readings; the same observations were
made at sea level before and after every excursion and the elevations calculated
from tables I had received from the late Professor AXEL HAMBERG. This method
does not, of course, give exact results, but it gives more reliable figures than
the altimeter. Our large series of photographs has been a great help. Neverthe-
less the need of a map, based on a real survey, was deeply felt, and when, in
1951, I was going to put my notes in shape for publication, I approached the
Chilean government through the Swedish Legation in Santiago and asked for
assistance from the Chilean Air Force. This was most generously granted. During
a flight on April 8, 1952, Masatierra was photographed; unfortunately it was rather

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 93

late in the season and much of the island was hidden by clouds." Though some
corrections could be made along the south coast and around Cumberland Bay,
the result was not what I had hoped for. A second attempt was planned, but had
to be given up, and the work was discontinued. Fortunately the Swedish engineer
Mr. BERTIL FRODIN, then a resident of Santiago and a most helpful channel during
my negotiations with the authorities, had been invited to join the first flight,
and his series of Kodachrome pictures, most of them taken from the plane, was
graciously put at my disposal. They have proved to be of very great help; the
reproductions here will, I dare say, testify to their high value. A comparison
with our photographs allowed me to identify practically every single forest patch
shown, and their size and shape was almost the same in 1952 as in 1917. With the
aid of all this material a new sketch map, reproduced here in reduced size, was drawn
(fig. 1). I want to emphasize that this map is a sketch only.

Main geographical features.

Masatierra can be inscribed in an obtuse-angled, isosceles triangle with the
hypotenuse (the distance from Pta Isla to Pta Hueso Ballena) 12.5 nautical miles
(23 km) long on the sea chart and the greatest width (from Pta Salinas to Los
Chamelos) 4.2 miles (7.8 km); circumference 34 miles (=53 km), area 93 sq. km.
These are the figures generally quoted, but others are also found: length 15.5,
width 3.75 miles (Lopez), 22 and 8 km (Ermel), 25 and 9 km (Branchi), etc. The
figures obtained from my map are: length (=hypotenuse) 18.5 km, width 7 km,
deed 5.2> sq: km.

Masatierra is a deeply eroded and very rugged mountain range (fig. 2) rising
abruptly from a submarine ridge running S—N and bordered by deep water; the
bathymetrical conditions will not be discussed here. There is hardly any level
land on the island worth speaking of. Where the soil is not covered by forest,
as on the barren seaward slopes of the valleys, on the coast escarpments and
on the precipitous ridges rising high above the continuous forest cover, hundreds
of lava beds overlying one another can be distinguished, varying in thickness
from a few m (in cases less than one) up to 20 or more (Quensel 2 p. 40). The
location of the main summit ridge and, as a consequence, the trend of the val-
leys, depends on the dip of the lava beds. From the east highland to Mt. Yunque
and from Pta San Carlos to Co Alto, the tilt is N to NE, above Pto Frances
14—18° (fig. 13), at Centinela and Pangal about 20°, between San Carlos and Pto
Ingles 12—13° (fig. 3), at Co Alto 20° or a little more. From the east end to in-

* One of Mr. Frédin’s photographs of Masatierra seen from the air in 2000 m altitude
was reproduced in the daily paper ‘Dagens Nyheter’. The explanation says: “. . . covered with
white clouds that later lifted, enabling us to map the islands accurately’. And in the text we
read: “We flew to and fro over Masatierra and took series of photographs which will be put
together to form maps in scale 1:15 000. We had the good luck to get the summits quite free
from clouds ...’’ This story is confirmed by Mr. Frédin’s kodachromes which show the central
and northern parts of Masatierra very clear. On the aerial map in 1: 28 500, submitted to
me by the Chilean Air Force, the island is, however, more or less covered with clouds, and
not one of the conspicuous mountains could be identified with certainty.

C. SKOTTSBERG

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VUAILVSVW

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 95

Fig. 2. Eastern half of Masatierra, seen from the air. — Photo B. Frédin 8/4, 195

Fig. 3. End of Cordén Salsipuedes close to Pta San Carlos, showing the dip of the lava beds.
At the foot of the ridge the Cemetery. — Photo C. Skottsberg 3/, 1917.

96 C. SKOTTSBERG

cluding the Yunque massif, the crest follows the south coast, an imposing bar-
ranca several hundred m high, with almost vertical gorges. As a rule the saddles
are impassable (figs. 11, 14, 31). From the saddle between Yunque and Piramide
(figs. 21, 22) the ridge turns NNW and in a shallow curve runs right across the
widest part of the island. The reason is that the beds, at least the middle and
upper ones, are horizontal; at the foot of Mt. Yunque a very insignificant tilt of
s—g° was observed in one place (fig. 7). The central part of the island receives
the greatest precipitation, and erosion has worked inland from two opposite direc-
tions, but the Cumberland valley system has been considerably more deepened
than the Villagra system. The ridge is 600 to 700m high in this section. A
narrow pass, Portezuelo de Villagra, often spoken of as “Selkirk’s lookout’’, forms
the only practicable passage between the two sides of the island. West of Vaqueria
the ridge reaches the north side of the island, turns SW and follows the coast,
rising to at least 500 m in the highest peaks, then getting lower and lower and
disappearing as we approach El Puente, flat and sandy and only 50 m above
sea level. The small peninsula forming the extreme west of Masatierra is crowned
by a hill at least twice as high. At Carvajal the beds appear to be horizontal
(fig. 47), but E of the isthmus they are tilted SE, and consequently all the valleys
trend toward the south coast. The dip is slight.

The change in position of the backbone, combined with its decreasing eleva-
tion, has a profound influence not only on the morphology, but, as a consequence
of the direction of the prevailing winds, also on the local climate and thereby on the
vegetation. Climatic dates will be found in my paper on the vegetation (3 pp. 812—
818); the common wind direction is SE to SW (together 78 %). Along the east
and central section the air currents are suddenly forced up over crests 500—
goo m high, cooled and condensed, and rain drenches the ridges (fig. 4). This is the
forest country, where the deep valleys are covered with verdure. The region
around Mt. Yunque may be shrouded in mist while all the country west enjoys
sunshine. Very often the lower cloud limit is knife-sharp (Skottsb. 3 fig. 2 on
p. 808). Fig. 4 shows clouds also over the West and on Santa Clara. But W of
Cerro Chumacera, where the main ridge forms the upper edge of the long north-
ern escarpment, the air does not hit a high, precipitous wall but rises gradually,
and the elevation is too modest to allow the rain-bringing clouds to gather ex-
cept now and then during the winter months. This is the barren, treeless, grass-
covered land.

Geology and morphology.

No extensive geological survey has been made in these islands. Our know-
ledge is mainly based on QUENSEL’s short visit in 1908, when he studied the
stratigraphy at a limited number of places and later gave an account of the
geology, petrography and mineralogical composition of the rocks, but his mater-
ial was too small to allow us to trace the different kinds of strata from one end
of the island to the other. For my own part I had no geological training, but
during our 1916—17 campaign I collected rock specimens in many places. They

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 97

Fig. 4. Masatierra and Santa Clara (right, behind the promontory) seen in SW from 3000 m alt.
— Photo B. Frodin §/, 1952.

were described by HAGERMAN. The joint material served Quensel for a renewed
study, enlarged to a discussion of the geotectonic connection between Juan Fer-
nandez and other volcanic islands of the East Pacific.

Even a casual visitor cannot fail to observe the difference in appearance
and colour between the lower brownish, yellowish and reddish slopes and profiles
and the higher, light to dark gray ridges; see the water colour sketch in Skottsb.
2, opposite p. 52. These two horizons can be followed from the east end to Tres
Puntas, possibly to Cerro Enrique, but no samples were brought from the ex-
treme western section with the exception of a few from Bahia del Padre. The
island is “in the main formed by a rather uniform series of basaltic lava beds,
only diverging in respect of coarser and finer grain or of a higher or lower
content of olivine’ (Quensel 2 p. 44). Rocks with a very high content of olivine
(picrite basalts) seem to be restricted to lower elevations; higher up more nor-
mal basalts, less rich in olivine, predominate, but between the two extremes there
is every transition. Of the lower lava beds, up to 200 m above sea level, “many
show a coarse-grained ophitic texture and may be classed as dolerites” (I.c. p. 45);
these lavas have been traced from Pto Frances to Tres Puntas. The dolerites are
very resistant and show, at least where observed by me, a columnar structure.
They form thresholds in some of the valleys. The most conspicuous ones were
met with in Vaqueria (fig. 5) — a piece of a column was figured by Hagerman
p. 28 — and on the south side of the island below Chumacera and Tres Puntas.
At Chumacera the bed is about 3 m thick. The pillars appear as long and narrow,

98 C. SKOTTSBERG

Fig. 5. Vaquerfa valley, Masatierra, showing bed of doleritic basalt. — After a photograph
bought in Valparaiso.

Fig. 6. Spheroidal weathering of basalt, Valle Colonial, Cumberland Bay. — Photo C. Skotts-
bere), 1917.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 99

Hard basalt
TTT Less hard basalt
20 Agglomerate beds

Dikes black

pout

2 ee ee oe

\
TITRE
WYYTIIABUDLLLD Y THTiTiy tee)
TEU n SD us
NULL °
ae x

s—i Til ——saen
TOR a ES, 3 Z :
BELL DEELEY OPAL EL GLI

Fig. 7. Diagrammatic sketch of a profile of the seaward base of Mt. Yunque. Height c. 350—400 m.

3-sided prisms. The dolerites have a very fresh appearance and are supposed to
represent intrusions between previously consolidated flows (Quensel 2 pp. 45, 47).

The lava beds at lower elevations in east and central Masatierra are inter-
bedded with agglomerate layers, formed by tuffaceous material as explained by
Quensel and illustrated by his fig. 11 (2 p. 54). The photograph was taken near
the entrance to Anson’s valley at approximately 50 m above sea level. In a
clayey ground mass of a deep brick-red colour hard blocks of various shapes
and sizes are embedded, showing spheroidal weathering; l.c. fig. 10 is a fine
example from the same bed. Another, from the floor of Valle Colonial near the
trail to Portezuelo, is seen in my fig. 6. The entire exposed surface weathers in
this fashion; the soft ground mass is washed out and carried into the sea and
the hard blocks left lying. The same kind of agglomerate is found also in other
valleys. Stratification is quite distinct in the profile illustrated by Quensel. His
opinion is that these beds are pyroclastic sediments of recent volcanic origin.
According to my notes the volcanic agglomerate was observed overlayered by hard,
gray basalt in Pto Frances and at the seaward base of Mt. Yunque (fig. 7). In
the profoundly eroded Cumberland valleys all the upper strata have been removed.

All through the island the lower horizons are traversed by vertical dikes of
hard lava striking approximately N—S. Nowhere are they better observed than
on the imposing perpendicular escarpment between Vaqueria and Juanango
bay, where the wall is ribbed with innumerable dikes which, thanks to their greater
hardness, project above the rim to form a serrated edge (figs. 8, Q).

The geology of Bahia del Padre was considered by POEHLMANN to be of
particular interest and importance. The lowermost bed at the entrance to the bay
was identified by him as an andesite representing a much older formation than
the overlaying basalts and tuffs and exposed only in this place, but Quensel came
to the conclusion that we have no reason to classify these beds as andesites of
an older formation (1 p. 266, 2 p. 56); they “have been subjected to alterations

in connection with thermal processes during some intermediate phase of volcanic
activity’.

SKOTTSBERG

Ce.

Ioo

, showing the numerous.

a and Juanango, Masatierra

Fig. 8. The escarpment between Vaquert

Photo C. Skottsberg 26/, 1908.

vertical dikes.

COT

A |

NISRERUMERIeSw en

and Juanango; comp.
arly.

a
gul

aqueri
do not run quite so re

arpment between V

; In nature they

8. Dikes black

oo
5:

Fig. 9. Diagrammatic sketch of part of the esc
fi

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS IOI

Fig. 10. El Puente seen from the shore in Bahfa del Padre, Masatierra. — Photo C. Skottsberg
*9/, 1917.

Fig. 10 shows the profile of the Puente. The foot of the cliff is more or
less hidden under talus material. A tuff bed overlayers a sequence of basalt
and agglomerate beds; one of these, rather distorted, can be followed righ across
the slope on Pl. 102: 1, Skottsb. 3. The tuff was described by Hagerman p. 26.
It resembles a coarse-grained sandstone and disintegrates easily. The Puente
(figs. 45, 46) and the adjacent slopes on both sides are covered with white, mob-
ile sand where small dunes and ripplemarks announce wind action, and the wind
carries the sand out into the bay. On the sand, standing more or less upright,
are numerous peculiar more or less tube-shaped concretions (see fig. 7 in Hager-
man’s paper). Hagerman p. 29 calls them sinter concretions formed by a number
of minerals in a cement of CaCO,: “Die wahrscheinliche Deutung dieser Phano-
mene ist wohl, dass mit Calciumkarbonat gesattigte thermale Gewasser uber eine
Vegetationsdecke geflossen sind, wobei Wurzeln etc. mit einer Kruste von oben
angegebener Zusammensetzung tiberzogen wurden.” This should bear witness of
late volcanic activity contemporaneous with the existence of a more humid cli-
mate than the present one, permitting a vegetation cover to thrive. Now there
are neither any hot springs saturated with lime nor is this region covered with
native plants; the Puente is a field of pure sand with large patches of weeds
along the edge (fig. 46). There are indications that the west part subsided in
geologically recent time, when Masatierra and Santa Clara hung together. With
greater elevation rains were more frequent and where the land is now barren, it
bore shrubs and trees.

102 C. SKOTTSBERG

More normal olivine basalts, less rich in olivine than the rocks character-
istic of the lower horizons, are “widespread up to the highest part of the island”
(Quensel 2 p. 49). They are dark gray and as a rule vesicular, scoriaceous or
slaggy, but they are hard, more resistant to denudation, and form the elevated
crests and crags all along the ridges. At intermediate horizons, approximately
between 400 and 500 m (Cordén Chifladores 400 m, Portezuelo 500 m), feldspar
basalts seem to predominate (Quensel |.c.). These lavas are ash gray, aphanitic and
aphyric in texture and have a tendency to develop a columnar structure. Rocks
of the same type were found near Pta Larga at less than 100 m in the form of
a dike, “which may signify a channel for the analogous lavas at higher levels’.
These beds are supposed to “represent a definite epoch of intrusion, intermediate
between the doleritic basalts and picrite basalts of the lower parts and the sco-
riaceous olivine basalts of the higher horizons’. Very likely the thick bed seen
on Pl. 97:2 in Skottsb. 3, about 450m above sea level, belongs to this type.
I admit that the photograph selected by Quensel to illustrate this formation (fig.
9, p. 51) has the same outward appearance, but the altitude is approximately
625 m, and as this place is out of reach — the climber cannot depend on the
shallow-rooted shrubs and ferns — no specimens were taken there; the samples I
brought came from 500 to 575 m, and whether or not the beds are intermediate
is impossible to tell, as no rocks from a higher elevation than 575 m have been
examined. The samples from this level are vesicular aphyric feldspar basalts. We
know nothing about the mineralogical composition of the rocks forming the
highest summit. Mt. Yunque rises about 350 m above Portezuelo de Villagra.

A geographical reconnaissance of Masatierra.

Pta Hueso de Ballena, where we shall start our circuit of the island, plunges
abruptly into the sea, forming an escarpment of perhaps 300 m. A dominant
feature of the coast is that talus deposits are insignificant or lacking, so that
the surf is able to undermine the wall and to excavate caves. Only in the coves
where a valley has been eroded down to sea level, a beach is found which leaves
a passage along the foot of the escarpment.

Between the east cape and the Frances valley the land rises to 500 m or
more. Some shallow quebradas, filled with forest, descend north toward the sea
but do not reach very far down (fig. 11).

Pto Frances does not deserve to be called a harbour; it offers no protection
even as an occasional anchorage (Instrucc. ndut. p. 226). It is a small, open cove
facing N and E, but with winds from other quarters landing is easy. The beach
consists of rounded stones and coarse shingles; here as elsewhere the surf re-
moves all minor particles. The lower slopes of the valley are very barren, the soil
is exposed or covered with patches of weeds, and the marks of running water
and the tracks of cattle are everywhere to be seen (fig. 12). Some little distance
from the shore and about 50m above sea level is a small shack. The streambed
occupies the entire narrow bottom of the broadly V-shaped valley to which sev-

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 103

Fig. 11. Most easterly part of Masatierra, seen from the air, looking east. The detached, coni-
cal rock is El Verdugo. Ridges a (centre) and 6. — Photo B. Frédin §/, 1952.

Fig. 12. Pto Frances seen from a spur about 350 m above the sea. — Photo C. Skottsberg
5S Fee OLO:
12

104 C. SKOTTSBERG

Fig. 13. Ridge a overlooking the E branch of Frances valley. — Photo C. Skottsberg ™*/, 1917.

Fig. 14. Pto Frances, W valley-branch. Left, c; right, ¢@ — Photo C. Skottsberg 17/4 1917.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I05

Fig. 15. Prospect of the E part of Masatierra, seen toward SE from an airplane at low eleva-

tion, showing the land from Pto Frances to Quebr. Pesca de los Viejos; between them, the

small and steep Q. Lapiz. The dotted line follows the crest of Cordén Chifladores. From left to
right ridges a, 6 and c. — Photo B. Frédin 8/, 1952.

Fig. 16. Continues fig. 15, to Pta Pescadores. From left to right Quebr. Lapiz, Pesca de los
Viejos, Laura and Piedra Agujereada; ridges a, 4, c, ad. — Photo B. Frédin 8/, 1952.

7 — 537351 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. Tf.

106 C. SKOTTSBERG

eral quebradas belong, filled with maqui-infested, grazed luma forest* lower down
but higher up with fine primeval stands. Some gullies cut deep into the ridge
and end in an impassable saddle; on the south side is a precipice. Figs. 13 and
14 show two of these gorges. Fig. 15 is a general view of the Frances system;
the letters a, 2 and c denote three conspicuous crests, easy to identify also on
figs. 11, 13, 14 and 16. The stream has some water also during the drier sum-
mer half year.

From Pto Frances the coast runs in an almost straight line to Pra Pescadores
(Fishermen’s point). The trail crosses three valleys, Pesca de los Viejos, Laura and
Piedra Agujereada, none of which has been eroded down to the sea (fig. 16), but
all have been cut back deep and widened to form a basin furrowed by numerous
small tributaries. To call these valleys “apenas unas grietas’’, as Guzman does
(p. 26), is not to do them justice.

Cordon de los Chifladores (Whistlers’ ridge, fig. 16) is, as most of the ridges
extending N—S, wide near the sea and narrows inland, in the steeper rise up
toward the crest approaching the knife-edge type. As everywhere along the north
side of the island the country near the coast is treeless. Possibly the forest never
went clear down to the cliff. This question was raised by Johow and commented
on by me (3 p. 800). I forgot then to mention Ermel’s theory that, as the islands
are so much younger than the mainland, a vegetation cover has not yet had time
to spread to the coast! There is no need for a discussion. On C. Chifladores the
first forest is met with about 300 m above the sea, covering the ridge another
50 m and then passing into the usual low scrub.

Near the seaward slope of the ridge is a dry crevice where a few stunted lumas
linger. We named it Q. del Lapiz (because I lost a pencil there).

O. de la Pesca de los Viejos (Old folks’ fishing place, fig. 17) had very
little water in the stream in December and not much more in April. Along the
outer slopes the inclination is gentle; the middle section is steeper, up to ae
Scattered trees are remnants of the once closed forest. The interior is well wooded
up to the main ridge. Not much maqui was seen in this valley.

O. de Laura (fig. 18). Near the sea the valley sides slope 23—24°, a little
farther inland about 30°. None of these valleys is deeply eroded. There was some
water in the stream in December, but nothing in April when the picture published
in Skottsb. 3 fig. 32 on p. 889 was taken.” Fig. 18 shows the valley in August
The outer section is desert-like, the soil naked or covered with weeds, but a few
solitary trees may be seen. The slope facing E is more barren than the opposite
slope. The interior is filled with good, thick-stemmed forest.

A high, well wooded crest, rising to at least 650 m, forms the background
of QO. de la Piedra Agujereada (fig. 19) which got its name from a rock pierced

* For some of the leading species the local names are used. Canelo, Drimys conferti-
folia, Chonta, Juania australis; Luma, Nothomyrcia fernandeztana (Masatierra) and A/yrceuge-
nia Schulzei (Masafuera); Maqui, Ardstote/ia magui (chilensis; a macal is a maqui grove); Naran-
jillo, Fagara mayu (Masatierra) and /. externa (Masafuera); Pangue, Gunnera peltata (Masatierra)
and G. Masafiuerae (Masafuera).

2 The text is incorrect. The photograph was taken by the author 17.4. 1917.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 107

_
\O
“NI

Fig. 17. Quebr. Pesca de los Viejos, looking S; ridge d. — Photo C. Skottsberg 17/,

Fig. 18. Quebr. Laura, looking N. — Photo K. Backstrém Aug. 1917.

108 C. SKOTTSBERG

+ py
td Lr Fe 4 ‘{ a e 4

Fig. 19. The interior of Quebr. Piedra Agujereada. — Photo K. Backstr6m Aug. 1917.

by a hole (aguja, needle). The exterior section (see Skottsb. 3 fig. 33 on p. 890)
has the same character as in the valleys mentioned, while the interior is filled with
fine forest covering the ridges on both sides and above 400 m very damp and rich
in tree-ferns. The vegetation cover acts as a sponge and only a minor part of
the precipitation will feed the stream, its lower course being dry during the summer;
in winter the water rushes down to the sea as a cascade, mentioned by Anson
(“The Spout’) and referred to in Instrucc. naut. p. 227, where it is said to run quite
dry at the end of the winter.

El Rabanal (rabano = Raphanus sativus, formerly abundant here) is very unlike
the other valleys with its wide, almost level floor; in Johow’s time it was densely
wooded, but in 1905 it was ravaged by fire, and the forest never came back. In
March 1917 the dry soil was covered with the dead stalks and innumerable young
rosettes of Silydum marianum (see Skottsb. 3 fig. 34 on p. 891). Pl. 86 (l.c.) shows
the same spot in August, when a vigorous new growth had sprung up. The shadow
across the S7/ydetum indicates the streambed. There are a few dying lumas, maqui
is plentiful in the quebradas, succeeded toward the interior by degraded luma-canelo
forest. Higher up are better stands (l.c. Pl. 94: 1).

Rising at an angle of about 85° Pta Bacalao forms the end of the Centinela
ridge. It got its name because the bacalao (‘‘stock-fish’’), the commonest and most
valuable fish in these waters (fig. 100), is very abundant here.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS

‘2161 "Jy, Saaqsnoys ‘9 oY — "[esurg 0} ddUVIVUD GYSRY “asp ayy jo
do} UO UONY]s ssafatIM ay, “AUO[OD 94} Wor Udas ‘vJaUNUID UOpIOD pur ovRorg VIq ‘OZ

“BI

TIO C. SKOTTSBERG

Fig. 21. View W from the summit of Centinela ridge, c. 780 m, toward Mt. Yunque and Mt.
Piramide (right), in front of Yunque Mt. Damajuana; below Piramide the crest between Quebr.
Minero and Pangal. — Photo C. Skottsberg ™/, 1917.

Cordon Centinela (Sentinel ridge). The broad northernmost part of this ridge
forms a small meseta, upon which, 320 m above the sea, stands the wireless station *
(fig. 20). The ridge, which is steep on both sides (45—55°), is very barren and shows
the stratification on the yellowish-gray lava beds very plainly. A zigzag trail, cut
in the rock, leads from Pangal to the station, where one has a splendid view of
Cumberland bay and the mountains behind (fig. 27). Following the ridge, which
gradually gets very narrow, we have the rare opportunity to walk right across the
island to the top of the main ridge and to look down on the south coast, almost 800 m
below our feet (fig. 21). The gradient is gentle all the way up, from 5° a little south
of the station to about 20° farther up. From about 300 m there is forest on both
sides, closing over the ridge a little higher up. A very dense scrub covers the crest
(Reameisoo22);

Pta Loberta (lobo, seal; place where sealing was practised in old times) and
Pta San Carlos (fig. 27) are the headlands of Cumberland bay. The distance
between them is about 2 km.

Bahia Cumberland or San Fuan Bautista is the only harbour in the islands
where large ships find good anchorage; see the charts and descriptions by Walter,
Sutcliffe, Lopez, Vidal Gormaz, Giinther etc. The bay is open toward N and NE;

* Acc. to Guzman p. 16 not used now.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS Tere

Fig. 22. Cumberland Bay from Pta Loberfa to Cordén Central, seen from the air in 1000 m
elevation. From left to right, Pangal, Cordén Escarpado, Quebr. Minero-Damajuana, La Dama-
juana, Valle Anson with El Yunque. — Photo B. Frédin 8/, 1952.

but winds from these quarters are uncommon also during the winter. The sudden
squalls coming down through the valleys with great force should be looked out for.
Four valleys end in the bay, from E to W Pangal, Minero-Damajuana, Anson
and Colonial. The scenery round the bay is very impressive, a semicircle of bold
mountains, green gorges and gray precipices (figs. 22, 27). Many of the names quoted
by Guzman are unknown to me, for inst. Cord6n de la Falda Larga (‘‘falda’’ in the
sense of “long skirt’), perhaps = the back wall of Q. del Minero, Picacho con la
Piedra con Letras (?Cerro Pirdmide), Cerro de los Mufioces, Picacho de la Mona
(= she-monkey) and Cerro el Tope (“‘top’’). Pico Central, a long established name,
is not mentioned by Guzman.

El Pangal. The entrance to this picturesque gorge is crossed by the trail to
Centinela, but it is also easy to land on the stony beach. The main branch is a
blind alley and the only true canyon on Masatierra, recalling on a small scale the
magnificent gorges of Masafuera. The U-shaped gorge ends in a high wall, luxuriant
with verdure, through which a small waterfall leaps down. The altitude of the val-
ley bottom at the foot of the fall is only about 200 m. Pls. 78, 87 and 98 (Skottsb. 3)
give a good idea of the vegetation with its stately tree-ferns and giant pangues
(Gunnera peltata) which gave its name to this valley. No wonder that WALPOLE
found that the “rhubarb” grew so luxuriantly on Masatierra, that it was too coarse
to be good! Above the waterfall the valley widens and is filled with primeval forest
(lc. Pl. 89:1). A branch with a patch of forest comes down from the Centinela

ridge (fig. 23).

I12 C. SKOTTSBERG

Fig. 23. Centinela ridge and Pangal, seen from the slope of Salsipuedes above the Cemetery. —
Photo B. Frédin 29/, 1952.

Cordon Escarpado (= steep ridge) with its Picacho, 365 m high (fig. 22), sepa-
rates Pangal from the next valley.

Q. del Minero (Miner’s v.) comes down to the water rather steeply (figs.
22, 24). We did not survey this valley, but to judge from the colours on the
Kodachrome film only the higher slopes have native luma forest, the lower being
covered by maqui. There were no plantations here in 1916—17. A steep spur
separates Q. Minero from

Q. de la Damajuana (figs. 25, 26). The lower slopes have been cleared, but
between 150 and 200 m elevation a dense macal fills the bottom, followed by a
mixed luma-maqui forest and finally a belt of native wood.

La Damajuana (The Demijohn, figs. 25, 26) is a very characteristic landmark.
The cone crowns a short, high and narrow spur, continued toward the sea by a
long ridge, which is barren on the west side. This ridge can be followed up to
the base of the cone, alt. c. 430 m, and from there round the bend into the valley
and up to c. 530 m. The gradient is steep, nowhere under 35°. There are ledges
of harder rock, on both sides with a small waterfall, the first at 230 m when going
up the valley. The sides of the cone are precipitous, the height 2338 ft (739 m)
according to the English chart, 712 m on the latest Chilean map, but only 570 m
according to Branchi; this figure is too small. As far as I know this mountain has
not been ascended. An attempt should be made from the south.

Valle de Anson drains the loftiest part of Masatierra and is watered by two

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 113

Fig. 24. Quebr. del Minero. — Photo C. Skottsberg *5/,. 1916.

permanent streams. Bounded by Damajuana on the east and by Cordon Central on
the west it is dominated by the square Yunque massif (figs. 22, 31). Due to the
scant water supply in the side gullies near the sea, the distal part of the valley floor
inclines steeply, whereas the middle section has been levelled, being almost hori-
zontal at the small clearing known as Plazoleta (or Plazuela) del Yunque (‘small
square’), situated about 220 m above the sea (Skottsb. 3 Pl. 89: 2), but from here
the gradient gets very steep. It is evident from the pictures in Walter's narrative
that quite some clearing was done by Anson's party, very likely as far up as to
Plazoleta, and there is little native forest below this point. In 1916—17 the trail
went through an extensive macal, and the lower slopes near the sea were barren.
Very few people lived in this valley (fig. 27).* The forests of the interior were in
good condition and the upper montane type luxuriated below the saddle between
Damajuana and Yunque from 400 to 600 m.

El Yunque (The Anvil) presents, from all directions, the same venerable
appearance (figs. 21, 22, 25, 31, 51). The walls are everywhere steep with gradients

« Plazoleta is where, in 1930, HUGO WEBER went to live all alone asa modern Robinson,
as told in his interesting book. He built a hut, cleared the ground, made a garden and raised
chickens. He married in 1932, built a more substantial home and extended his cultivations. In
1942 he left the island and settled on the mainland. The little farm is still his property but without
a tenant, and I am told by Dr. GUILLERMO KUSCHEL that the place was overgrown with Aubus
ulmifolius when he visited the island a few years ago.

mie? C. SKOTTSBERG

Fig. 25. Damajuana and Yunque from the trail to Pangal, looking SW. — Photo C. Skottsberg
/4 1917.

from 55 to 70, in places almost perpendicular. The exact height is not known;
the figures vary from 1700 ft (§37 m, Walpole) to 983 m (Viel); Branchi has 805 m,
the English chart 3005 ft (913 m), the new Chilean 915 m, Instrucc. ndut. 927 m.
The first ascent was made in 1795, a second shortly after, the third in 1923, after
which the attempt has been repeated with success a few times (Guzman p. 30).
Only the route from the Damajuana-Yunque saddle seems to lead to the summit.
The figure 838 m (doubtless too high) on Friederichsen’s map corresponds to a small
pinnacle set on the saddle, the ‘““Camote’’ (= sweet potato, bulb). The ascent, which
is difficult and dangerous, was described by TENZ, who was the first to give us
any information on the topography and plant life of the summit; see Skottsb. 3
pp. 897—898.

Cordon Central (figs. 22, 27, 31, 99). From the broad gable of Mt. Piramide,
E of its centre, this sharp-edged ridge runs down to the coast, separating the Anson
and Colonial valleys and widening to a fan-shaped, barren and sandy front, slop-
ing down to the stony beach. Prco Central, c. 570 m, marks the end of the harder
basalts belonging to the upper horizons; l.c. Pl. 89: 2, Johow Pl. IV.

Valle Colonial (V. del Polvorin of Guzman; p. = powder-house), the seat of
the village, is the only valley on Masatierra that approaches maturity. The floor
rises gently to the foot of Mt. Piramide, a distance of about 2 km, and is watered

A GEOGRAPHICAL SKETCH OF THE JUAN FERN

JEZ ISLANDS Il5

Fig. 26. Mt. Damajuana seen from Anson’s valley. In the foreground a section through an
agglomerate bed. — Photo C. Skottsberg 75/12 1916.

116 C. SKOTTSBERG

Fig. 27. Valle Colonial seen from the Centinela ridge, c. 350 m above sea level. Left, Cordén
Central; above, Portezuelo de Villagra and Cordon Salsipuedes, ending in the vertical escarpment
of Pta San Carlos, behind which, on the west side of Pto Ingles, towers Cerro Alto. — Photo
Cz Skottsberg 24),21917.

by two permanent streams, which receive several small tributaries from the
surrounding gullies. The valley bottom was cleared of its native forest centuries
ago; already 80 years ago practically nothing was left below 250 m. Macales and
maqui-luma stands fill the interior, and only farther up the gullies we find better
forest. Material washed down from the sides have contributed to build the wide,
stony and sandy beach, the largest piece of level ground on this island (figs. 3,
20, 27). I cannot remember having heard of any names for the streams. Guzman
mentions 3 streams emptying in Cumberland Bay, Arroyo del Hospital, A. de la
Turbia (turbio = turbid) and A. de la Reina; the first is, I suppose, where Anson
had his hospital.

El Piramide (figs. 21, 29, 31), separated from Mt. Yunque by a narrow saddle
(W Portezuelo del Yunque), towers above the colony. Johow’s Pl. V dates from von
Rodt's reign and shows the scenery when no village existed. The figure 809 m
on Friederichsen’s map stands for the summit of Mt. Piramide. From both N and
S the ascent is rather steep, 40—50 the first stretch, but gets more gentle higher
up, 25—20°; toward the valleys on both sides the slopes are precipitous (Skottsb.
Bc Resor

Portesuelo de Villagra (fig. 28). No visitor, even if he only has a single day
at his disposal, fails to visit this famous spot with the Selkirk memorial tablet. The

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS Pry

ce tg

Fig. 28. Portezuelo de Villagra, seen from above the settlement in Valle Colonial. — Photo
C. Skottsberg *°/,, 1916.

altitude of the pass is generally stated to be 550 m; an average of 20 aneroid read-
ings gave 593 m, which I believe is too much; about 575 ought to be approximately
correct. The north “door-post’ rises with an angle of 80° (Quensel 2 fig. 9 on
p. 51), the south is formed by the slope of Mt. Piramide; see fig. 29, a transverse
profile with a gradient on either side of about 70°. The trail across has, in some
places, been worn down in the red forest soil and has been deeply eroded by
running water. From the saddle one has a grand view of the south side of Masa-
tierra from Mt. Yunque to Pta O'Higgins and Santa Clara (figs. 49, 50).

A volcanic island, especially if considered to be of late Tertiary origin, often
has craters or shows other signs of recent activity, and it is not surprising that
visitors to Masatierra have tried to locate old centres of eruption. Walpole writes
p. 93: “We skirted up the western side, which shows three semicircular craters,
whose sides toward the sea are broken down, thus forming bays within their basins.
Of these, Cumberland Bay is the central...’ The other two were, I presume, Pto
Frances and Pto Ingles (if not Bahia del Padre). And Quensel (1 p. 256), referring
to Cumberland Bay, speaks of “die lockeren Tuffmassen, die noch einen alten
Kraterboden bedecken ...” and p. 257: ‘‘“Machtige, oft rot gefarbte Tuffablagerun-
gen fillen den Talboden, was darauf hindeutet, dass es sich nicht um ein Erosionstal,

118 C. SKOTTSBERG

Fig. 29. South side of Portezuelo with the Selkirk memorial tablet, seen from the pass. —
Photo C. Skottsberg #5/,. 1916.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 119

sondern um eine teilweise erhaltene Kraterbildung handelt.’’ Finally, “Ob das west-
lich von dem Portezuelo sich 6ffnende cirkusformige Tal auch eine Kraterbildung
darstellt, lasst sich nicht entscheiden. In einer vorlaufigen Mitteilung habe ich diese
MOglichkeit angedeutet, und vieles scheint mir darauf hinzudeuten, dass urspriinglich
ein Doppelkrater hier gestanden hat, wobei gerade der enge Riicken des Portezuelo
die Scheidemauer bildet.’’ This was written after our visit in August 1908, and at
that time I shared my companion’s opinion (1 p. 136). During my later expedition,
when I came to know most of the island better, I arrived at a different conclusion
(see e.g. 2 p. 57). The geological structure is the same all the way from Pto Frances
to Bahia Juanango, we find the same agglomerate beds more or less well exposed
in the valley bottoms, but nobody would be inclined to deny that they are typical
erosion valleys where denudation, thanks to the scant supply of running water, is
very slow; in many cases abrasion has worked faster than erosion. In Pto Frances
the stream has barely managed to lower its bed down to sea level near the coast.
In Pto Ingles, where the surrounding ridges are very high, the distal part of the
valley has been widened and levelled, and quite some alluvial soil has been formed.
The same procedure is, I think, responsible for the formation of the Cumberland
valley system; there is nothing that speaks in favour of a crater theory or against
its origin as the result of erosion. The streams drain the wettest part of the island,
where erosion is greater and faster and accumulation a factor of some consequence.
As I said, my first impression was that the Colonial valley was the old crater and
its wall formed by Cordon Central, Mt. Piramide and the Salsipuedes ridge, and
I think this also was what glimmered in Quensel’s mind. However, the thick, strati-
fied deposits of brick-red, deeply weathered tuff with “‘Bruchstiicke von Olivin-
krystallen, Lapilli-artige Lavabruchsticke, Erzkorner und Glas’ also occur in
Anson's valley, separated from Valle Colonial, the supposed crater, by the over
500 m high Cordon Central which runs down to the shore of the bay. I mentioned
above that my photograph of the section through an agglomerate bed published
by Quensel was taken in Anson's valley. Either this is another broken-down crater,
or the deposits are ejections from the “Colonial” volcano. This would make us
postulate that the Anson valley had been excavated even to a lower level than the
present one when the volcano was active, and this seems questionable. In his
second paper Quensel quotes his earlier discussion (p. 53) and adds: “‘It is over
40 years since I visited the locality and naturally I cannot now rely on any personal
recollection”, and “But the composition of the formation, as well as my notes from
the field, offer indications that the tuffaceous material of Bahia Cumberland also
represents pyroclastic sediments of recent volcanic origin”. He refers to the sub-
marine eruption 1835 off Pta Bacalao. I willingly admit that I lack the necessary
training to get to the bottom with a geological problem, but my observation near the
foot of Mt. Yunque (see above p. 99) suggests that the agglomerate bed underlying
the basalt is of the same nature as the formation in Cumberland Bay. With regard
to Villagra, there is no semicircular valley corresponding to the opposite one, for
only on the north and east sides are ridges resembling a crater wall. Until a geo-
logist-vulcanologist has had an occasion to study the island it is better to leave the
question of the nature of Cumberland Bay open.

120 C. SKOTTSBERG

patente amen

Fig. 30. Cordon Salsipuedes from Cumberland Bay, with the still wooded quebradas nos. 3 to
5 (counted from Pta San Carlos); beyond, the deep Quebr. Gutierrez. — Photo B. Frédin 29/, 1952.

Fig. 31. Cumberland Bay in SSE, seen from the air. From left to right Quebr. Minero-Dama-
juana, Damajuana, Portezuelo del Yunque, Valle Anson, El Yunque, El Piramide with Cord6én
and Pico Central, Salsipuedes with Pta San Carlos. — Photo B. Frodin 8/, 1952.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 121]

Fig. 32. From the trail across the Salsipuedes highland, looking SE. Photo C. Skottsberg 24/, 1917.

Cordon Salsipuedes forms the western boundary of Cumberland Bay and runs
from the main range to Pra San Carlos (figs. 3, 27, 31), along the east side of the
Salsipuedes highland between the bay and Pto Ingles. Whether the name, which
means “get out, if you can’’, was originally applied to the coastal escarpment, 380 m
high on Friederichsen’s map, I cannot tell. A well-worn zigzag trail, also used
as a bridle-path, unites the colony with Pto Ingles and crosses the ridge at about
410 m. From here it is not difficult to follow the ridge up to over 700 m; who
attempts to continue until the rib abuts on the backbone takes his life in his hands,
for the ridge is a knife-edge, studded with crags. Already at about 600 m it is only
I—2 m wide, and the rise, quite gentle farther down, increases to 2 —30. As
we climb, scrub, brushwood and forest succeed each other on the slopes.

The declivity facing the Colony is sculptured by a series of shallow gullies, of
which the two nearest to the coast are treeless, whereas the others are wooded.
A comparison between fig. 30 from 1952 and Skottsb. 3 Pl. 90: 1 from 1916 makes
it clear that the forest patches have undergone no perceptible change in size. The
floor of the fourth quebrada (counted from Pta San Carlos) slopes 30°; the forest
is of an open, degraded luma-canelo type with much maqui. There is no water in
these gullies. The gullies of the main ridge, Q. Gutierrez and Q. del Monte
Maderugo, were thickly wooded in 1917 (monte maderugo = forest full of timber).

The trail to Pto Ingles ascends the buttress between the gullies no. 2 and 3,
descends on an undulating slope (fig. 32), crosses Loma de los Muiioces and descends

8 — 537351 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I.

122 C. SKOTTSBERG

rather abruptly into V. Ingles. The quebradas coming down from the main ridge
are filled with a remarkable Dicksonza forest near the range — note Q. Helechos (“Fern
gully’) — followed by luma groves and macal; I refer to my description, 3 p. 909.
Fog is a very important climatic factor here.

Pto Ingles offers no protection and is not used as a harbour (fig. 33). As a rule
there is a heavy swell and the boulders on the beach are shifted to and fro making
landing uncomfortable, perhaps dangerous. Close to the east headland a rock pro-
jects, separating a miniature cove from the main bay. The beach is a steep wall
of boulders, but with a tolerably calm sea landing is easy at the foot of the rock,
which is pierced by a tunnel, “una roca agujereada sobre una playa de piedras
grandes’, as this place is described in Instrucc. naut. p. 229; referring to Selkirk,
Guzman calls the tunnel “la portada del Solitario” (p. 23). Through the tunnel or,
at high tide, across the rock, we gain the bay, and immediately to the left, about
5 m above sea level (Branchi), is the famous “Robinson’s cave’, a favourite goal
for visiting tourists (fig. 34). It is hardly probable that the cave, described in some
detail by Guzman l.c., served the recluse as his permanent abode.

Fig. 35 is a general view of the valley seen from an airplane, but this picture
does not show the extent of the beach flat and of the wide, gently sloping valley
floor. The stream has water at all seasons. A ridge extending halfway down the
valley divides it in two; the east branch comes from a crescent-shaped saddle, equally
conspicuous from both sides of the island (figs. 35, 36) and reported to be impassable.
The dividing cordén can be followed along the crest up to about 550 m, where it
gets so narrow that further advance becomes too hazardous (see Skottsb. 3 PI.
go: 2). All the low land in the valley (fig. 36) has been cleared by fire and the forest
replaced by extensive weed fields, but in the branch valleys and side gullies is some
good forest, where a few chonta palms have been spared (Skottsb. 3 Pl. 88). There
is much naranjillo, but little maqui. The continuous cover of herbs and grasses
testifies that erosion is slight, and so is the inclination of the cleared valley floor.
It would perhaps be possible to reforest this valley with luma and other native trees.
At present (or at least in 1917) it is grazed; a tropilla is seen in fig. 36.

Cerro Alto boldly terminates the dividing ridge between Pto Ingles and Vaqueria
(figs. 37, 38). Possibly this ridge gives access to the summit which is about 600 m
high; an older figure says 627 (Friederichsen, Johow). All other sides are almost
or quite perpendicular. Patches of forest are seen on the flanks of the cone.

Bahia de la Vaqueria serves, as the name suggests, as a cattle ranch. The
cove is useless as an anchorage, but landing is easy enough with a calm sea. There
is no level beach; the stream, which is permanent, gropes its way between a wall
of boulders of all sizes. Fig. 39 is a general view of the valley seen from the air.
The outer part is grass-land with scattered trees on the slopes, closing to form groves
higher up, the interior is densely wooded (Skottsb. 3 fig. 35 on p. 894). The ani-
mals in Vaqueria have been left to run wild, and the visitor should look out for
the bulls.

As far as I could see, the geological structure is the same as in Cumberland
Bay, with the same red volcanic agglomerate, and it was during our visit to Vaqueria
that I began to doubt the crater character of the former. Halfway up the valley

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I

oA
<

-
3

Fig. 33. Pto Ingles with the foot of Cerro Alto (note tilt of the strata). Mrs. S. seated on an
old Spanish gun. — Photo C. Skottsberg ?°/, 1917.

Fig. 34. The cave (‘‘Robinson’s grotto”) in Pto Ingles. — Photo P. Quensel *5/, 1908.

124 C. SKOTTSBERG

Fig. 35. The surroundings of Pto Ingles, seen from the air; note the crescent-shaped saddle
overlooking the south coast of the island. — Photo B. Frédin 8/, 1952.

Fig. 36. The gently sloping floor of Valle Ingles with the main range in the background; right,
the dividing ridge where, in 1908, the last living Saztalwm grew. — Photo K. Backstr6m 1917.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I2

Fig. 37. Cerro Alto, seen from Cordon Salsipuedes. The small eminence above the escarpment
is the same shown at the extreme right in fig. 38. — Photo C. Skottsberg */12 1916.

Fig. 38. Cerro Alto seen from a point on the crest of the central ridge in Valle Ingles, c. 400 m
above sea level, looking N. — Photo C. Skottsberg 19/, 1917.

126 C. SKOTTSBERG

Fig. 39. The Vaquerfa cove and valley, seen from the air. Left, Cerro Alto, nght, ridge
between Vaqueria and Juanango; the top of Cerro Chumacera visible behind. —
Photo B. Frédin 8/, 1952.

6 7 8 9

1 Co Agudo; 2 El Piramide; 3 El Yunque; 4 Co Chumacera; 5 Co Tres Puntas; 6 V. Villagra;
7 Villa Alemana; 8 M. Juanango; 9 V. Juanango.

Fig. 40. Bahfa Juanango with Pta Negros and Morro Juanango, seen from the air looking SE.
— Photo B. Frédin 8/, 1952.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 127

we meet the conspicuous dolerite bed mentioned above (p. 97 and fig. 5). A pas-
sage across the steep rocky and scrub-covered ridge leads us down into the
Juanango valley.

The spectacular escarpment between Vaqueria and Pta Negros was described
and illustrated above (p. 99 and fig. 8). Near the entrance to Vaqueria is a low rock
and a little farther west, off Pta Norte, another called Morro del Viudo (Widower's
rock). These rocks rest on a submarine abrasion terrace, clearly distinguishable at
low tide. The coast wall projects west in a long, narrow and curved spur, not unlike
a saw-blade, Pra de los Negros, and SE of this lies Morro Fuanango (fig. 40).

Bahia del Fuanango is a wide bight, protected against winds from N and E,
but otherwise open. Seen from some distance out to sea it looks forbidding, and
the name “Desolation Bay” seems well chosen, but on a closer view two green
quebradas come in sight, V2//a Alemana (Germantown), accessible with difficulty,
and Q. Juanango.

QO. del Fuanango. The beach in the little cove is of the same kind as in
Vaqueria, a low escarpment and large, angular stone-blocks, but with a suitable
wind landing is easy. My diary calls these blocks ‘‘conglomerate’’, without much
doubt identical with the agglomerate found in Vaqueria. The valley is full of weeds
near the sea, but as it isn’t grazed native grasses are abundant farther in. At about
200 m above the sea are the first forest patches, and a little higher up, at the small
waterfall, is good forest. The threshold is, I suppose, formed by the same dolerite
bed as in Vaqueria, but I did not bring any specimens.

From Fuanango to Bahia del Padre the coast escarpment trends SW in wide,
slightly concave curves to Pra Lemos and Pta Tunquillar and thence to Pra
Meredaxia, the east head of B. del Padre. Figs. 41 and 42 make further descrip-
tions superfluous. Two conspicuous mountains tower high above the coast-line,
Chumacera and Tres Puntas, overlooking both sides of the island; the latter with its
three peaks ought to be a fitting goal for expert climbers (fig. 43). On the slope
of Chumacera are found the most westerly forest patches on Masatierra; from here
the country is treeless.

Bahia del Padre (Parson’s Bay) got its name from the configuration of the
rock W of the entrance. The old Spanish name B. Herradura (Horseshoe B.) is still
used by some authors. The diameter is about 250 m. It is a convenient harbour
for small boats, but the entrance, guarded by rocks, is narrow (fig. 44). Landing
on the beach of sand and shingles (Skottsb. 3 Pl. 102: 1) is comfortable with a
moderate swell, but may be difficult. The cove is a natural amphitheatre, but we
do not find very much of the “risuefias representantes del mundo vegetal’ praised
by Guzman (p. 24), except a patch of salt-meadow (Sa/scornia) along the beach,
because the flora is poor and mainly consists of weeds, among which the gilly-
flowers are conspicuous. Nowhere is the climate drier. // Puente was described and
illustrated above (p. 101, fig. 10); the flat, sandy surface is seen on figs. 45 and 46,
the former showing the wind-polished tuff beds.

Looking at the maps and photographs one is struck by the peculiar appearance
of B. Padre. It is evident that it is no valley, no result of erosion. Branchi wrote
(p. 168): ‘‘Un crater muy pronunciado puede suponerse en la Bahia del Padre’, and

128 C. SKOTTSBERG

Figs. 41—42. North coast of western Masatierra from B. Juanango to Pta Lemos, seen from
Quebr. Juanango. Cerros Chumacera (note patches of luma), Tres Puntas and Enrique. — Photo
C. Skottsberg °/, 1917.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 129

ipseAeeeGerromlres Puntas.—— Photo Hans Frey.

130 C. SKOTTSBERG

Fig. 44. The entrance to Bahia del Padre. — Photo C. Skottsberg 15/, 1917.

I am tempted to endorse his opinion. Both Quensel and Hagerman came to the
conclusion that the rocks have been subject to post-volcanic thermal processes, and
Hagerman, after a description of the palagonite tuff from Puente, writes: “Stellt
man diese verschiedenen Bildungen aus der Nahe der Padrebucht zusammen, so
gelangt man zu der Auffassung, dass dieses Gebiet frische Spuren vulkanischer
Tatigkeit aufweist’’ (p. 26).

On the naked sand we found many living beetles (no reference is made to them
in vol. II) and empty shells of four species of landshells, /ernzandesta tryoni Pils-
bry, Succinea fernandi Reeve (also in sand at Tierra Blanca), S. ¢er¢a Odhner and
S. semiglobosa Pfeiff. (also in sand on Santa Clara). These delicate creatures of the
humid forests are entirely unfamiliar to these dry and barren surroundings, and it
is difficult to account for the presence of these shells here as well as at Tierra
Blanca and on Santa Clara. Are they a testimony of a more humid climate per-
mitting some kind of brushwood to exist, a period during which the concretions
mentioned above (p. 101) were formed? Or did dwarf trees such as Dendroserts
litoralis, Rea pruinata and Chenopodium Sanctae Clarae, on which landshells lived,
grow here in historical time but before goats were introduced? Certain facts do speak
in favour of this theory. A few specimens of Rea and Dexdroserzs still occur on the
south coast of Masatierra, particularly on M. Vinillo and on M. Juanango, where the
goats cannot get them. What did this now barren country look like when the islands

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 131

Fig. 45. View from Puente, looking S. Wind-eroded tuff beds and sand. Behind, Pta O’ Higgins
with its morro, in the background Santa Clara. — Photo K. Backstrém }/, 1917.

ees

itn

Fig. 46. Mobile sand on Puente, in the foreground a dense growth of weeds, mainly Chenofo-
dium multifidum. C. S. as measure. — Photo K. Backstr6ém 15/, 1917.

132 Cc. SKOTTSBERG

were discovered? Not as to-day I am sure, but more or less like Morro del Spartan
at Santa Clara, as already suggested by Johow p. 261. I would think that 80, in
some valleys 90 per cent of the soil is now occupied by introduced weeds. We
have no reason to believe that there was a desert when the immigration of aliens
began.

It is to be regretted that we did not study how the shells found did occur,
if only on the surface or also deeper down embedded in the sand. And we have
to find out if landshells live on the plants mentioned above.

With Pra de la Isla we reach the end of the north coast. Time did not
permit us to visit the small peninsula W of El Puente, and we shall now pro-
ceed along the south side of Masatierra.

Pta O Higgins, watched by a nameless morro (small skerry), and the inhosp-
itable coast cliffs of Bahia Carvajal are seen in fig. 47. With a NW wind
boats engaged in langost fishing at Santa Clara find shelter here. From P%a
Aguabuena to Co Negro the coast shows several well-marked bights.

Bahia Tierra Blanca or Tierras Blancas has its name from the white sand
above the bay. Just as the other bays on this coast it is bordered by cliffs and
beaten by a never-resting surf. To land anywhere on this side of the island is
possible only under very exceptional conditions (fig. 48).

Pta Larga (Long Pt.) separates Tierra Blanca from the next bay; in the
background rises a rounded hill which I take to be identical with Guzman’s Co Ezrv-
gue. From a distance it seems to be formed by basalts of the higher horizons, but
no specimens were brought.

Bahia Chupones derives its name from “chupon’, in Chile a vernacular name
for Greigia sphacelata Reg. (Bromeliaceae), which has edible fruits (chupar = suck),
and applied by the fishermen of Masatierra to the extremely rare Hesperogreigia
as well as to Ochagavia; in this case the former, an inhabitant of the wettest
and loftiest ridges, is not to be thought of, whereas the latter, a typical xerophyte,
very likely occurs here, even if we did not observe it W of Tres Puntas, where
it covers rock faces (see Skottsb. 3 Pl. 97). The slopes round the bay are grass-
land, in the western half almost pure Avena barbata, otherwise with extensive
patches of the native Si#petum. All streambeds in this western section are dry
most of the year.

Loma Escarpada’, 385 m high where we crossed it near the main range,
separates B. Chupones from Villagra Bay, this taken in a wide sense. Off the
point lies Morro Vinillo (vifilla means “small hill planted with vines’, but the
name must refer to something quite different in this case); the gradient of the
surface shows the tilt of the lava beds (fig. 49).

Bahia de Villagra and its valleys, the bay taken in a wide sense and

‘In my field notes I called this ridge ‘Cordon Escarpado’’, and this name also appears
on the Chilean chart, and on my map I had used the same name for the ridge between Pangal

and Q. Minero, and as it was published in this sense (Skottsb. 3 pp. 890, 915), 1 have renamed
the other ridge Loma Escarpada.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 133

Fig. 47. View from Puente toward Bahfa Carvajal and Pta O'Higgins. Behind, Santa Clara. —
Photo K. Backstr6ém }/, 1917.

Fig. 48. Bahia Tierra Blanca seen from Pta Larga. — Photo C. Skottsberg °/, 1917.

134 C. SKOTTSBERG

Fig. 49. View from Portezuelo de Villagra looking SW, with Santa Clara in the background.
Morro Vinillo in the middle, right the south coast of Masatierra from Bahia Villagra to Pta
O’Higgins. — Photo C. Skottsberg 18/,, 1916.

extending from M. Vinillo to Los Chamelos. The broad slopes E of Loma
Escarpada, locally known as Los bajos de Villagra, are strewn with lava boulders
and furrowed by dry, stony streambeds. In the background two very conspicuous
mountains, already seen on the north coast, Co Tres Puntas and Co Chumacera
(Rowlock Mt., with a deep vertical slit), rise to a height of perhaps 500 m
(Branchi’s figure 650 is certainly too high). In the dry streambed below Tres
Puntas we found a patch of pangue, which reaches its farthest west here. Chu-
macera looks like an enormous rock slab standing on end. At the foot water
was found also during the dry season, and here is the westernmost luma forest
on this side, and below a threshold (see above p. 97 and Skottsb. 3 fig. 36 on
p. 896) with a small waterfall a grove of Boehmerza. The next gully has forest
down to about 300 m above the sea. Fig. 50 gives a good idea of the nature
of this country.

On all the earlier charts and maps the bay is presented as forming a regular
curve, but the aerial survey proved that this is not the case; see fig. 2. The
coast cliffs are lower here than farther west, but there are very few places where
it is possible to get down to the water. It can be done not far from Cerro
Negro, where we found access to the beach.

Villagra is watered by three permanent streams and densely wooded. The
scenery is even more grand than on the Cumberland side, with the sequence of

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I

Ww
Nn

Fig. 50. View from Portezuelo de Villagra looking W. From W to E Cerros Enrique, Tres
Puntas and Chumacera. — Photo C. Skottsberg 1°/, 1917.

summits from Co Agudo (sharp) to Mt. Yunque, more imposing perhaps from this
side than from any other (fig. 51). A peak between Agudo and Chumacera is
called Oreja del Conejo (Rabbit’s ear) by Guzman. The higher slopes are pre-
cipitous, with gradients of 60—v7o0° and sculptured with numerous hanging
gullies, carrying water after every rain, when many little cascades tumble down
from the summit of Mt. Yunque (fig. 2) between the carpets of ferns and pangue.
The Villagra valleys have not been levelled by erosion as much as Valle Colo-
nial, V. Ingles or V. Anson; the inclination in the middle and lower sections is
20—30. The forest in Q. de la Choza (responsible for this name is a small
shack below the lower timber-line, l.c. Pl. 85: 1) and on the slope of Mt. Piramide
(l.c. Pl. gt) is primeval; it extends up to the level of the Villagra pass and is
very wet and rich in species. Some maqui is seen here and there at lower eleva-
tions. The forest comes to a sudden stop 200—250, in cases 300 m above the
sea. On a former occasion (3 p. 895, Pl. 85:1) I have discussed the nature of
this timber-line. Below the forest degraded grass-land with foreign grasses and
herbs dominates over the natural S7zpetwm. Along the streams a fringe of pangue
runs down toward the sea (l.c. Pl. 85:2). Nobody lives in Villagra, but it is
grazed over by cattle on the lower slopes.

Mt. Yunque has already been described; I shall add here what Tenz l.c.
has to say about the summit. ‘Se ve arriba una altiplanicie muy extensa y suave-
mente inclinada hacia oeste a poca profundidad, de forma rectangular, rodeada

136 C. SKOTTSBERG

Sey
Seep, oe

Fig. 51. El Yunque seen from the Villagra slope c. 175 m above sea level. — Photo C. Skotts-
bergse Feroz.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I

Fig. 52. South coast of Masatierra from Cerro Negro to beyond Corrales de Molina. After a
water-colour sketch by the author.

de cordones en los cuales sobresalen varias cimas. Desde ellas nacen, en distintas
direcciones, quebradas en que corren cristalinas vertientes.’’ The buttress project-
ing S is prolonged to a ridge ending in a low cone, Co Negro, 190 m high (one
reading only; lc. Pl. 92: 1).

From £/ Vungue to El Verdugo the main ridge of the island presents the
picture of a sky-high rock wall (figs. 2, 11, 52). E of Mt Yunque several nearly
vertical, trough-shaped gorges have been dug out, each with a cascade and a
patch of forest and known as Corrales de Molina (corral = enclosure; probably
named for Padre IGNacio Mo.iNa, an Italian-Chilean naturalist of the 18th
century and author of a Compendio in which 3 plant species from Masatierra
were mentioned). Goat-hunters cross the ridge here and descend into the gullies,
which have been described with much detail by Weber who went there several
times. Fig. 52 shows that it is no easy going. Hence follows a naked vertical
wall, exposing the regularly stratified lava (fig. 52, right). Along the shore Playa
Larga extends, marked ‘‘Low beach” on the English chart, presumably a low
abrasion ledge. The country farther E is a succession of cliff walls and gorges;
see figs. 2 and 11. The sinister name £/ Verdugo (executioner, hangman, fig. I1)
reminds of the dangers on this coast, where no light warns the sailor.

9 — 537351 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I.

138 C. SKOTTSBERG

Fig. 53. Santa Clara from the air, looking SE. — Photo B. Frédin 8/, 1952.

Santa Clara.

Santa Clara or Goat Island is a barren, desolate islet separated from Masa-
tierra by a shallow, about 1500 m wide strait, dreaded for its strong currents
and turbulent waters. The depth is 19—20 fathoms according to Anson’s map;
I sounded 20—45 m.

The older literature gives little information about this island, but it was
described in some detail by Guzman pp. 48—53, to whom the reader is referred.
Various bays and morros are mentioned, with names probably given by the
fishermen. The circumference of the island is said to be 9 km, the area 500
hectares. Measurements on the new map gave a length of 3.5 km, a maximum
width of 1.25 km and an area of 2.5 sq.km. This map is based on the aerial
survey of 1952 and on Mr. Frédin’s photographs (fig. 53). The length profile is
seen in its full extension on fig. 48. A comparison shows that all the older maps
are very defective.

The island is everywhere bordered by steep cliff walls (fig. 53). Above is
a sandy table-land, studded with hills, of which the easternmost, incorrectly
placed on other maps, is about 375 m high (366, Lopez). Guzman calls it Co
Negros, but as we have one Co Negro on the south coast of Masatierra, I
named it Cerro Fohow in commemoration of the author of the well-known work
on the flora of Juan Fernandez. Close to the coast are several morros. The west
side of the island is beaten by a heavy surf and inaccessible, but on the inner
side, behind M. del Spartan, landing is — but not always — possible.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 139

On Anson’s map of 1741 Santa Clara is shown as covered with some kind
of arboreous vegetation. Walter does not tell if a landing was made, but the
name Goat I. shows that there were goats on the island, and Ulloa’s map of
1742 leaves it treeless. Just as Johow we found the place very barren and the
vegetation mostly formed by weeds with dAvenxa in dominance, all very dry
during the summer. When Giinther says that there were “algunos drboles en la
pendiente del este’ he either did see some specimens of Dendroseris on the cliffs
above the water or refers to Morro Spartan (also called M. de los Alelies;
fishermen are said to have collected seeds of Ja/thzola in B. del Padre and to
have scattered them on Santa Clara). Relics of the original flora found a refuge
on the morro, which is separated from the island by a channel less than 10 m
wide and blocked by huge boulders which are exposed at low tide. The current
rushes through the narrow channel, and to judge from the vegetation the goats
are unable to cross. I believe that Johow (p. 261) was right in assuming that,
before the introduction of this pest, Santa Clara must have looked much like
M. Spartan. The altitude of the island is sufficient to catch the trade clouds, as
shown on fig. 4, and we are told that rains are not unfrequent during the winter
months (Johow p. 261), giving origin to an abundant vernal flora of annuals.
During our brief visit in January we did not see any trace of water; but we
read in Instrucc. naut. p. 230 that a stream leaps over the cliff at the NW point,
a statement repeated by Guzman who calls it Chorro de Dona Maria |p. 50).

To judge from our passing observations, the geology is the same as of
western Masatierra, uncounted, light yellowish or brownish volcanic beds tra-
versed by numerous vertical dikes of a hard gray basalt. I regret having neglected
to bring samples of the lava beds for comparison with the rocks from Puente.
The dike rock was described by Hagerman p. 28 and found to be identical with
the basalt from the top of Co Negro. The sample came from a dike on M. Spartan.
We landed at the foot of this dike which forms a flight of steps leading to the
table-land of the morro; see Skottsb. 3 p. 924 and Pl. 103. The dikes project
as flat slabs above the softer beds.

Masafuera, former surveys and maps.

Until our visit in 1917 Masafuera was much less known than her sister island.
It had been inhabited more permanently only during the period of the penal
settlement I909—1I91I3, a misfortune that befell this ocean castle a second time
in 1927—1930. Only few scientists have visited Masafuera.

Of the early navigators few paid a visit to this island. One of Commodore
Anson’s captains, on his way to Cumberland Bay, happened to come up under
Masafuera and reported that the island was not, as former navigators had imagined,
a barren rock, but “‘almost every where covered with trees and verdure, and
was near four miles in length’. He had not been able to land, but added that
“it appeared to him far from impossible, but some small bay might be found
on it, which might offer sufficient shelter for any ship desirous of refreshing
ashore” (Walter p. 134). As four ships of the squadron were missing the Com-

I40 C. SKOTTSBERG

MASAFU ERA Ens. Toltén 20° Norte
P. ve

% Q. Sanchez
¢

Q. Negra
+Q. Sandalito

, Wee,
Loberia = SQ one

Nueva

} V { le Q Seca

.
Loberia
Ventana

Loberia Vieja =

oo, = Chorro
—— 2” Dona Maria

Scale 1:100000

GENERALSTABENS LITOGRAFISKA ANSTALT
STOCKHOLM 1953

Fig. 54. Map of Masafuera. After Skottsberg.

modore sent a sloop to Masafuera to look for them there, and when she returned
after a fruitless search, during which the island was circumnavigated, a report
was drawn up with the first real description of the island. Anchorage had been
found on the north side close to the shore but protected from S only (l.c. p. 156).
Pl. XXI is a view of the NE side of “Masa-Fuero’, as the name is spelled. It
shows the table-land up to the highest hills well forested, and we have little
reason to doubt that such was the situation 200 years ago. Four places along
the coast are marked with the letters a, 6, 6, and c; a seems to indicate a reef,
6 and @ are the entrances to two of the canyons. | believe that a also indicates
©. Casas and that 6 are Q. Ovalo and Q. Sanchez; c is the waterfall coming
down from Q. Larga. Pl. XXII, pretending to show the west side, is less easy
to read. I think that it represents the north-west section of the coast from Cabo
Norte south toward Loberia Vieja, but nothing like the perfectly cylindric tower
on the extreme right in the picture exists — the only thing I can think of is

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I4I

a very irregular rock close to the beach in Loberia. The promontory in the
middle is Buque Varado. Groups of trees are seen only above this place and
above Loberia Nueva.

Ulloa found the island very inhospitable: ‘La Isla de afuera de Fuan Fer-
nandez es toda muy alta, y tan escarpada, y escabrosa que no tiene parage
conmodo para desembarcar’’ (p. 287). On a former occasion (3 p. 796) I quoted
the narratives of Byron and Carteret. The island was well wooded in 1765.

The hydrographic expeditions despatched by the Oficina in Valparaiso paid
little attention to Masafuera. When Lopez reports that the island was “‘cubierta
de arbolado” in 1875 (p. 67) he must have included the Decksonia jungle on
the slopes of Mt. Inocentes. Johow (p. 96) asserts that the luma was common
all over the island except near the coast and in the highland, but this does not
necessarily mean that it formed extensive woods. We find the same statement
in Instrucc. naut. of 1896. It is not known when the destruction of the forest
started in earnest, but it is probable that the exploitation of the sandal-wood,
which led to its complete extermination, had serious consequences for the forest
as such. The names of two valleys and a place on the south coast testify that
a species of Sazfalum grew on the island. The possibility that forest fires have
ravaged the woods must not be forgotten. Foreign grasses spread into the cleared
spaces and prevented the germination of the seeds of the native trees. The
direct influence of the goats, introduced, I believe, during the 17th century,
remains to be found out. They greedily devour the arboreous Compositae and
the endemic herbs, but I cannot tell if they eat the leaves of the luma, naranjillo
and canelo. I can testify from my own experience that there was better forest
in 1908 than in 1917, an undisputable consequence of the activity of the con-
victs during the intermediate period. Nevertheless Giinther (1920) repeats the
old statement that Masafuera was covered with trees.

In 1895 the Oficina Hidrogrdfica published the first map showing the prin-
cipal topographical features. New dates had been provided by Johow. During
our visit in 1917 we soon discovered that this map was too defective to be used
as a basis for a map of the vegetation, and we did what we could to correct
it. The result was a sketch put at the disposal of the Oficina, which used it
for a new chart. When the flight over the islands was undertaken in 1952, Masa-
fuera was covered by clouds. A new map is urgently needed.

Main geographical features.

In shape and general appearance Masafuera differs profoundly from her
sister island. The island (fig. 54) is a solid rectangular block with slightly rounded
corners, tilted NE so that the escarpment along the west side is very much
higher than on the east side, where it is quite respectable (figs. 55—57). The
table-land is traversed by numerous parallel deep valleys running NE—E (figs.
57, 58), the high west wall by a number of precipitous gorges; this side has
much the same appearance as the south side of Masatierra E of Mt. Yunque

(fig. 52).

142 C. SKOTTSBERG

Fig. 55. North half of Masafuera looking W. Quebr. de las Casas at the extreme left. — Photo
K. Backstr6m Feb. 1917.

The question may be raised if not the eroding forces have been more active
in bygone times than they appear to be at present when so many of the streams
are dry or carry very little water during the summer half year. If it is true, and
this is believed by many, that the Ice Age was characterized by high precipita-
tion values, the climate of Juan Fernandez, otherwise not much influenced by the
glaciation in the southern Andes, must have been more rainy than now and, as
a consequence, the eroding forces stronger. The end of the Pleistocene left the
islands much as we see them now.

The coast lacks bays, there is no harbour, not even a sheltered cove to
accomodate small craft. The landing places from where the interior can be reached
are the entrances to ©. Sanchez and Q. Casas, but only under favorable con-
ditions with regard to wind and sea. It happens that an expedition has to return
with its errand unaccomplished. Abrasion has created a terrace along the west
side, continued by a reef studded with rocks (fig. 59). At the foot of the coast
cliffs a talus has accumulated, sloping down to a beach where the surf washes
out the finer material leaving the boulders. It is possible to scramble along the
shore from ©. Sanchez south round the island and from there north along the
west side to the north extremity of the Loberia Vieja plain, separated from the
Loberia Ventana beach by an impassable obstacle. Figs. 60 and 61 show that
it is no easy going, and with a heavy swell it may not be possible to pass the
southeast corner of the island. The opportunity to make this circuit round 3/,
of the shore and to proceed .from one canyon entrance to the next greatly

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 143

simplifies matters for the surveyor where, in contradistinction to Masatierra,
boats cannot be used to move from one place to another.

The longitudinal axis is directed N by W—S by E. The statements regarding
the size of the island vary a great deal: 9 miles (14.4 km) by 4 miles (6.4 km) with
a circumference of about 46 km and an area of 85 sq. km, all according to Lopez;
11.7 by 7.4 km (Johow’s map); Branchi, 10 by 6 km, area 84 sq.km; Giinther,
9.25 by 6 km, and Guzman, 17(!) by 7 km, a circumference of 55 km and an
area of 85 sq.km. The figures taken from my map are 10.3 by 6.2 km and the
area 57.6 sq.km.

Geology and morphology.

In 1908 Quensel, the only geologist who ever visited Masafuera, made a
hasty survey of the rocks along the coast from Q. Casas to Loberia Vieja and
north of Casas from the shore to perhaps 1300 m altitude. His field notes and
specimens served him for a description of the principal geological features. In
structure the two islands differ considerably. Both are built up of innumerable
volcanic beds of varying thickness and tilted about 20°, both exhibit two more
or less distinct horizons, both are traversed by vertical dikes, but here the
resemblance ends. On Masatierra the lower strata consist of lavas very rich in
olivine, interbedded with tuffaceous deposits, deeply weathered and denuded,
resulting in the formation of V-shaped valleys. The lower horizon of Masafuera
consists of hard, dark gray to black vesicular feldspar basalts (Quensel 2 p. 58), less
rich in olivine and very resistant to the denuding forces. These rocks prevail
up to approximately 1000 m. My notes from 1917 contain a statement that W
of Vicente Porras, where Playa Larga begins, a bed of “‘conglomerate’’ was
observed, traversed by harder dikes, but unfortunately no specimen was pre-
served. In a paper not quoted by Quensel, who did not mention the occurrence
of agglomerate beds on Masafuera, R. A. PHILIPPI described 14 rock samples col-
lected by Germain, most of them “‘losen, von der Hohe heruntergerollten Blécken
entnommen’’. I doubt that anybody will be able to identify these rocks from
Philippi’s unscientific description, but two of the samples suggest the occurrence
of tuffaceous deposits: they consist of ‘‘Rapilli’’ cemented together.

At about 1000 m a lava bed of a different aspect was met with, a light gray,
“iddingsite-bearing phyric olivine basalt’? (Quensel 2 pp. 58—63). Between 1000
and 1100 m a rock of quite another type occurred, described as a light yellow-
ish green soda trachyte. Apparently a closely related type is found also at
lower levels; boulders of this rock are so common in the talus along the
south shore that the place was called Tierras Blancas. Quensel arrived at the
conclusion that “‘lavas of soda-trachytic composition have been emitted at different
times and that they have alternated with lavas of more normal basaltic com-
position” (2 p. 66).

The upper horizon, from about 1100 to 1400 m, is formed by dense, ash
gray feldspar basalts, which probably form the whole upper complex of lava
beds with exception of the summit of Mt. Inocentes; these rocks very much

144 C. SKOTTSBERG

we Wg

Fig. 56. East coast of Masafuera between Pta Negra and Playa Ancha, with entrance to Q. del
Varadero. — Photo C. Skottsberg #4/. 1917.

Fig. 57. Part of the east side of Masafuera seen from the air. The highland is covered with
thick fog. — Photo B. Frédin §/, 1952.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 145

Fig. 58. View from the high land above Las Chozas looking toward the Ovalo, Pasto and

Sandalo valleys. — Photo C. Skottsberg 3/, 1917.

Fig. 59. The shore at Loberfa Vieja with a crag on the reef- — Photo K. Backstrém 17/, 1917.

146 C. SKOTTSBERG

Fig. 60. At the landing-place in Las Casas. Our party is about to embark, assisted by the crew
of the schooner. — Photo K. Backstrom 4/3 1917.

Fig. 61. As fig. 60, showing beach of large boulders. — Photo kK. Backstrom 9/3; 1917.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 147

Fig. 62. Slope of Mt. Inocentes seen from Cordon Atravesado, c. 1350 m above sea level. —
Photo C. Skottsberg 15/, 1917.

resemble the hard basalts between 400 and 600 m on Masatierra (I.c.). The sum-
mit of Mt. Inocentes consists of a still different rock, a slaggy olivine basalt
supersaturated with iron oxides (l.c. p. 70). The extension and thickness of this
bed is unknown.

As on Masatierra numerous vertical dikes traverse Masafuera, in this case
in W—E direction. The rock is extraordinarily rich in olivine and the dikes
may represent, at least in part, channels for the beds forming the summit of
the island (l.c. p. 74).

The difference in geological structure between the two islands helps, I think,
to explain the profound difference in morphology. The streams, eroding deeper
and deeper into the land, have excavated valleys which, in their distal parts,
are V-shaped, with steep sides, which rapidly become steeper inland, where the
perfect U-shape is retained in the canyon, with the entire narrow width of the
gently sloping bottom occupied by the streambed. In the innermost part, where
the land reaches its greatest elevation and the precipitation its highest figures,
lateral erosion has widened the valley and a fan-shaped series of hanging gorges
has been formed. The canyon ends in a high, almost vertical wall with a water-
fall. Figs. 63—65, 68—71, 73, 75 and 77 serve to illustrate the valley formation.
In the northern, drier and less high half of the island no deep canyons were
formed. The ridges left standing between the valleys are very unlike the ridges

C. SKOTTSBERG

148

2161 ‘qo WoNSy

ove

wo C

{ “NM OjoYd — ‘sesed Se] 9p ‘rqan() 0} aouejUy “fg “SIy

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 149

Fig. 64. In the Casas canyon after a few days of dry weather. The rhubarb-like plant is Gumnera
Masafuerae. — Photo K. Backstrom Feb. 1917.

Fig. 65. Looking down into the interior of Casas canyon from the crest of Cordén del Barril.
— Photo K. Backstrém 4/3 1917.

150 C. SKOTTSBERG

Fig. 66. The grass-covered table-land between the Casas and Vacas valleys, looking N. The
stone-building, an observation post overlooking the ocean, dates from the first convict settlement.
— Photo C..Skottsberg: 19/5. 19107.

in Masatierra, because the hard basalt layers have not been removed by denuda-
tion, but are left as a cover right down to the sea, forming gently sloping, wide
plains. Farther inland the ridge gradually narrows. See figs. 57 and 58. Beds of
greater resistance form ledges along the valley slopes and thresholds across the
streambed.

The south half of Masafuera.

Shrouded in clouds most of the time, Los /uocentes rises above the west
wall, an imposing dome forming the south half of the island. Opinions have
differed very much regarding the altitude: 2000 m on the older charts as well
as on Johow’s map, 1836 (Lopez), 1850 (Viel; 850 is a misprint), 2300 (v. Rodt),
1624 (Branchi) and 1840 (Guzman). My single reading, carefully worked out,
gave only 1500 m, a figure possibly too low. Toward W, N (fig. 62) and NE
the summit falls precipitously; it is connected with the north table-land by a
ridge, C. Atravesado, bordered on both sides by a precipice and so narrow that the
stretch above the Vacas valley, where a pinnacle rises from the knife-edge, is not
passed by the goats. Above Casas the highest point is c. 1370 m. Where Ermel
got the idea from that the summit is covered with eternal snow (p. 113) is hard
to understand.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS ISI

Fig. 67. Cordon del Barril, looking W toward C. Atravesado. Note goat track along the ridge.
— Photo C. Skottsberg 4/, 1917.

Q. de las Casas, once the headquarters of the convict settlement and the
ordinary landing-place, is called O. Baquedano by Guzman, commemorating the
corvette frequently sent to the outlying islands of the Republic. From the
beach, bordered by lofty headlands down to the water, we enter the valley, which
is about 200 m wide here, with slopes receding under an angle of 40—45° (fig,
63). It gradually narrows so that the streambed offers the only access into the
canyon. After a downpour the stream overflows and the valley is closed; a couple
of days with dry, sunny weather and the bed, filled with boulders of all sizes,
resting on the bedrock, lies dry, with pools of crystal clear water in the depres-
sions and back of the thresholds (fig. 64). The most spectacular part of the gorge
begins about 1 km from the entrance, where the width at the bottom may fall
below 10 m, while the walls are many hundred m high (fig. 65). From a botan-
ist's viewpoint Casas is one of Nature’s conservatories; see Skottsb. 3 pp.
639—640 and PI. 110. Johow called this place ‘“E] Pangal’’, and nowhere is the
pangue (Gunnera Masafuerae) more luxuriant (I.c. Pl. 111). About 2 km from the

152 C. SKOTTSBERG

Fig. 68. Entrance to Quebr. de las Vacas. — Photo C. Skottsberg 14/, 1917.

entrance and 210m above the sea the passage is obstructed by a threshold dam-
ming up a deep pool which cannot be passed round, and into this pool the
water comes down in a cascade. 0. Casas drains C. Atravesado. The eroding
power of the current is great, and I suppose that the stream is more or less
permanent during the winter half year, but it must have taken a very long time
to dig a gorge 1000 m deep through the hard basalt.

C. del Barril. Between Casas and Vacas lies a sloping, triangular table-
land, about 1.5 km wide along the top of the coastal escarpment, which is 125 m
high in this place, and extending west c. 2 km, where the cord6n proper begins.
The ruin seen on fig. 66 lies 190 m above the sea; the tilt, corresponding to
the dip of the lava beds, is 20° E. Three small gullies break the monotony of
the grass-land, Q. Chica, Q. Blindado and Q. Cabreros. Q. Chica (=small) is very
shallow and dry, but some luma trees are seen. QO. del Blindado (B. means
ironclad cruiser, but may have a very different significance here) is much larger
and the water comes down over a threshold about 500m upstream and has dug
out a miniature canyon. There is good forest between 400 and 500 m and some
groups of trees a little farther down. There is some forest also in Q. de los
Cabreros (Goat-hunters’ valley). Where the narrowing rocky Barril ridge, which
has its name from a barrel-shaped monolith, begins, the elevation is 730 m. By
and by the crest gets very narrow, but the rise is gentle, and there is a goat
track all the way up to C. Atravesado (fig. 67). The ridge slopes 50—60° on

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 153

both sides, and we look down into the Casas and Vacas canyons (figs. 65, 69).
From C. Atravesado the road N to Plano de la Mona lies open.

QO. de las Vacas (=cow; this place-name is old, but no cattle existed on
the island in 1908. Reintroduced in 1909, they had been killed off before 1917
and were introduced again in 1927). Following the beach south from Casas
past Pra Langlois we reach the entrance to this second large canyon, which
drains the highest part of Mt. Inocentes. It happened that when Casas was dry,
the Vacas stream was still alive. The entrance is quite picturesque thanks to
the deep side-valley separated from the main quebrada by a high spur ending
in a sharp peak (fig. 68). The interior has the same morphology as Casas; fig.
70 shows the drainage basin viewed from the Barril ridge, fig. 71 a typical, low
threshold of harder basalt. Some witches’ cauldrons were observed (fig. 72).

Little Vacas is accessible to a point about 1 km from the sea, where a very
high cliff barrier impedes further progress.

The easiest way to reach the table-land S of Vacas is to climb a low bar-
ranca at Playa Ancha, a \evel stretch of stony beach, from where Q. de los
Lnocentes with its untouched forest and fern-groves is gained without difficulty.
Crossing this valley high up we climbed the Inocentes-Vacas ridge which leads
to the top. To our surprise we were not stopped by the fern forest we had reason
to expect judging from our experience farther south; the stony and rocky ridge
was covered with Alpine heath which continued right to the Inocentes summit.
The ridge can also be gained from Little Vacas. The slope is, as seen on fig. 68,
steep, but it is terraced, and on our return we made a quick descent here.

From the ridge south of QO. Inocentes a good view is obtained across the
interior of the narrow Q. Angosta (fig. 73). The morphology is the same as in
Casas and Vacas, but the valley got its name because it is narrower than any
other, only about 5 m wide at the bottom, a magnificent blind alley where, c. 1 km
from the entrance, we are confronted with a precipice and waterfall once more.
The photograph shows the columnar structure of the threshold. At the foot of the
waterfall the vegetation is, as in all these gorges, luxuriant.

Quebr. del Varadero (varadero = ship-yard, which does not fit here; varar
also = to be stopped, and a strong surf may prevent passing the escarpment
south of the canyon entrance). Our first attempt to ascend Mt. Inocentes was
made from this valley at the place seen in fig. 74, where we gained the ridge
at 400 m above see level. On account of the slippery grass the climb was a little
hazardous. To begin with, the ridge itself was easy until, at c. 740 m, the tree-
ferns, which had begun to appear on the sides, gathered on the very crest, form-
ing an impenetrable thicket. Having crawled through the soaking wet barricades
of decaying trunks under the closed roof of the fronds and made perhaps 500 m
in an hour, we had to give up. The altitude was c. 950 m and Dzcksonia con-
tinued in every direction as far as we could see from a solitary canelo rising
above the fern roof.

A short walk into the Varadero gorge offers a strikingly wild scenery (fig.
75). A picture of the short and narrow side valley was published by Quensel
(2 fig. 3 on p. 43). Gradients of 60—8o0° are the rule in these gorges.

IO — 537351 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I.

SKOTTSBERG

154

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156 C. SKOTTSBERG

Fig. 71. A low threshold in the Vacas canyon. — Photo C. Skottsberg 13/, 1917.

Fig. 72. Two witches’ cauldrons in the Vacas canyon. — Photo C. Skottsber

; 8], 1917.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 157

Fig. 73. A view of the interior of Quebr. Angosta, seen from the ridge N of this valley, 550 m
above sea-level. — Photo C. Skottsberg 9/,; 1917.

The south coast. The strip of beach is narrow S of Varadero, and the escarp-
ment reaches an imposing height at Pra Negra (fig. 56). A short km beyond
this point a gorge with a cascade is passed, Chorro de Dona Maria, whether the
same lady to whom the chorro on Santa Clara was dedicated I cannot tell. The
talus W of this place derives its name, 7zerras Blancas, from the light colour of
the deposits. Along Rodado del Sandalo (rodar, to make a turn) the land rises gently;
the shore is fringed by a reef, on which the sea breaks with a thundering roar
(fig. 76). At Vicente Porras we arrive on the west coast. The long and broad abra-
sion terrace, Playa Larga, was referred to above (p. 142); a level plain like this is a
rare phenomenon on Juan Fernandez. At Loberia Vieja, “the old sealing grounds”,
where some fur-seals were still to be seen, the width is 1 1/, cable (277.5 m) ac-
cording to Ginther, a figure in good accordance with our estimation. Several hang-
ing gorges face the playa, coming down from Inocentes and C. Atravesado, and two
streams find their way across the plain to the sea. One of the gorges, described as
QO. de la Loberia Vieja (Skottsb. 3 p. 941) was surveyed by us. The entrance is
barred by very large boulders, and one has to climb along the wall to get into
the valley. Quensel published a photograph of the entrance (2 fig. 15 on p. 58).
The short canyon ends in the ever-present cliff wall with its waterfall (fig. 78).
The altitude of the valley floor is only c. 190 m.

A short but impassable stretch prevents us from reaching the remaining

158 C. SKOTTSBERG

inte
Pt 2

Fig. 74. Entrance to Quebr. del Varadero, from where, up the south wall (left) an ascent was
made. The -knob on the extreme right is shown in fig. 75. — Photo C. Skottsberg */, 1917.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 159

Fig. 75. Looking into the Varadero canyon. — Photo C. Skottsberg ?4/, 1917.

160 C. SKOTTSBERG

aed

Fig. 76. Coast of Masafuera between Tierras Blancas and Vicente Porras. — Photo K. Back-
Strom, 1%/5: 16017.

“loberias”’, L. Ventana (=window, a hole in the rock), Z. Nueva and L. del Bugue
Varado.

Playa del Buque Varado (“the beach of the stranded ship’) is a wide tongue
formed by the talus deposits; probably one or several landslides have occurred
here, where the barranca is of formidable height and steepness (fig. 78). A stream
coming from the gullies above has cut a winding bed (fig. 82).

Ensenada TLoltén is a shallow open cove W of Cabo Norte. When it is im-
possible to approach the east coast boats anchor here, where landing, as a rule,
meets with little difficulty. Consequently, numerous weeds mix with the native
herbs and grasses covering the slopes at the foot of the barranca. Unfortunately
Toltén is quite unfit as a base camp, as the highland cannot be reached, nor
is there a passage along the shore to ©. Sanchez.

The table-land N of Q. de las Casas.

The Plano de la Mona (“she-monkey’s plain’), a name of unknown origin
(there are no monkeys on Juan Fernandez), and the rim of the coast escarpment
can be reached from several valleys. The usual route takes us up a zigzag trail
from Casas, across a dry gully and through the large, branched and well for-
ested Q. del Mono (another queer name) to the abandoned convict settlement
Las Chozas, situated among the remnants of luma forest 550—650 m above sea

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 161

Fig. 77. Interior of the Loberfa Vieja canyon. — Photo C. Skottsberg 1%/, 1917.

162 C. SKOTTSBERG

SRI

Fig. 78. Pta del Iman and Playa del Buque Varado, seen from the sea looking S. — Photo
C. Skottsberg +°/, 1917.

level; another 50—100 m and the undulating grass-, fern- and pangue-covered
Mona plain, crossed by many little gullies, dry during the summer, lies open in
every direction (fig. 79). At one of the tributaries to Q. Pasto, 1130 m above
the sea, a pool of water in the rocky stream-bed made the place a convenient
camp site (“Campo Correspondencia’’). A short walk brings us to the edge of
the plain and to Las Torres (Towers), c.1370 m, and Co Correspondencia,
c.1420 m. Fig. 80 is a view of one of the Loberia gorges seen from Corres-
pondencia. The Alpine flora is well represented on these hills. Near the top of
the high western barranca some shallow depressions almost lack vegetation (fig. 81).
The soil is clayey and the surface, which shows distinct signs of water erosion,
strewn with stones and cracked in places. Some small boulders rested on short
pillars of clay. The ground is perhaps flooded during the winter. I was told that
patches of snow have been observed here, but this needs corroboration. Wind
erosion might be responsible for the formation of the small “tables”.

Avenida de las Cabras, “Goat Avenue’, is a well-beaten track running along
the very edge of the abyss from C. Atravesado to Co Verde. From about
1100—1250 m alt. one looks down on Buque Varado (fig. 82) and Q. del Iman
(Magnet gorge, fig. 83). One of the former inmates of the convict settlement told
me that he had been employed cutting a trail down the precipice to Buque
Varado, where also huts were built, but we saw no signs of either. There are

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 162

Fig. 79. View across Plano de la Mona toward the Larga and Sanchez valleys, elev. 940 m.
— Photo ©.) Skottsberg 25/, 1917.

Fig. 80. Looking down into one of the upper gorges of Quebr. Loberfa from an altitude of
c. 1400 m. — Photo C. Skottsberg 14/, 1917.

164 C. SKOTTSBERG

Fig. 81. Small depression at the west edge of the Masafuera table-land, c. 1400 m; naked
soil, on the stones mosses and lichens. Left, a patch of Lophosoria. — Photo C, Skottsberg 19/, 1917.

Fig. 82. Looking down on Playa del Buque Varado from c. 1200 m. — Photo C. Skottsberg
7/3 1917.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 165

Fig. 83. Quebr. del Iman seen from c. 1100 m. Left, Buque Varado. — Photo C. Skottsberg
He WONG.
3

Fig. 84. Plano de la Mona, looking N, with Cerro Verde in the background. — Photo C. Skotts-
berg 7/, 1917.

166 C. SKOTTSBERG

Fig. 85. Looking S toward Loberfa Nueva from Avenida de las Cabras N of Buque Varado.
— Photo C. Skottsberg 7/, 1917.

A GEOGRAPHICAL SKETCH OF THI AN FERNANDEZ ISLANDS

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168 C. SKOTTSBERG

only scattered patches of trees on this side, while ferns, especially Lophosorza,
pangue, grass etc. form a more or less continuous cover. A picture taken from a
point between Buque Varado and Iman gives some idea of the scenery (fig. 85).

At the north extremity of the plain stands a green, rounded hill, Cervo Verde
(fig. 84), mentioned as Pico Norte in Instrucc. ndut. p. 230 and said to be 1340 m
high. This figure is too high, I believe.

From Cabo Norte to Q. de las Casas.

The coast cliffs from Cabo Norte to the Sanchez shore are almost perpen-
dicular and about 300 m high. Half way a cascade tumbles down to the beach
in two leaps, coming from the gap of Q. Larga. Just as the other valleys on the
east side this has a forest belt above c. 300 m and grass-land below. Here the
trees cover also part of the ridge between Larga and Sanchez, apparently the
largest continuous patch of luma forest on Masafuera.

O. de Sanchez is very long, narrow and twice furcate, and there is forest
in all the: branches. None of the other valleys is quite destitute of trees. The
small QO. Negra (also called Q. del Plan Negro) does not reach the shore. Q. del
Sandalito and QO. del Sandalo do not seem to offer anything of particular interest.
The talus slope permits a fairly comfortable passage as far south as to the large
QO. del Pasto. ““Pasto’’ means pasture, and there is better grass-land with abun-
dant native grasses along this part of the coast than anywhere else. Pasto is a
large valley with its sources near the edge of table-land. The main stream runs
over a high threshold. S of Pasto a land-slide is crossed before reaching Q. de/
Ovalo, which derives its name from a conspicuous hill inside the entrance, a good
example of columnar structure. The passage is a little fatiguing until we are op-
posite QO. del Mono, another big valley emptying its waters through a narrow
crevice (fig. 86). From here going is easy to Casas, where we have completed
our circuit of the island.

Human influence on Juan Fernandez.

The discovery of the two virgin and uninhabited islands in 1574 was soon
followed by the first inroad into its living world. If we are to believe the histo-
rians cited e.g. by Guzman, Fernandez returned and settled on Masatierra where he
introduced 60 Indians (Weber), a few goats and pigs and devoted himself to
agriculture, fishing and sealing. His stay is said to have been short, but his goats
remained and multiplied. After Fernandez’ death the island was turned over to
the Order of Jesuits, and, in the beginning of the 17th century, the first fruit-
trees were planted and vegetables introduced (Weber p. 162). During the “era
of the buccaneers’ the islands served as a place of refreshment, but the damage
was, I daresay, confined to the surroundings of the harbours. Their first visit is
said to have taken place in 1680. A change for the worse came when L’Heremite
reported that the precious sandal-wood was abundant. I refer to Johow’s instructive
account of the history of this ill-fated tree, pp. 127131. L’Heremite’s visit took
place in 1624, and already at the middle of the century a lucrative traffic was

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 169

Fig. 87. Ruins of the old Spanish fort in Cumberland Bay. — Photo C. Skottsberg *8/, 1917.

in full swing between the islands and Peru. Toward the middle of the 18th century
there cannot have been very much left on either island, or the sandal-wood should
not have escaped Anson and his men who spent months there and must have
known of the former trade.

The hunt for sandal-wood ended in the extermination of this species and
had, I presume, serious effects on the forest in general, with the final result that
all the lower slopes became cleared of trees, leaving the field open for an in-
vasion of weeds; Ulloa for instance was struck by the abundance of Avena.

Having suffered serious losses from the raids of the martial English navi-
gators the Spaniards took possession in 1750 and made up their mind to trans-
form Masatierra into a stronghold. I shall not dwell upon the history, enough said
that in 1779 7 places were fortified. Part of the walls in Cumberland Bay are still
standing (fig. 87), and old guns lie about in other places (fig. 33). Nor have I any
good reason to retale the melancholy history of the time when Masatierra served
as a prison for banished patriots during the years of resurrection and later. Its role
as a depository for undesirable citizens came to a definite close in 1855. Then
came the tenants.

A source of destruction of the forest, not at all negligible, were the fre-
quent visits of American whalers during the 19th century who called to supply
themselves with water, wood and goats’ meat. The large herds of goats had been

Il — 537351 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I.

1710 C. SKOTTSBERG

a great asset during the era of the corsairs, and in order to stop this traffic
mastiffs were landed as early as in 1686 or 87 (Burney IV p. 210). The result
was that the goats had become very shy at the time of Anson's visit; he estim-
ated them to be about 200 only, but the dogs were plentiful, savage and dan-
gerous. His men saw no dogs on Masafuera, whereas goats abounded, but later
dogs were introduced also to this island which was spoken of as ‘‘Isla de Perros’’.
There were some dogs left when Johow went to Masafuera in 1892, but none
in 1908. In 1830 they were exterminated on Masatierra. Freed from their worst
enemies, the goats increased in number; in 1877 Viel estimated them at 3000.
The new lessee von Rodt persecuted them for the sake of their hide, and in
1892 the number had sunk to about 1000, while there were 200 on Santa Clara
and 4000 on Masafuera; if these figures are reliable is hard to tell. We came
across a small herd on the south side of Masatierra in 1917, and once in a while
a daring hunter brought down a buck from the inaccessible crags where they liked
to hide themselves. At that time they enjoyed the protection of the Government —
the descendants from Robinson Crusoe’s goats were regarded as sacred. They
were abundant on Masafuera, and even if their number was reduced during the
time of the convict station, they soon recovered. To judge from what Weber tells
us the island swarmed with goats after the departure of the convicts in 1930,
and they went right down to the beach. Their damage to the indigenous vegeta-
tion (and thereby to the fauna) cannot be estimated nor disputed. Their ravages
had left their marks everywhere, and several peculiar endemic plant species
were on the verge of extinction in 1917.

Beside goats Juan Fernandez brought pigs which ran wild; but they were
killed off long ago, nor are there any wild asses left, while cattle and horses
came to stay. I suppose that ever since the first Spanish colony 200 years ago
domestic animals have existed on Masatierra. In 1813 there were 1000 sheep, 100
goats, 500 cows, 200 horses, 10 mules and 40 pigs (Guzman p. 178), but a decade
later very few seem to have been left. Again the Government stocked the island
(Sutcliffe 2 p. 206), and in 1833 there were 350 sheep, 120 tame goats, 70 cows,
15 horses, 8 mules and 30 pigs (Guzman l.c.), but the tenants took no interest
in farming, and in 1860 Mackenna registered 15 sheep, 98 cows, 15 horses and
15 mules only. In 1878 and 1892, only horses remained.

It is easy to understand that the early visitors got a very favourable impres-
sion of Masatierra, its agreeable climate, volcanic soil, evergreen forests, luxuriant
verdure and good water, and concluded that the island was fertile if only its
natural resources were utilized. Certainly they did not fail to observe that there
was very little level and moderately sloping land, but the valleys of Cumberland
Bay and Pto Ingles must have looked inviting. We can also understand that in
those times it did not occur to anybody that the living world was unique, but
it is strange that educated people like Governor Sutcliffe or Mrs. GRAHAM could
dream of Masatierra as fit not only to feed a considerable population but to be
able to export its products. Mrs. Graham, cited by Sutcliffe (2), wrote in 1824:
“The island might maintain easily 2000 persons, exchanging the surplus of beef,
wines, brandy, for bread and clothing; and its wood and water would render it

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS i fB

invaluable as a port.’’ And Sutcliffe (1 pp. 442—443) cites a letter written to him
by one MANUEL DE SALAS, Sept. 1, 1835, when Sutcliffe had been appointed
governor:

“The clays and soils of various colours brought to my memory the minerals of
Golconda and Brazil... their (the islands’) geographical position places them in a like
situation and renders probable the same effects and equal productions . . . One occurs
to me which might be of great importance; such is the making of bricks, of which
immense quantities are brought from England and the United States, and for which
you have all the elements in abundance; there are, the firewood, the clays, and, above
all, the idle hands.”’

Still the tenants, who succeeded each other, did not coin money. Guzman
made an estimation of the arable land, 110 hectares on level ground, 70 on the
lower slopes; in addition there were 20 on Santa Clara and 100 on Masafuera,
and, if the forest was cleared, grazing land of much greater size would be obtained.
The colonists took no interest in cattle. Lopez argued (1878) that if the tenant
were more enterprising, he would go in for cattle and sheep, of which the island
could maintain 10000 — the expectations had grown! Viel’s report (pp. 19—22),
founded on his observations in 1877 — a few days I presume — reflects the
ideas of his time regarding the future of Masatierra. He admits that its fertility
was more apparent than real although chonta and sandal-wood grew in the for-
ests, both highly appreciated for their valuable wood and a possible source of a
small trade — it was 50 years since anybody saw a sandal tree! A road around
the island would become a great asset, because various industries could be started
in different places... Timber was plentiful, but the contract forbade the lessee
to use it, because the disappearance of the forest would endanger the water
supply. Viel’s personal opinion was that this danger did not exist, “the island,
situated in mid-ocean, accumulates the clouds and condensation produces the
rains which feed the streams and irrigate the land; thus there is no reason why
the lack of trees would have a contrary effect’. He forgot all about erosion. He
recommended to His Excellency to authorize the lessee to utilize the forest; be-
sides, he wrote, ‘‘if dead trees are left lying, the fresh growth will be poor and
raquitic (!)’’; one cannot help wondering why, after thousands of years, when the
fallen logs were left to decay, there was still dense, healthy forest. Unfortunately,
“the quality of the native timber was not good, for which reason it should be better
gradually to replace the indigenous trees by better ones, which later would become
a source of income to the Republic. Pines should be planted at once on a large
scale; it would be wise to take advantage of the present tenant who would be
glad to look after the plantations, provided that some minor behalves which
wouldn't in any way harm the State or render his contract more favourable were
conceded him.’’! It was necessary to supply good timber; if not, the whalers would
cease to come. Fruit-growing ought to be improved, wheat grew well, potatoes
could be exported to northern Chile with great gain. In this connection Viel says
that zarzaparilla — here = Acaena argentea, a noxious weed — was common and

1 Translated from Spanish.

172 C. SKOTTSBERG

maqui not less abundant; it is not clear whether or not he regarded them as a
source of wealth.

As a more promising industry than forestry Viel recommends raising cattle.
The island could easily maintain 1000, during the Spanish time there had been
800, and now the number of domesticated animals was 98 cows and 50 horses,
but many had run wild. The 3000 wild goats ought to be protected by a closed
season; 800 skins had been exported recently. Apparently the fur seals did not,
in his opinion, need protection: “the output was mediocre, a consequence of the
settlers not being sufficiently intelligent and industrious’! — he forgot that the
poor result of sealing was a consequence of ruthless persecution during a cent-
ury. The average number of seals killed per annum was 700 — one would call
this a fair number considering the small size of the islands. Fishing was neg-
lected, only very little salted or dried fish had been exported. There was plenty
‘“langosta’, and it ought to be made into preserves. Viel concludes his report
with the following words: “‘El] estado actual es bien lamentable.’’ Had the Govern-
ment listened to his recommendations the situation would have become much
more lamentable.

Of this official report to the Minister of Finance and to the Governor of
Valparaiso FELLENBERG published a translation, but he protests against Viel’s belief
that the disappearance of the forest would not harm the water supply. As an appen-
dix he published 3 letters from the new tenant, addressed to his relatives in Bern.

ALFRED VON RODT, of Swiss descent, an ex-officer in the Austrian army
and a well-educated gentleman, settled on contract on Masatierra as tenant of
the islands. Encouraged by the many favorable reports on their resources, he ex-
pected to make his fortune, and he had some money to invest in the enterprise.
His first letter, written shortly after his arrival and dated June 5, 1877, contains
dates on the size and position of the place; he tells that there were 7000 wild
goats and pasture for 1000 head of cattle and that he intended to start “‘verschie-
dene Industrien’”’. Evidently he went to work without delay; on March 13, 1878,
he writes that he had timber worth 3000 Dollars ready to be shipped to the coast.
He reported from Masafuera 17.6. 1878 that this island was considerably larger
than Masatierra (!) and that there were large forests and the most beautiful grass-
lands where it would be easy to feed 20000 sheep. He had killed more than
700 fur seals.

The castle in the air vanished, von Rodt never founded any new industries
on Masatierra, there never was a sheep-farm on Masafuera. He lost his money,
but he remained true to his beloved Masatierra, where it is easy enough to make
a living, but perhaps not a fortune. The von Rodt dynasty was still going strong
when we visited the islands; the sons of Alfred made their living as lobster? fish-

1 Translated from Spanish.

2 Professor KARL LANG, head of the Dept. of Evertebrates in the Nat. Hist. Museum, Stock-
holm, on my request kindly made a diligent search for an English equivalent to the Spanish
word langosta as name of the large Decapod /asus Lalandei (formerly known as Palinurus
Jrontalis), which from a scientific viewpoint is no real lobster. He reports that Palimurus vul-

garis and related forms are called “‘rock-lobster’’; “spiny lobster’ is another name. For the sake
of brevity “lobster’’ or “‘langosta” will be used here.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS LTS

ers, cultivated their corn-fields and gardens and reared a new crop of sturdy
fishermen.

The damaging influence of von Rodt’s reign on the native flora and fauna
must not be overrated. It is true that the maqui spread, but also that the goats
decreased in number. The population remained very small, and the cultivated
soil did not extend much. In comparison with recent disasters the encroach during
the last decades of the 19th century was of little significance, except, perhaps, with
regard to the chonta palm, one of Nature’s great treasures. And BURGER tells
us (p. 111) that when von Rodt failed to “develop” the islands, he became a
protector of their nature.

In 1877 the population of Masatierra counted 64 persons, 29 of whom were
male, children not included (demographic figures taken from Guzman p. 37), in
1878 74 (22), in 1879 141 (51), and in 1880 147 (55) — the rise probably due to the
arrival of labourers contracted by the tenant. Then it began to sink: 82 (24) in
1885, 61 (23) in 1886. In 1885 the German merchant ALEXANDER ERMEL visited
Masatierra as partaker in a tourist excursion. As many before and after him he
fell for the beauty. of the scenery and he returned with fantastic ideas of the value
of the natural resources, which nobody had understood to utilize properly.1 Con-
sequently he did not hold a very high opinion of von Rodt. Even barren Santa
Clara became fertile in Ermel’s eyes. For Masafuera only no great future was in
store; it was too inaccessible. ““Daher wird Masafuera ftir die Zukunft allein die
traurige Bestimmung haben, als ein in Wolken eingehiillter, schauriger Felsen
dazustehen, dessen Nahe die Seefahrer soviel als moglich vermeiden werden”
(p. 115).

In spite of the very short duration of his visit (3 days), Ermel judged him-
self competent to discuss the economic importance of the islands for Chile. Masa-
tierra might well serve as a health-resort. Its principal industry would be fishing.
Here he was, as the future showed, right, and the same is true when he argues that
the seals needed protection. In former days the seals had been extremely abund-
ant, not only the fur seal, but also the sea elephant. L’Heremite saw thousands
of sea lions and seals (Burney III p. 18). Walter reports on the “sea lions’, which
occurred in large herds, but Pl. XIX shows that his “sea lions’ were sea elephants.
Ulloa tells us (p. 287) that “the beaches and rocks were everywhere crowded
with seals in such abundance that no free space was left where one could walk;
they did not leave a passage between them’.2 Captain ROGERS (1709) says
that a compact string of seals lined the beach of Cumberland Bay (Guzman
p. 215). The main slaughter of the fur seal commenced in 1797; it is reported that
in 1801 a single ship carried one million skins to the London market. If this be

1 These illusions have been very tough. In an American newspaper (Meridian Star, Miss.)
the following paragraph appeared on March 1, 1928: “Crusoe island fruitful. — Juan Fernandez
Island, on which Alexander Selkirk, the reputed original of Robinson Crusoe, lived for four
years, is one of the most fruitful spots in South America, according to a recent survey. Every
known plant seems to grow there. One Frenchman who was shipwrecked there 40 years ago
refuses to leave.’ No comments needed, but the climax is priceless.

2 Translated from Spanish.

174 C. SKOTTSBERG

true, it is a wonder that sealing could be practised with a profit over a period of
40 years. The seals have disappeared from Masatierra, the sea elephant is extinct
in the waters of Juan Fernandez. As I told above, we found a small herd of
fur seals at Loberia Vieja. Since 1891 it is protected by law, to what effect I shall
not say.

But let us return to Ermel. Even if fishing ranked first, the wealth of the
land was by no means contemptible: timber and fuel, chonta, sandalo, charcoal,
“womit Herr von Rodt in den ersten Jahren seiner Pachtzeit einen lohnenden
Anfang gemacht hatte’; all kinds of fruit, probably also grain and wine, and
breeding cattle, llamas and vicunas should be tried. He also underlined Masatierra’s
strategic position.

However, nothing happened except that the population continued to dwindle,
reaching its lowest figure, 35 (12), in 1893. When Johow revisited the islands in
1895 it had grown to 54. In 1891—g92 Johow headed an official commission and
he outlined a program for the future management of the islands. This document
was published in his book, pp. 267—274. It is not without interest, and a sum-
mary will be given here. It begins by stating that the utilization must be based
on the principal marine products, the langosta (Yasus Lalandez), confined to
Juan Fernandez and the Desventuradas Islands (San Ambrosio and San Felix), and
the bacalao (Polyprion prognatus), also absent from the coast of the mainland. Even
the latter would become a valuable article of export, the more so because it is
one of the worst enemies of Yasus. The waters abound in many other kinds of
savoury fishes. When, however, the commission recommends to repeal the law of
1891, prohibiting the destruction of the fur seals, with the motive that seals are
the most dreaded enemies of the fishes, no responsible authority in our days
would agree. Poachers can be relied upon to keep the number of seals down.

Johow states that agriculture will never become profitable, for the simple
reason that there is very little arable land and that the soil, once deprived of
its natural vegetation cover, will be subject to erosion. Wheat should be imported
from the mainland and land utilization limited to cultivation of potatoes and veget-
ables for local consumption. Nothing is said of the aspects as grazing land.
There were very few animals in Johow’s time. After these introductory remarks the
commission proceeds to answer a number of questions.

With regard to the administration, some kind of authority should be estab-
lished on Masafuera. A small steamer would become necessary to serve the com-
munications between Masatierra and Valparaiso. An “inspector de colonizacion”’
should be appointed. The question whether it would be necessary to prohibit
the cutting of chonta and sdndalo is answered in the affirmative. Johow estimated
the number of full-grown chontas at 100 to 150, and he had occasion to visit
the last living sandalo. They were already under provisional protection and it
was recommended ‘no solo por motivos meramente ideales o sea cientificos, sino
tambien por consideraciones de cardcter comercial i econdémico” that the law
should remain in force and violation be subject to severe penalty: “‘No seria, a
nuestro juicio, exajerado castigar la destruccion de un solo ejemplar de chonta
o de sandalo como delito de robo comun, i prohibir.tambien su corta bajo pre-

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 175

testo cientifico.’’ And the commission made public that, at repeated occasions,

the crew — they forgot to add “‘and officers’’ — of vessels belonging to the Armada
Nacional had possessed themselves of large quantities of chonta trunks — with

the assistance of the colonists. (Such is the attitude of the majority of people
all over the world that if an individual stands before the last living specimen of
a plant or animal, he will grab it, because if he doesn’t, someone else will have it.)

In the 1890's there were no plantations needing protection, but in Johow’s
opinion the existence of the native forest was jeopardized by the ravages of a
parasitic fungus, Axtexnarta scortadea Berk. (Limacinia fernandeziana Neger ex
Johow), and he feared that it would ‘“‘concluir poco a poco con la vejetacion de
muchas quebradas’’. The fungus is still plentiful, but its killing capacity was, as
far as I could see, greatly overrated. The commission recommended to make pun-
ishable to set the woods on fire, and to enjoin the inhabitants and visitors in
need of wood to use only the fallen trunks. If I remember right there later was
a regulation that only the maqui could be used for fuel. In the report nothing
is said about the damnable habit of visitors to rob the forest of the stately en-
demic tree-ferns which, always without success, they tried to plant in their gardens
on the mainland. Ermel saw the lifeboats of his ship return laden with young
chontas and tree-ferns and looking like gardens afloat. I have witnessed the same
traffic myself, but I do not know if it still flourishes — there is a long way now
to the young chontas and to the tree-ferns.

A question if fishing with dynamite ought to be forbidden was answered in
the affirmative, but not because it would reduce the fish supply very much —
though it is admitted that the demand for fish might be greater in the future —
but rather because “‘la dinamita... manejada por personas de poca o ninguna
ilustracion, facilmente orijina funestos incidentes .. .’’. With regard to the langosta
no scarcity had made itself felt, but it was recommended to leave it in peace
during the breeding season.

Among the remaining questions, of which many were referred to specialists
and not answered by the commission, one is of interest: it was recommended to
exterminate the wild dogs and the native buzzards on Masafuera. The motive was
to take better care of the goats, still considered to be an important source of
meat; the meat is, I admit, very palatable. A bounty was therefore suggested.
Nobody will grieve the loss of the wild dogs; they are not good company, but
in the case of the buzzard, Buteo erythronotus exsul, the care for the goats gets
in conflict with the interests of science. In his catalogue of the insular Ornis Johow
remarks that the “aguiluchos quizas representan una variedad endémica si no una
especie distinta de la del continente’’ — nevertheless he would not hesitate to
exterminate it. Had he known the flora of Masafuera better, he would have been
less kindly disposed toward the goats.

Finally the commission underlines the urgent need of regular communications
with the mainland with rapid steamers permitting the products of the fishing in- -
dustry to arrive fresh at their destination.

Thus the wants and needs were stated, but little was accomplished. The
wild dogs on Masafuera were exterminated, but fortunately the buzzard escaped.

176 C. SKOTTSBERG

Fig. 89. Lobster fishers working for Recart y Doniez by their boats during the closed season.
Cumberland Bay. — Photo K. Backstro6m Dec. 1916.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I

Fig. 90. The take is brought into the Recart y Doniez factory; left, live lobsters; right, the
metal cooking baskets. — Photo C. Skottsberg °/, 1917.

Fig. 91. The harbour in Cumberland Bay, showing three schooners at anchor, a motor launch
and a number of fish-chests used for storing the take until shipped alive to Valparaiso. —
Photo C. Skottsberg 3/, 1917.

12 — 537351 The Nat. Hist. of Juan Fernandez and Easter Est. Viole fe

178 C. SKOTTSBERG

Fig. 92. The village in Cumberland Bay, seen from the slope of Salsipuedes. — Photo C. Skotts-
berg 24/, 1917:

Masafuera got no supervisor, the management of the forests continued as before,
the chonta continued to be cut unlawfully —- a common occupation during the
winter months, Weber says p. 116 — nothing was done for the fishing industry,
no steamer came and went to the benefit of the inhabitants. Life went on as before.

Let us return to the langosta. Ever since the days of the discovery of the
islands this magnificent crayfish had been appreciated as very good eating, and at
Anson’s time the beaches literally swarmed with large-sized specimens. No in-
strument was needed to catch any amount in one or two feet of water. How
long this happy state of affairs lasted I do not know, nowadays the best catch
is made in depths from 40 to 80 meters. In spite of the value of this marine
product, which from time to time was shipped to the mainland, none of the
tenants had the means to organize the industry on a larger scale. A company
was formed in 1893, but even with a catch of 35 to 40 thousand annually and
exporting some 40000 tins of preserved tails, the business did not pay, and a
new company formed in 1900 also failed (see Guzman p. 209). In 1914 Messrs.
Recart y Doniez started their establishment. We had the very best opportunity
to watch the fishing from the catch of the sharks used as bait to handling the
langosts in the factory (figs. 88—g91) and to the export of the living animals, in
the company of which we made our return voyage to Valparaiso onboard one of
the schooners. The stern is built as a well with free circulation of the water, and

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 179

Fig. 93. Carica papaya in the garden of Mr. Charpentier, Valle Anson. — Photo C. Skotts-
layers By G7

if the passage could be completed in 21/, to 3 days all went well, but in too
calm or too bad weather the motor alone did not help very much, the crossing
was slow and it happened that very few of the lobsters arrived alive. Two minor
companies were also in operation. The closed season was strictly observed. On
the 2d of January at 6 a.m. a rifle shot signalled the open season, and the
launches towed a string of boats, each party rushing to gain the best grounds
which it monopolized during the season. Recart y Doniez also held the licence
to fish at Masafuera and sent a schooner thither several times. I don’t remember
having heard that any kind of fish was exported at that time, but fish was the
staple food on the island. About 200 people lived there, most of the men being
employed by the companies. The village in Cumberland Bay (fig. 92) looked
quite inviting with its neat, if not always too well kept houses, vegetable gardens,
fruit trees and flower-beds, set among exotic trees like araucarias, eucalypts,
poplars, pines, A/b/zzta, Eriobotrya and so forth. Johow has published a paper
on the plants cultivated in the islands, and also a list in his book pp. 263—266.
With the exception of the fig and the quince most of the fruit trees yielded in-
different fruits. The climate is of the Mediterranean type, but at the same time
pronouncedly oceanic, and the lack of a period of real warm weather seemed to
be responsible for the failure of Citrus fruits. The more surprising was it to find
the papaya in cultivation in a garden belonging to a colonist of French descent.

180 C. SKOTTSBERG

‘ig. 94. Threshing the wheat with a tropilla of horses, Cumberland Bay. — Photo K. Back-
strom I917.

Fig. 95. Chaff before the wind. Same place as fig. 94. — Photo K. Backstrom 1917.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 181

Fig. 96. A rodeo in Cumberland Bay. — Photo K. Backstrém 1917.

The specimens were small, and so were the fruits, which we had no opportunity
to taste (fig. 93). This seems to be the only tropical tree thriving in this warm-
temperate climate. Several ornamental plants have become naturalized in the neigh-
bourhood of the village, e.g. Zantedeschia aethiopica, Pelargonium spp., Tropaecolum
majus, Matthiola incana and Lochnera rosea.

Wheat-fields were few and small and harvesting methods quite primitive;
see figs. 94 and 95. Most of the flour was imported, and there was also a shortage
of potatoes. Unfortunately the number of animals had increased very much, much
more than would be deemed necessary, but milk-cows were few. Imported wine
was the daily beverage, here as in other parts of Chile. Herds of cattle roamed
through all the valleys from Pto Frances to Vaqueria on the north and from the
foot of Mt. Yunque to Puente on the south side of the island. I regret having
neglected to ask their number; every animal had its owner. Fig. 96 shows a
rodeo in Cumberland Bay. Only the herd in Vaqueria was left to run wild. Horses
were seen in some valleys, but I cannot remember having seen any sheep, and
we were never offered mutton.

After 1917 the population remained about the same for years, but in 1930
it had increased to 298, of which 155 were male, including the boys. Ten years
later it had grown to 434 (225).

I shall leave Masatierra for the present and try to follow the fate of Masafuera.

182 C. SKOTTSBERG

Fig. 97. View from the south slope of Casas valley, Masafuera, showing the headquarters of the
first convict settlement. — Photo K. Backstrom Feb. 1917.

The first house was built in 1867 (Weber). Goat-hunters or fishermen used to visit
the island. In 1908 we found some people staying there, but there was no perma-
nent settlement. The next year saw a radical change. The full history is told by
Guzman pp. 87—101. A short summary is given here. Guzman begins with the
following words: “‘Tal vez la descripcién de la flora natural de la isla de Mas Afuera
que hiciera el sabio sueco Skottsberg, hizo creer a un Ministro de Justicia del
Presidente Pedro Montt, que dicha isla era el sitio ideal para la fundacion de un
presidio agricola.’ Certainly Guzman is joking, the more so as I tried on my
return to start a campaign in favour of preserving the native flora and fauna
— not the goats however! — and never said a word about Masafuera as a suitable
place for agricultural or any kind of commercial experiments. Two months later
a decree was signed, transforming the island into a settlement of criminals, and
in April 1909 “‘Carcel Pedro Montt”’ received its first inmates and soon after a second
lot, together 170. For the State it was no cheap affair to erect substantial headquar-
ters at the entrance to Casas valley, to build a jetty in the turbulent water of the so-
called landing-place and a schooner to serve the colony, but the parents of the
scheme were optimistic and thought that once in operation, the establishment
would soon become self-supporting, a miracle to be performed by cutting trees
and cultivating the soil. Goats were plentiful and easy to get hold of, at least to
begin with. ‘Tal era la conviccidn que tenian los creadores del presidio en la fe-
racidad de la estéril Mas Afuera, que al alcalde se le designo con el singular titulo

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 183

a A Ee

Fig. 98. The ruined forest and village of Las Chozas, Masafuera. — Photo K. Backstrém Feb. 1917.

de Director y Jefe de Cultivos’’, Guzman writes. This official was a thrifty fellow,
but he could not call up arable land where there was none. Houses were built in
many places, vegetables sown and potatoes planted, cattle introduced and killed
surreptitiously by the convicts, but much food had to be imported, and when the
boarders obtained permission to send for their wives or relatives to keep them com-
pany the population rose to 350 persons. Good luck did not favour this humanitarian
enterprise, the situation on the island became, to say the least, unpleasant, the
schooner was shipwrecked and lost, and in 1913 Masafuera was abandoned. The
buildings in Casas were in tolerably good condition in 1917 (fig. 97), but the
wooden huts more or less fallen to pieces. Ruined forests, abandoned potato
fields and a host of new weeds told the story (fig. 98). Of domestic animals only
two horses could be discovered. We caught them because we could use them.

But, as Guzman says, “los juristas no podian aceptar que una Naturaleza
tan prdodiga en helechos y en plantas sub-alpinas, no fuera generosa también para
con las legumbres, hortalizas y frutales . . ."’, and in 1927 ‘‘Prision Carlos Ibanez’’
was established, this time not reserved for ordinary criminals — in addition 150
political offenders were exiled to Masafuera. More trees were cut, new houses
built, new seeds sown, but the harvest was no richer than before, and in 1930
the colony was discontinued. Guzman seems to fear a repetition of this sad story,
to judge from his concluding remark “‘ . . . el hombre con sus disposiciones legales

184 C. SKOTTSBERG

Table J.
Lobster catch 1940—46. After Guzman.

Gross weight in kg

Number

Total Living Tinned
TGA ORR crs) i or tea 109 250 BIZe5a ir 15 533 96 998
Koy Wire am I ato Oo Gino 103 605 100 590 14 415 86 175
ICV iZ3Gh ig: Uo ath 56 6 & 134 589 112 5311 20 903 gi 628
TOAS pir a ens Hel ela 62 287 64 160 14 929 49 231
LO AA. sem beats 1 (2 alte 27 959 28 600? II 947 16 633
IVI G 5 5 GG Golo 9 < 23 516 24 220 13 643 10 557
TOA#O Sy pee eck ey oe en ee Fs 56 431 58 120 — —

y con su persistencia, indiferente del pasado, espera el momento propicio para
construir un nuevo penal, sobre las ruinas del séptimo presidio”’ (there had been
5 on Masatierra).

On January the 16th, 1935, President ARTURO ALESSANDRI signed a decree
making the Juan Fernandez Islands (together with Easter I.) a National Park, and
two German residents, Weber and CARLOS BOCK, both addicted to the study of the
fauna and flora, were appointed honorary forest inspectors. Bock soon died, and
Weber left the islands after some years. The regulations were strict; had they been
followed the goal for which I had fighted so many years, supported by Chilean col-
leagues, would have been attained. The rules did not infringe upon the reasonable
rights of the colonists, nor was the fishing industry affected. In order to collect
specimens of the indigenous plants and animals, also for scientific purposes, a license
was required. No permanent habitations could be erected on Masafuera and Santa
Clara. It is evident that Weber, in spite of his earnest intentions, did not quite
understand what effective conservation involves or he would not have written the
following words (p. 138): “Die Tierwelt von Juan Fernandez war leider sehr
sparlich, es sollte mehr Leben in die schweigsamen Walder kommen” — it did
not occur to him that the introduction of foreign animals, even a few birds, would
disturb Nature’s balance in a way never properly to be foreseen.

Unfortunately a law has little effect without adequate means to enforce it.
Without money and men with authority a conservation program cannot be realized,
and in the case before us there was neither. Serious inroad upon the forests
might, however, be prevented, and with the declining lobster industry and the
non-existent possibilities to expand agriculture there seemed to be no danger that
more people would settle on Masatierra. But what happened? Guzman’s interesting
account gives the answer, l.c. p. 37.

1 This figure, which is identical with the figure for 1940, must be incorrect. In all other cases
save one (1941) the average weight of the langosts slightly exceeded 1 kg, and it was very little
below that weight in 1941.

2 On p. 202 Guzman quotes a very different figure: 47 238 kg obtained during the period
Jan.—Aug. The balance cannot, I am sure, have been consumed locally, because the people
fish for the companies, so that some other explanation has to be found.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 185

“>

Fig. 99. The central part of Masatierra, seen from the air and showing the cultivations and

8- 99 Pp : ving

plantations around Cumberland Bay: from left to right V. Colonial, V. Anson and Quebr. del
Minero. — Photo B. Frédin 8/, 1952.

In 1940 the population amounted to 434 persons (225 men and boys), in
1943 771 (198 men; the number of women was 123, of children no less than 450);
1944 shows a drop to 653, a result, one would think, of the heavy drop in the
number of lobsters caught. In 1948 the population was about 600. The total
export of lobster, living and tinned, is shown in Table I (after Guzman).

As we see, the proportion of preserved lobsters underwent a rapid decrease
during these years; the export of live animals was undoubtedly more profitable.
The reasons for the sudden rise in total output in 1946 are not known to me,
nor if it indicated a reliable increase or just a transient improvement; after 1946
no figures have been available to me.

The growth of the population between 1940 and 1943 cannot have had
anything to do with the fishing industry, and the drop from 1943 to 1944 does
not run parallel with the quantity of lobsters; besides, the effect would not show
before 1945, when no census was taken.

More people meant more houses, more gardens, more timber and fuel used,
etc. The effect is shown by comparing figs. 24, 27 and 92 with 22, 30 and gg.
In 1917 the lower slopes of Q. del Minero were barren, in 1952 there were plant-
ations; Anson’s valley, where few people lived in 1917, bears sign of much
activity, and the aspect of the settlement in the Colonial valley has changed a
great deal from 1917 to 1952. Everybody welcomes that the devastated slopes
near the sea, where erosion is a serious problem, have been planted, nobody
objects to the growth of the population as long as fishing provides a decent

186 C. SKOTTSBERG

livelihood for the majority. This, however, is not the case. The time has come
when the old dreams to utilize the land, fostered by well-meaning patriots, are
coming true. The statistics tell us that Masatierra is being transformed into a
cattle and sheep farm, a new and strange type of National Park. In 1945 the
island had 5000 sheep, 600 head of cattle, 500 tame (?) goats and 300 horses,
and even if the horses and cows mostly graze the open country with grasses and
herbs, mainly introduced weeds, they do not despise the light forest which, and
this was easy to observe already in 1917, suffers. The habits of the sheep and
goats are different; wild sheep were observed by Weber in the thirties (p. 116).
There are no fences and nothing to prevent these animals to run wild among the
crests, where it is difficult to follow them. No palatable plant species is likely to
escape these gluttons and in contradistinction to the naturalist, they need no license.
It goes without saying that they will take to the mountains when, at the end of
the drier season, food becomes scarce. Guzman recommends introducing new kinds
of grasses, better adapted to the climate, a measure probably unheard of in combina-
tion with the management of a national park. It should not be forgotten that one
Mr. Orro RIEGGEL, “el gran amigo de las ciencias naturales’ as Guzman calls
him, celebrated the creation of the national park by introducing 6 pairs of rab-
bits (!) which, as everywhere else, will take what the sheep leave; nor that a parti-
cularly dangerous weed, the zarzamora (Rubus ulmzfolius), a wellknown pest on the
mainland, was introduced on purpose to be used as living fences. The success was
complete. The thrushes took care of the dispersal of the fruits, invasion is going
on everywhere, and I have just been told that the entire Plazoleta del Yunque pre-
sents the picture of an impenetrable “living fence’. From the same source I have
the information that the unlawful cutting of the Chonta has not been stopped —
I doubt that anybody tried to stop it — but is going on just as before and with
the same method: before a tree is felled a cut with an ax is made near the base
in order to know the thickness of the wood cylinder; if it is too thin to be of
value the tree is left standing, a potential victim of fungus attacks.

It remains to be mentioned that 17 colonists settled on Masafuera some
years ago (72 persons in all). Their principal occupation is said to be lobster
fishing.

Basing his opinion on his personal knowledge of the islands and on a wealth
of material from various sources, Guzman discusses the present situation and the
possibilities to improve it. There can be no doubt that he is interested in the
preservation of the nature, but it is also evident that he greatly underrates the
dangers jeopardizing the survival of the indigenous flora and fauna. The pros and
cons in agriculture and silviculture are set forth in detail. In his appreciation of
the fruit produced he differs considerably from Johow who did not hold a very
high opinion of the quality, and our impression was not too favourable. Oranges
and lemons, not grown in our time, are said to prosper in the valleys — this may
be true, but what are the fruits like? The wheat, of poor quality according to
Johow, is excellent, which must mean that new strains have been introduced. A
circumstance in favour of agriculture, Guzman says, is that as a national park all
the land belongs to the State, and the concessioned farmer needs not worry about

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 187

rents or amortizations. On the other hand, Guzman continues, the factors render-
ing agriculture difficult are also conspicuous. The men working for the companies
— there are three of them, but the Recart y Doniez Co. was taken over “‘en
estado ruinoso” by Cia Santa Sofia — earn much more money than the farm hands,
with the result that everybody prefers to be a fisherman. The schooners cannot
accommodate the export of fruit or vegetables. Good soil is rare and shallow,
because the islands are too young (!). The zarzamora is taking possession of the
best agricultural soil and is becoming a serious problem. The strong winds are
another inconvenience, to be overcome by planting hedges.

The most serious obstacle is the very limited extension of arable land, referred
to above p. 171; 300 hectares, in 9 different localities, including Masafuera and
Santa Clara, might be utilized, with an annual output of cereals amounting to
2300 quintales. If 800 are reserved for local consumption and 300 for seed-corn,
1200 remain for export (l.c. p. 160) to compete on the market with millions of
quintales produced on the mainland, and having to carry the cost of freight! It
it disgusting to find that such imaginary hopes have been carried from genera-
tion to generation. How much of the land is level? Almost nothing. And what
would happen if the slopes, where the inclination rarely falls below 15 to 20%,
and the soil is very thin, are plowed? The badly eroded areas seen everywhere
give the answer. When Guzman recommends the declivities for tree-planting he
gets on safer ground. And he believes that reforestation, large olive plantations
and the creation of orchards and artificial meadows would transform Masatierra
into an ‘‘alegre huerto isleno, deleite de los turistas” (l.c. p. 164).

Masatierra is already blessed with sheep, cattle, goats and rabbits, all in excess
—- will it also be necessary to add a host of tourists? Guzman holds a very high
opinion of the attractions of Juan Fernandez. He quotes “el célebre viajero Carlos
Rowsel” (I regret never having heard his name before) who once said that if
Masatierra were situated 60 instead of 360 miles from Valparaiso it would be “‘la
Reina de los Balnearios’” in the Pacific — and this in spite of the fact that there
is not the slightest trace of a bathing beach on the island. And when Guzman
compares Juan Fernandez to Hawaii and finds that the one is just as marvellous
as the other, he has lost all contact with reality. I should add that I happen to
be very well acquainted with both, and that nobody could appreciate the beauties of
Juan Fernandez more than I do. Call it a miniature Hawaii if you like, but deprived
of the tropical luxuriance, the colourful Royal history of the natives, the white,
palm-shadowed beaches of coral sand, the gigantic mountains and active volcanoes,
not to mention the comfort and luxury offered by the busy cities, the large hotels,
beautiful camp sites, excellent motor roads and easy and rapid communications
by sea and air with the outside world! The nature of the indigenous living
world is, in principle, of the same type in the two cases, but incomparably richer
and more varied in Hawaii — there is room for half Masatierra in the caldera of
Haleakala. To the average tourist Masatierra remains Robinson Crusoe’s island
— this is its main attraction. I have no doubt that the traffic will grow and the
tourists become a moderate source of income which nobody will envy the kind and
hospitable colonists, but I am afraid that Guzman is too optimistic when he thinks

188 C. SKOTTSBERG

that the day is near when Juan Fernandez will become “el balneario habitual de
los sonadores del mundo” (p. 229) and when large passenger planes will land on
the big aerodrome on Masatierra. At present not even a tiny plane is able to land
without crashing, but there are some places where an autogiro could come down
safely. The roadstead in Cumberland Bay is so far the only place where a plane
(a Catalina) has alighted, but with a high wind and a heavy swell the situation
gets unpleasant. The level land by the sea in front of the colony is the only
place where a small airport could be constructed. Its length will be 500 m or, if
the front slope of Cordén Central with the old prison caves be blasted, perhaps
800 m, but the buildings belonging to the fishing companies and a good many
other houses would have to disappear. On the table-land of Masafuera, about
1200 m above sea level, another airport could be built at enormous cost, another
one, probably a little cheaper, on the Loberia plain. Neither would serve any
sensible purpose if not a strategic one. Let us limit our plans of making Juan
Fernandez a popular goal of the tourist to an improvement of the communications
and to a couple of modest guest-houses — and to impress on the visitor that
he finds himself in a sanctuary where he has to keep his hands off. An unknown
number of plants and animals barely manage to hold their own, and obviously
many are on the verge of extinction, pronouncedly stenotopic as they are. The
trail to Portezuelo ought to be improved and kept in repair, for everybody will
want to see Selkirk’s Lookout, to read the memorial tablet and behold the grand
views. And there is no point within easy reach where the endemic flora is — or
was, at least, in 1917 — better displayed.

: *
6

If we want to preserve a unique living world of very great scientific interest
and as such belonging not to a single country but to the whole world, these are
the rules:

to limit plantations and fields to the waste-land on the north side of Masa-
tierra;

to encourage gardening for local consumption;

to declare war on the introduced noxious weeds, goats and rabbits;

greatly to reduce the number of domestic animals and to keep them out of
the native forest;

to reduce the number of wild goats on Masafuera and keep it on a minimum
or, which would be the best, to exterminate them;

to teach the inhabitants not to disturb Nature’s equilibrium;

to enforce the Law of Jan. 31, 1935, by appointing a sufficient number of
salaried supervisors and guardians.

Human influence has cut its mark deep and it has changed the natural scen-
ery greatly without adding to its beauty. In part this has been inevitable, if man
was to live on the islands, but there has been and is too much senseless destruc-
tion. This is the more to be regretted as the welfare of the population need not
at all depend on either breeding cattle or sheep-farming. Once the lobster meant

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 159

i eat il

wae’
Ren i

|
i
4

Fig. roo. Large specimen of bacalao, Cumberland Bay. The gentleman is Don NATALIO SAN-
CHEZ, subdelegado civil and the highest authority in the islands at the time of our visit. —
Photo K. Backstrém 1917.

Igo C. SKOTTSBERG

everything. To-day we can speak with Guzman of the “‘Crepusculo de la langosta”’.
The situation is alarming, he calls for measures to put an end to the decline, and
he proposes certain ways and regulations. I would like to add that in a case as
serious as this the best thing to do is to proclaim the whole year a closed season
during a sufficiently long period. An industry based on the enormous supply
of fish, with the bacalao (fig. 100) heading the list, would, if carefully handled,
become a profitable enterprise. A ship with modern equipment would bring the
frozen fish to the mainland where it would find a ready market. The new methods
of cold storage permit deep-frozen food, fish not excepted, to keep absolutely
fresh during any length of time. Beside the bacalao, the most abundant of all and
considered to be the most delicious, there are several other kinds of commercial
value unknown on the coast of Chile. I shall add one more remark. Cumberland
Bay ought to be an ideal place for a biological station. The fauna and flora,
terrestrial as well as marine, offers a wide field for scientific research, no less
important from a practical viewpoint.

The present management of these precious islands is not in good accordance
with the intrinsic meaning of the Law and, in some cases, directly violates it.
If the responsible authorities do not change their attitude, Juan Fernandez will
become a second Saint Helena and a disgrace to an enlightened world.

Speaking as on behalf of the Chilean nation, I would like to say, with a
slight alteration, what Guzman said about the threatened langosta (p. 223):

No permitamos que nuestras islas se transformen en el recuerdo de una
riqueza extinguida en forma irreparable. Asi lo exige nuestro prestigio de nacion
civilizada, asi lo esperan de nosotros las generaciones futuras.

December 1953.

Bibliography.

ANDERSON, G. W. A new, authentic and complete collection of voyages round the
world. London. No date. (Byron’s and CarTeret’s visits to Juan Fernandez.)

Anson, G. A voyage round the world in the years 1740—44. Compiled by RicHarRD
Water. London 1749.

Brancut, FE. C. La Isla de Robinson. Valparaiso 1922.

Brtccen, M.J. Fundamentos de la geologia de Chile. Santiago 1950.

Burney, JAmes. A chronological history of the discoveries in the South Sea or Pacific
Ocean. London. Vol. II (1813, L’Heremite), IV (1816).

BurGer, O. Die Robinson-Insel. Leipzig 1909.

Ermet, A. Eine Reise nach der Robinson-Crusoe-Insel. Hamburg 1889.

Fropin, B. Expedition pa svenskt uppdrag kartlade sagoén i Stilla havet. (Expedition
on Swedish initiative mapped the fairy isle in the Pacific Ocean.) Dagens Ny-
heter 72/, 1952.

Grar Marin, A. Problemas economicos de Juan Fernandez. Departamento General
de la Produccioén. Oficina de Enlace Agricola. Memorandum 158. Santiago,
29 de Abril de 1944. (Typewritten.)

GrauaMm, Marta. Journal of a residence in Chile during the year 1822. London 1824.

Guntuer, E. Derrotero de la costa de Chile. V. Anuario Hidrografico de la Marina
desChile: 32. 1920.

GuzmAn Parana, J. Cumbres ocednicas. Santiago. No date.

A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS Ig!

HacerMan, T. H. Beitrige zur Geologie der Juan Fernandez-Inseln. Nat. Hist. of Juan
Fernandez and Easter Island. 1: 1. 1920.

HawkeswortH, J. An account of the voyages undertaken ... for making discoveries
in the Southern Hemisphere. I. London 1783. (Byron's and CarTERET’s journeys.)

Instrucciones nduticas de la costa de Chile. Anuario Hidrografico de la Marina de
Chile. XX. 1896.

Jouow, F. Estudios sobre la Flora des Islas de Juan Fernandez. Santiago 1896.

Juan, G. y ULLoa~, A. DE. Relacién historica del viage a la America meridional. Pt. 2,
vol. III. Madrid 1748.

Lopez, JuAN E. Esploracion de las islas esporddicas de la costa de Chile. Anuario
Hidrografico de la Marina de Chile. II. Santiago 1876.

Puiuipp1, R. A. Geognostische Beschaffenheit der Insel Masafuera. Neues Jahrb. fiir
Mineralogie etc. Stuttgart 1857.

Puiippson, A. Die Erosion des fliessenden Wassers und ihr Einfluss auf die Land-
schaftstypen. Geogr. Bausteine herausgeg. von Dr. HERMANN Haack. Gotha 1914.

PoEHLMANN, R. Noticias preliminares sobre las condiciones jeograficas i jeoldjicas del
Archipiélago (de Juan Fernandez). In: Jonow, Estudios.

QuensEL, P. (1) Die Geologie der Juan Fernandez-Inseln. Bull. Geol. Inst. of Upsala.
RE 1912.

—— (2) Additional Comments on the Geology of the Juan Fernandez Islands. Nat.
Hist. of Juan Fernandez and Easter Island. I: 3. 1952.

Skortsserc, C. (1) The Wilds of Patagonia. London rort.

—— (2) Till Robinson-6n och viarldens ande. Stockholm 1918.

—— (3) The vegetation of the Juan Fernandez Islands. Nat. Hist. of Juan Fernandez
and Easter Island. II: 29. 1953.

Surc.irre, Tu. (1) Sixteen years in Chile and Peru from 1822 to 1839. London 1841.

—— (2) Crusoniana or Truth versus Fiction, elucidated in the history of the Island
of Juan Fernandez. Manchester 1843.

Tenz, O. Una ascensién emocionante a la cumbre del Monte Yunque. El Mercurio
8/, 1921. Valparaiso.

Vipat Gormaz, F. Jeografia ndutica de la Reptiblica de Chile. Anuario Hidrografico
de la Marina de Chile. VII. 188r.

Viet, O. Islas de Juan Fernandez. Anuario Hidrogrdfico de la Marina de Chile. IV.
1878.

WALTER, R. See ANSON.

WaLpoLe, F. Four years in the Pacific in Her Majesty's ship “‘Collingwood’’, from 1844
to 1848. London 1850.

Weser, Huco. Signalmaat Weber. Zehn Jahre auf der Robinson-Insel. Reutlingen 1940.

Addition.

This paper was in page proof when my friend Mr. G. Looser sent me a report
written by Dr. Arperto Grar Marin, Head of the Oficina de Enlace Agricola in Sant-
iago, and kindly put at my disposal (see Bibliography). Dr. Graf Marin visited Masa-
tierra in January 1944 in order to inform himself of the economic situation of the
population, 452 persons in 1943, a figure very different from the one quoted by Guzman
for the same year, 771; both cannot be correct. Graf Marin judged the living condi-
tions to be comparatively good, even if much remained to be improved in the way
of housing, teaching, medical care, entertainment, travel facilities, etc. It goes without
saying that lobster fishing gave the main income; again, there is a discrepancy be-
tween his figures and those of Guzman: 84 100 lobsters shipped to the mainland in
1943, whereas the total catch was 62 287 according to Guzman. The average annual

192 C. SKOTTSBERG

profit derived from the export of wool was about 1/, of the income from fishing, the
number of sheep 3000. Finally, some mutton was sold to Cia Oto Hnos, apparently
the only fishing enterprise in operation (comp., however, p. 187). With regard to the
lobster Graf Marin was told that it breeds the year round whence follows that the
fears later expressed by Guzman would seem groundless and a closed season — after
1931 1/6—30/9 — umnecessary.

Although Graf Marin recognizes the fishing industry as the main source of wealth
he revives the old dream of Masatierra as a future rich farming country, and his plans
were drawn up without regard to the fact that Masatierra forms part of a National
Park, protected by a law which, nevertheless, he does mention in passing. He recommends
to import farmers from the region of Concepcion, where the climate is similar. As
the soil, save for the houses and gardens, is fiscal property, anybody is entitled to
supply himself with wood in the forest — only maqui is used for fuel, however. The
forest should be subject to economic management by multiplying the few good timber-
trees and by introducing new ones; the Mediterranean cork-oak is particularly recom-
mended beside Acacia melanoxylon and Pinus radiata. Of the endemic trees two are
(or rather were) of commercial value, the chonta palm and the sdndalo; the former —
and also the latter, if it could be rediscovered — ought to be propagated in nurseries
and planted on a grand scale. He tells us that the last living sandal tree seen by
Johow in 1892 and by me in 1908 was ordered by the Subdelegado VeERA in 1918
or 19 to be cut down — but in 1916 I was assured by the same man who brought
me to the tree in 1908 that it had died and the wood been taken care of. I have
no reason whatsoever to doubt the correctness of his statement.

With regard to the black pest Avfennaria (Limacinia) Graf Marin shares my
opinion that Johow exaggerated its dangerousness. A more serious enemy of the luma
is an insect, Sazssetia oleae, and he recommends the introduction, successfully arranged
for on the mainland by himself, of its parasites.

Even if various fruits can be produced in sufficient quantity to be exported —
the water-supply is good and the streams might be canalized and used for irrigation —
raising live stock should constitute the main occupation of the farmers, but it is
jeopardized by the spread of the zarzaparilla (Acaena argentea) and of the zarzamora
(Rubus ulmifolius). They ought to be exterminated, which is just as desirable from
the naturalist’s viewpoint, but much easier said than done. In order to improve the
pasture alfalfa and new and more nutritious foreign grasses should be introduced. Above
all, to stimulate general utilization of the land and to prevent that a single thrifty
person take possession of most of it, the island ought to be parcelled and the size
of the lots fixed with regard to their rentability. The total value of Masatierra, settle-
ments not included, is said to be 220000 pesos, of Masafuera 120 000 (considerably
more to-day that 1o years ago). From a scientific and scenic viewpoint the value of
these islands cannot be expressed in figures — it is inestimable. If the plan set forth
in Graf Marin’s interesting report materializes, Juan Fernandez will stand as a unique
example of a National Park, the home of a unique fauna and flora, offered for sale.

THE NATURAL HISTORY
OF JUAN FERNANDEZ
AND EASTER ISLAND

BDLITEDABYVS DR. CARL: SKO LPS ii tee

VOL. I
GEOGRAPHY, GEOLOGY,
ORIGIN OF ISLAND. LIFE

PART III

5. C. SKOTTSBERG: Derivation of the Flora and Fauna of Juan
Fernandez and Easter Island.

Part III completes vol. I.

UPPSALA 1956

ALMQVIST & WIKSELLS BOKTRYCKERI AB

ww.

5. Derivation of the Flora and Fauna of Juan Fernandez
and Easter Island.

By
C. SKOTTSBERG.

With 1 Map.

Part I.

THE JUAN FERNANDEZ ISLANDS.

Chapter I.
Composition, distribution and relationships of the Flora.

The statements made below are based on the results laid down in vol. II of
this work and in a number of papers published from time to time prior to the
survey in the field undertaken by myself and collaborators during a three months’
stay in the islands, Dec., 1954—March, 1955. Later on some minor changes
will have to be made in the lists of the Vascular Plants, but as they will not alter
the conclusions drawn I have preferred not to include them here, more so as the
species in question, some of them at least, have to be subjected to a close taxo-
nomical study. With regard to the Cryptogams, to be referred to a number of
specialists, the revision of the collections will take some time, but even if a few
new species will be described, and other additions made to the lists, the pro-
portions between the various geographical elements will, I think, remain much the
same.

1. Angiospermae.

In the islands 42 families are represented of which one, Lactoridaceae, is
endemic and monotypic. Until recently Juan Fernandez (Masatierra) was the only
oceanic island possessing an endemic family, but after the discovery of the genus
Degeneria, which forms the monotypical family Degenerzaceae, it shares this
honour with Fiji; the Fiji group is, however, not as ‘oceanic’, in the current
sense of this word, as Juan Fernandez.

The largest families are Compositae with 28 (13 gen.), Cyperaceae with 14
(7 gen.), and Gramineae with 13 species (10 gen.), but these are very large fami-
lies almost everywhere; Campanulaceae (2 gen.) and Juncaceae have 6 (2 gen.,
but some species of Fucus may not be truly indigenous), Rubiaceae 5 (4 gen),

13 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

194 C. SKOTTSBERG

Umbelliferae 5 (3 gen.), Chenopodiaceae 4 (2), Myrtaceae 4 (4), Piperaceae 4 (1),
Rosaceae 4 (3), Solanaceae 4 (2), and Urticaceae 4 (3), Caryophyllaceae 3 (2),
Cruciferae 3 (1), Gunneraceae 3 (1), Halorrhagidaceae 3 (1), Berberidaceae 2 (1)
Bromeliaceae 2 (2), Convolvulaceae 2 (2), Labiatae 2 (1), Leguminosae 2 (r1),
Plantaginaceae 2 (1), Rutaceae 2 (1), Scrophulariaceae 2 (2); represented by 1
species only: Aizoaceae, Boraginaceae, Callitrichaceae, Empetraceae, Ericaceae,
Euphorbiaceae, Flacourtiaceae, Iridaceae, Lactoridaceae, Loranthaceae, Palmae,
Ranunculaceae, Rhamnaceae, Santalaceae, Saxifragaceae, Verbenaceae, Wintera-
ceae, 17 families or 40 % of the entire number. Some of the families are alto-

gether small, but others are large and widespread also in Andean America—
as, for instance, Umbelliferae-Hydrocotyloideae (Azore//a), Caryophyllaceae, Cruci-
ferae, Labiatae, Leguminosae (Adesmia, Cassia), Plantaginaceae, Scrophulariaceae
(Calceolaria), Boraginaceae (Plagiobotrys and related genera), Euphorbiaceae, Iri-
daceae (S¢syrizchium), Ranunculaceae, Rhamnaceae, Verbenaceae. The absence
of such families as Amaryllidaceae, Asclepiadaceae, Fagaceae, Geraniaceae, Lau-
raceae, Liliaceae, Nolanaceae, Onagraceae, Orchidaceae, Oxalidaceae, Polygona-
ceae, Portulacaceae, Valerianaceae and Violaceae is noteworthy, being more or
less well, some of them very well, represented on the Chilean mainland.

Of the 42 families Masatierra possesses 39, seven of these also found on
its satellite Santa Clara, and Masafuera 30; 12 are confined to Masatierra (Plan-
taginaceae also on Santa Clara) and 3 to Masafuera; 27 are common to Masa-
tierra and Masafuera. The families are, in contradistinction to the genera and
species, evenly distributed over the group, as shown by the list of Genera and
Species, Dable I.

The number of genera is 89; of these 71 are found on Masatierra, 9 on
Santa Clara and 54 on Masafuera; 35 are known from Masatierra only, including
those known from Santa Clara, all also found on Masatierra; 18 are confined
to Masafuera, 36 shared by both islands. Expressed in percentage: Masatierra
39.3 %, Masafuera 20.2 %, Masatierra + Masafuera 40.5 %. Thus less than one
half of the genera are common to both islands. The floristic difference between
them is partly explained by the difference in altitude and thereby in climate—
Masafuera has an alpine and subalpine flora for which there is no room on
Masatierra, partly by the more varied topography of this island, which has a richer
flora. The difference becomes particularly obvious when endemism is considered.

Of the 89 genera, 17 (19 %) are endemic; of these 12 (70.6 %) are confined
to Masatierra, 1 (5.9 %) to Masafuera and 4 (23.5 %) found on both islands; see
Table II.

Of 71 genera known from Masatierra 16 (22.5 %) are endemic to the islands,
of 54 found on Masafuera, 5 (9.3 %); of 35 genera only found on Masatierra, 12 are
endemic (34.3 %), the corresponding figures for Masafuera are 18, 1 and 5.5 %
and, for the genera occurring on both islands, 36, 4 and 11.1 %. These figures
serve to illustrate the great difference in the distribution of the endemic genera
commonly looked upon as representing the most ancient element among the
Angiosperms in Juan Fernandez.

Of the total number of species, 147, 99 are found on Masatierra, 9 on Santa

DERIVATION OF THE FLORA AND FAUNA

Table I.

Endemics bold-faced, endemic genera in capital letters.

_Abrotanella crassipes Skottsb. .

Acaena ovalifolia Ruiz et Pav.

— masafuerana Bitter

Agrostis masafuerana Pilger .
Apium fernandezianum Johow
Azara fernandeziana Gay

Berberis corymbosa Hook. et Arn.
— masafuerana Skottsb.
Boehmeria excelsa Wedd.
Callitriche Lechleri (Hegelm.) Fassett
Calystegia tuguriorum R. Br. .
Cardamine chenopodiifolia Pers. .
— flaccida Cham. et Schlechtd. .
— Kruesselii Johow

Carex berteroniana Steud.

— Banksii Boott

CENTAURODENDRON dracaenoides Johow

Centella triflora (Ruiz et Pav.) Nannf. .
Chaetotropis chilensis Kunth

— imberbis (Phil.) .

Chenopodium crusoeanum Skottsb.

— nesodendron Skottsb.

— Sanctae Clarae Johow .

Chusquea fernandeziana Phil.

Cladium scirpoideum (Steud.) Benth. et Hook. f.

Colletia spartioides Bert.

Coprosma Hookeri (G. Don) W. R. B. Oliver .

— pyrifolia (Hook. et Arn.) Skottsb.
CUMINIA eriantha Benth.

— fernandezia Colla

Cyperus eragrostis Lam.

— reflexus Vahl

Danthonia collina Phil.
DENDROSERIS litoralis Skottsb.
— macrantha (Bert.) Skottsb. .

— macrophylla D. Don

— marginata (Bert.) Hook. et Arn.

Masatierra

Santa Clara

195

Masafuera

196 C. SKOTTSBERG

Dichondra repens Forst.

Drimys confertifolia Phil.

Dysopsis hirsuta (Muell. Arg.) Skottsb.
Eleocharis fuscopurpurea (Steud.) H. Pfeiff.

Empetrum rubrum Vahl

| Erigeron fruticosus DC. .
| — Ingae Skottsb.

— Innocentium Skottsb.

— luteoviridis Skottsb. .

— rupicola Phil. .

— turricola Skottsb.

Eryngium bupleuroides Hook. et Arn.
— inaccessum Skottsb. .

— sarcophyllum Hook. et Arn.

| Escallonia Gallcottiae Hook. et Arn.

| Euphrasia formosissima Skottsb.

Fagara externa Skottsb. .

— miayu (Bert.) Engl. .

Galium masafueranum Skottsb. .
Gnaphalium spiciforme Sch. Bip.
Gunnera bracteata Steud.

— Masafuerae Skottsb. .

|— peltata Phil.
| Halorrhagis asperrima Skottsb.

}— masafuerana Skottsb.

— masatierrana Skottsb. .

Hedyotis thesiifolia St. Hil. .

Hesperogreigia Berteroi Skottsb.
HESPEROSERIS gigantea (Johow) Skottsb.
JUANIA australis (Mart.) Drude

Juncus capillaceus Lam.

— dombeyanus Gay .

— imbricatus Laharpe

— planifolius R. Br.

— procerus E. Mey. .

Koeleria micrathera (Desv.) Griseb.

|} LACTORIS fernandeziana Phil.

| Lagenophora Harioti Franch. .

Libertia formosa Grah. .

ota Ue | 22
sneer &
s| 3 |
op)

4 +
+ - ot
) |
_>
= - +
+ ~ +
= = 35
= - +
= — +
= - +
= — 4
Reed

- = -
= = +
= = +
= = +
= = a
= = +
+ = ==
= = +
= = +

Pe ee |
— = +
reese |p
|)
¢ he Ae
| ae
ae
fees eer
, oe
= = +
+ - E

DERIVATION OF THE FLORA AND FAUNA 197

— O =
Pretselia ailleneay, IPSS oy” pCR Ac OR oe, Oa leew et at ACen geen eevee | = +
inigattlaytiASALICLANA SKOLLGD Spe. cy 4) sy ee ck ee es “ae es ws ey ee, ws NS ~ --
PMacoynicanpus disynus (Bitter) Skottsb. . . . . . - . . . 2 5.2 flees 4 = =~ |
~MEGALACHNE berteroniana Steud. |
|— masafuerana (Skottsb. et Pilger) Hubbard ms. ..........) — ; + |
| Vite ISee LAD TALUS PEL. ss ES TOnMAr 6 i) ty su ev ell eno) s,s) atte ee hte Be - = + |
| Mrraenventan Ss CUULZeCL | OHOWi a lel certs ahs) ou ck oso) a = © cele) SSO e - ~
Beier amiiimicatlatian( POI.) Bers os 53 2 a ew I ee — =
NegteLame EAN aACensis) (stills) MOruCe Vatet- seo = oy oe ns ue mc, OS -- — 3
ieaianagCOLGirolane nil sp. yic a ota cee ae Aine felt) — -
| NOTHOMYRCIA fernandeziana (Hook. et Arn.) Kausel ..... . +. - -
O@EAGAVTACelesansyP Dis 5 0) 08 Fen ee ce ok ok we ee ne oe SE + -- -
| Oreobolus ObmsanomlusnGant Ghee they) ot ilm ace fev ev: ea fone Mohs) 2h ce 3 clea — - +
Ee atk Gde Dial AMUNLITENUE LES Ue NACI sung ce ce chats) tea ea toy ee en Wai ies PMTs tee) Pe ee — — 2"
Pi aLory Chitane MiNENSISHD Css a2, von ney eit) tok 3, Ni Vol pied ey pce Slot oes iar on ep - — =
Peperomia berteroana Miq.
CRM ALCL eZ TAT am VIG a tsss, 3) cy 12, tee Poe Ge cich em 2p oh vom ep Su oi. ay Gayton vo | Shes + = +
| — margaritifera Bert.
= Skottsbergii C. DC.
PERLE 2) Teo FD Cs So aio auone irae Gai ae eM - = oF
| PHOENICOSERIS berteriana (Dene) Skottsb............-/ # | = | -
| — pinnata (Bert. ex Dcne) Skottsb. |
|— regia Skottsb. .
Pehayetanthus Berterot (Hook, eb Arm))\Reiche = 2's. 2 37. 3 3 3 3 - = =
| FipLoecncetmimpicolors (Vall) Preslgitawss - Geel 2 a sf) shiek (eu reas ~ = =
ania oOrernand eZiavbenc tury ey, Elecite. ee ieh ta ee ici oy ce fel sive sene a5 = ae |
UI CAA CHAM MaVAL YE <A sel a epee etiam ope ol ce ese) to [Peavas ek Ge es - + =
SPGHOPLHORUS bromiocidés Phils. sys oe! Amos 2) eee
GAIN CUMS CAPTALUMIPSKOELS DS vase chest eh ches) “eck Se eyo ec) ot oy Po | at
Eb wanicranths Beri ex Denes 8 8)..2 Ry Re ee wn, SES =
me trifolial Berti /exwener iy @ IDS. i es. 2. le - as,
= priinata (johow) Skottsb is 2b57.. 2a. ea So eee =
Rhaphithamnus venustus (Phil) B.L. Robins... ...-- +--+ =| F | = ‘icky
RHETINODENDRON Berterii (Dene) Hemsl. ........---/ + = — |
BOBINSONTACeveniay Paint) 2Rs ee. eh Fe ol ee
| — gayana Dene Slash Sg) oat th a al a Silenesre | | =
|— SEACHISUDEH EAL ee Ft eee en es SE Se eels Mom one + = ie
| — INEASHEMCT AG SKGOLESDAr. Reh edness eee ha tia as ne Mar OL marie 8 See oe | = = a5
P=Wiurifers Dead Lae ey ey Pee

198 C. SKOTTSBERG

g] bie

a nN a
Rab USTs COLES, S mas Me aM ass. ... 00s 0s a ea, fe | +
salicomia druticosas Km iyae mens. eS... os) Soars aoe. a late [0 cst. Wane +
Santalum fernandezianum F. Phil... .......2.2....., : Ste = +
Scinpusicenmuns pall MO, 54: sk, . <. . Eaneea oh Bato | + = =F
pe OdOSUS PRO Lt ep yeree Mears. - c.f o(c, 2. veakse ede ed £ Pog ee | ate e y
SELKIRKTA Berteroi (Calla) Mems|.. . . . . .. . sn ac A ee
Solanum@fermandezianumpPhil, ©... 1. 2s ns en ee) jatectael ees pane
== duasafueranmmuyBiteeret Skottsbs . : ss . i na cs, Quel eels ea aii
a= RODINSOnianUmMSBIEEeh eG a cvs a ws Ree ee ates | rae | |
Sophora fernandeziana (Phil.) Skottsb. ............... See | —
ey amlasatueranay(Pbal))pSkOttsb....., fis s<i.) 2.4 Ge cee ane =. eS cee |
| spetgulanajcomfertiflora,Stend. .. . .......... mee tanh, AN ||) = ar a
meet RAS ABUICT ANAT SOLES: ys. Svs. ss wks Aso ae oe = iS +
Stipa (Nassella) laevissima (Phil.) SPER? 28. oo Ast eat eae eae ee ae | = af
pee DCesiario min myetUR UP tess 506, s,s a! arc? cds wy GY rth =e | = hs
SYMPHYOCHAETA macrocephala (Dene) Skottsbs. = 47-0. Ne SEE Lo = =
ac Eea Bon vexpansa MUEE 28 saya x ty eva gb. Geis nn ote +
drisetum chromiostachyum Desv. ............ Syste es weak ae tel =
Went Selkiridi (Hook etArm,)-Berg . . . oe a pe ee) eee -
Wncimia brevicaulis Thouars s, .. 2... ..... .. 2 Bis ert) tan cal ata
mmc OSCAEACUCKeD th ga ean tice 2 ns. WOE Ny aceon Or
=e WOUSTASIIUBOOttises 05) co en se sare Nem sci, Megmek ee eevee ie Bes | ert | — ar
=== JOUUSONGIOS PGE 5S Gate ce) Oe a re ne en oe Pecceait — | = “ts
CCS RE OS DDS Mat esac coe ot sos! a, Gh ARE Pale ; Sit tet ets
Urtica fernandeziana (Rich.) Ross ........,...... atte ae + 4) aie
2 ME SEUIVOTAS EN ©” Ose a re ae Sa he +
Wahlenbergia Berteroi Hook.et Arn. .......... sid acinaculiok alae “eat wee = |
m= tetnandeziana (ADC. p.p.) Skottsb. . ...... 0.0 sRERERANISN MY SEAS HRS ee
—= (GIB iR rey Sikes ey EN sie oven Se ree
mean rAlmiig(Colla)sSkottsbasw awe et. exc. ss 2 nc kus ce wwe op =
mee asaimerae (Phil)pSkottsbe vs 6. ss vb ee peiwa yt = = a
VEODSIGLOIEAS 7USs CHEE Sol 108) cess a ea rr Oe)

Clara, and 74 on Masafuera; 68 = 46.3 % (68.7 % of 99) are restricted to Masa-
tierra, I = 0.7% to Santa Clara, and 47 = 32 % (63.5 % of 74) to Masafuera,
4 = 2.7 % are common to Masatierra and Santa Clara with the exclusion of Masa-
fuera, 4 = 2.7% found in all three islands and 23 = 15.6% on Masatierra and
Masafuera excluding those also occurring on Santa Clara. Thus only 27 species
= 18.4 % are reported from both Masatierra and Masafuera, a surprisingly small

DERIVATION OF THE FLORA AND FAUNA 199

Zable L1.

Distribution of the endemic genera.

Masatierra
Santa Clara,
Masafuera

(Gvalcchingeyelyavchroval GG ka CaaS nS ee a ee { re = -
Cuminia
Dendroseris
Hesperoseris
TET a CRN Se LO a ee Gey ey oo ee me —
Te AGCORISWE CE PIB Ae Ut calcite Wolect a ee 1 ike chon, Wks ak TA oF _

MEN EVeIoNAES = og “a. ole Soh lO! GS am EGkhon Ute lon emt = +
INGEHOMIVECIAt NSM cM iagr) Wl benwtbe serene + acy wou oo ye we oily hex --
Ocharaniane- mets artes encoten fe tsg8e Gy fe dered cz:|| Sent - -
IPNOEMICOSChISH AN SMNECIA ST Te dics Shiciteats Gaalie’ Stl + | = +

Podophorus

IRVEES “ce a! = Sg mae id Mia a oho, heh ta ene nee ns oe ae —
betin oclenclromy cmos © ckatia Mwai a ane hea calc) ey oss se lee
RODINSOMAM shower ctary pases Mak tomodecas hoe! 2 Ess ye Sigel does + — 4e
Selina terns flechette Meuse at ‘sey hell, ve) Shea ahs + Sa | lle ic
Symphyochaeta .

SEA STEG]IMEE IS Se LE ROAR Clee ce ec ne | —- | =

number, for even if we pay due attention to the physiographic difference between
them, it tells a story of effective isolation with little possibility for an exchange
across the 92 miles of water separating them.

Of the total number of species (147), 101 are endemic = 68.7 %; of the 99
species found on Masatierra, 66 (66.7 %) are endemic in Juan Fernandez; the cor-
responding figures for Masafuera are 74 and 47 (63.5 %), and for Santa Clara 9
and 5 (55.5 %). Thus endemism plays about the same role in all cases, and this
is apparent also when local endemism is considered. Of the 68 species restricted
to Masatierra 50 (73.5 %) are local endemics, of the 47 species restricted to Masa-
fuera, 34 (72.3 %); the single species restricted to Santa Clara is endemic. Of the
27 species common to Masafuera and Masatierra 13 (48 %) are endemic in Juan
Fernandez. If, to the 68 species restricted to Masatierra, 4 also found on Santa
Clara are added, the figures are 72 and 73.6%. Of the total number of endemic
species (101), 50 are, as we have seen, confined to Masatierra (49.5 %), 3 (3 %)
to Masatierra + Santa Clara, 1 (1 %) to Santa Clara, 34 (33.7 %) to Masafuera,
12 (11.9%) common to Masatierra and Masafuera, and 1 (1 %) found on all three
islands. The difference between Masatierra and Masafuera stands out even more
clearly. It is less pronounced when we come to the 46 non-endemic species: only
on Masatierra 18 (39.1 %), on Masatierra and Santa Clara 1 (2.2 %), only on

200 C. SKOTTSBERG

Masafuera 13 (28.3 %), on Masatierra and Masafuera 11 (23.9 %), and on all three
islands 3 (6.5 %). However, several of the species restricted to Masatierra are,
perhaps, not truly indigenous, whereas the indigenous character of the species
only recorded from Masafuera cannot be doubted; 11 of them are mountain plants.
With regard to the former, some were commented upon by JoHOw. In his table
“Continjente B, Especies autéctonas, pero no endémicas’’, pp. 221-222, he men-
tions several species regarded as doubtful natives: 3 species of Guaphalium, Mimu-
lus, Solanum furcatum (= robinsonianum), T7etragonia expansa, Spergularia rubra
(= confertiflora), Parzetaria humifusa; Trisetum chromostachyum and Danthonia
collina. With the exception of Guaphalium spp. I have listed them as native.
There seems to be little reason to regard 7e/ragonia, a wide-spread thalassocho-
rous plant, as anthropochorous, and the Spergudarza is known only from Juan
Fernandez and Desventuradas Is. Danthonia gave me the impression of being
just as autochthonous as P7p/ochaetiuim and the two species of S@pa, and I find no
good reason to exclude either MWimulus, Parietaria or Trisetum (this never seen
by us). With regard to Solanum “furcatum’’ (not furcatum of DUNAL) I share
Jouow’s doubts. It was described by BITTER as S. robzusontanum, a Morella micro-
species of S. xzgrum \.. coll. and found on all three islands. This assemblage
as represented on the mainland has never been seriously studied. Possibly S. vodzn-
sonianum is a case of the same kind as the dandelion discovered on the islands
in 1917 and described as Z7araxacum fernandestanum Dahlst., a microspecies of
the Eurasiatic Vulgaria. There cannot be the slightest doubt that it is an alien
introduced from Chile, where these weeds have not been studied. From JOHOW’s
list I have excluded Bahia, Amblyopappus, -rythraea, Monocosmia and Phataris.
Further, some species not recorded from Juan Fernandez by JOHOW or earlier
authors, but for the time being listed as native by me, are under strong suspi-
cion: Funcus capillaceus and planifolius, Centella and Hedyotis, perhaps also Paro-
nychia. If, in the future, we shall be able to purify the list, the percentage of
endemic species will rise to 72% or even more.

The genera richest in species are Lrigeron with 6, Funcus, Robinsonia, Unct-
nia and Wahlenbergia with 5, and Dendroseris and Peperomia with 4 species each;
eight genera have 3, 15 two, and 59 only one species. The average number of
species to a genus is 1.65.

The systematical position of the endemites and the distribution of the genera and
species also found elsewhere.

Gramineae.

Stipa L. About 250 sp. (BEWS), widespread both hemisph., subtrop—temp.
neestana Trin. et Rupr. Mex. to Boliv. and Chile, Braz., Argent., Urug. Poly-
morphous (249. 771).
laevissima (Phil.) Speg. Pert, Chile, Argent. With a number of S. Amer. sp.
referred to a separate genus (Vasse//a).

DERIVATION OF THE FLORA AND FAUNA 201

Piptochaetium Presl. 20; N. Amer. to extratrop. 5S. Amer.

bicolor (Vahl) Desv. Chile: Valdivia; Braz., Argent.
Podophorus bromoides Phil. In 227 I referred to Brachyelythrum Beauv. and Aphan-
elythrum Hack. as the nearest relatives. Mr. C. E. HUBBARD kindly informed me
that it resembles the former in a number of important features and that he regards
both genera as belonging to a relatively ancient group of grasses. The distribution
of Brachyelythrum is disjunct after a well-known pattern: 2. erectum (Schreb.)
Beauv., N.E. Amer., and &. japontcum Hack. ex Honda, China, Korea, Japan.
PILGER (798) brings this genus to Festuceae-Festucinae next to Aphanelythrum,
placing Podophorus with the Brominae.
Chaetotropis Kunth. 2 (23).

imberbis (Phil.) Near the following.

chilensis Kunth. Pert, Chile, Braz., Argent., Urug.
Agrostis L. About 125; very wide-spread in temp. and cold climates.

masafuerana Pilger. ‘“‘Gehort in die Verwandtschaft von A. canina L. In der
Tracht A. magellanica Lam. sehr ahnlich” (PILGER 797). This is recorded from
Magell., Falkl., N. Zeal. (55). F. B. H. BRowN (35.1. 84) says that his A. rapens7s
from Rapa Id. is “‘very closely allied” to magellanica, but to judge from his
illustrations they seem to have little in common.
Trisetum Pers. About 65; N. and S. temp. (also S. Braz.).

chromostachyum Desv. Centr—S. Chile.
Danthonia DC. About 100; essentially southern (S. Amer., S. and E. Afr., Austral.,
N. Zeal.), but extending north to N. Amer., S. Eur., and India.

collma Phil. S. Chile.
Koeleria Pers. 60 (DomIN); N. temp., S.E. Austral., Tasm., N. Zeal., S. Amer.
And. to Patag., Falkl.

micrathera (Desv.) Griseb. S. Chile.
Megalachne Steud. 2 very distinct sp., berteroniana Steud. and masafuerana
(Skottsb. et Pilg.) Hubbard ms. Reduced to Bromus by PILGER (797), but restored to
generic rank in his posthumous paper on the system of the Gramineae (798).
Chusquea Kunth. About 100 (?) sp. Mex. to Argent. and S. Chile.

fernandeziana Phil. Related to Chilean species.

Cyperaceae.

Cyperus 1. A world-wide, essentially tropical genus of about 600 sp. (KUKEN-
THAL).

eragrostis Lam. (vegetus Willd.). N. and Centr. Amer. to Braz., Urug., Argent.
and S. Chile; Easter I. Often introduced and perhaps not indigenous in Juan
Fernandez.

reflexus Vahl. Mex. and Tex. to Braz., Argent. and S. Chile. Indigenous?
Scirpus L. About 400; cosmopolitan.

nodosus Rottb. Circumpolar, S. temp. zone. Recently the Chilean plant was
segregated as S. molinianus Beetle, but I am unable to recognize this as specifi-
cally distinct (326).

202 C. SKOTTSBERG

cernuus Nahl. Subcosmopolitan and of variable habit; south to Fueg. and
Falkl.
Eleocharis 8. Br. A world-wide genus of about 150 sp. (SVENSON).
fuscopurpurea (Steud.) H. Pfeiff. S. Chile. Listed before as a subspecies or
variety of HY. maculosa (Vahl) R. Br. (W. Ind.—Braz.).

Oreobolus R. Br. An austral-circump. genus of 6 sp., extending north to Malaysia
and Hawaii.
obtusangulus Gaud. Andes of Colomb. and Ecuad. to Fueg., Falkl.

Cladium ®. Br. 48 (KUKENTHAL); widespread, trop.-subtrop. with preponderance
S.E. Asia-N. Guin.—Austral.—N. Zeal., north to Hawaii, W. Ind., Braz. 2 boreal sp.

scirpoideum (Steud.) Benth. et Hook. f. Nearly related to C. angustifolium
(Gaud.) Benth. et Hook. f. (N. Guin., Tahiti, Hawaii).

Uncimia Pers. 32; circump.—austr.-subantarct. with outposts north of the Equator
in Centr. Amer. and Philipp. Is.; greatest concentration of species in N. Zeal.

brevicaulis Thouars. S. Chile’ to Fueg.; Falkl:; Trist; da°@)> St Pauleand
Amsterd. Is.

Douglasii Boott.

costata Kuekenth. Related to the former.

pirleoides Pers. Andes from Colomb. to S. Chile and Patag.

tenuis Poepp. Centr. Amer.?; Andes of Chile to Fueg.

Carex L. About 1500; worldwide, but comparatively few trop. sp.

Banks Boott. Andes of Centr. Chile to Fueg. Belongs to the boreal section
Frigidae-Fuliginosae of KUKENTHAL.

berteroniana Steud. Sect. Echinochlaenae of KUKENTHAL, with few excep-
tions (Austral., Tasm., Norfolk I. and one Chilean sp.) confined to N. Zeal.

Palmae.

Juania australis (Mart.) Drude. Affinities undoubtedly trop. Andean. Included
in the Iriarteae by DRUDE, but perhaps nearer to the Morenieae, or possibly
regarded as type of a separate subtribe. HUTCHINSON brought ¥awania next to
Ceroxylon, following BENTHAM and HOOKER, CROIZAT (77) remodelled Morenieae,
including Fuanza and the Mascarene Hyophorbe, otherwise linked with Chamae-
doreae.

Bromeliaceae.

Hesperogreigia Berteroi Skottsb. 2 or perhaps more sp. Subtrop. Andean, related
to Greigia (8 sp., Costa Rica, Venez.-S. Chile). LoOosER called Hesperogreigia
monotypical, because my paper (2Z0) had escaped him; if not, he would not have
written ““:Quiza de la afinidad de Greigia?’” (770. 291). Inspired by L. B. SMITH
he suppressed Hesperogre7gia in an appendix to his paper (p. 299), an attitude
I cannot take. The two genera differ very much.!

1 Sepals free. Pericarp thin. Seed testa thin, with a thick coat of mucilage, forming a massive
pulp. Raphe inconspicuous. Scales with irregular, more or less isodiametric cells . Hesperogreigia
Sepals connate into a tube. Pericarp fleshy. Testa very hard, without mucilage. Raphe very con-
spicuous. Scales with long and narrow cells radiating from centre. ......... . Greigia

DERIVATION OF THE FLORA AND FAUNA 203

Ochagavia elegans Phil. Very near the Chilean genus Rhodostachys; see 229. 110
and 249.774.

Juncaceae.

Lusula DC. A world-wide essentially temp. and mainly boreal genus of about
80 sp., also represented on trop. mountains and in the S. Braz. highland; south
to Fueg., Falkl. and N. Zeal.

masafuerana Skottsb. Nearly related to some Andean sp. (Mex., Boliv.—
Fueg.) neading taxonomic revision.
Funcus LL. Mainly temp.subtrop.; about 225. Well represented S. hemisph. (about
50), especially Austral. and N. Zeal., less so temp. S. Amer.

wmoricatus Laharpe. Ecuador—S. Chile, Braz., Argent., Urug.

capillaceus Lam. Ecuador, Centr. Chile, Braz., Argent., Urug. Introduced?

procerus E, Mey. S. Chile.

dombeyanus Gay. Pert, Chile, Braz., Argent., Urug.

planifolius R. Br. S. Chile, Austral., Tasm., N. Zeal. Introduced?

Iridaceae.

Luveruvacopreng. 5 sps,°3 Chile, 2 N..Zeal.
formosa Grah. S. Chile.

Piperaceae.

Peperomia Ruiz et Pav. Possibly over 1500; pantropical.

berteroana Mig. Extremely like P. ¢v7stanenszs Christoph. from Gough L;
see 245.

margaritifera Bert. ex Hook. Possibly related to P. Remecket C. DC. (Samoa).

Skottsbergii C. DC. Allied to the former.

These species form a separate subgenus 77/denidium Skottsb. (247), which
seems to stand closer to palaeotropical than to neotropical groups. P. derteroana
and ¢ristanensis show affinity to P. urvilleana A. Rich. (Austral., N. Zeal., Lord
Howe I., Norfolk I.) and P. Ventenati Mig. (Java).

fernandeziana Miq. Central (Frai Jorge, Talinay) and S. Chile (Valdivia). Be-
longs to subgen. Sphaerocarpidium.

Urticaceae.

Urtica L. About 40 sp. widely scattered in temp. zones on both hemisph., well
represented in Amer. (13, Mex.—Fueg.).

Masafuerae Phil. Related to U. echinata Benth. (Ecuador).

fernandeziana (Rich.) Ross. A very distinct species without near relatives
(WEDDELL).
Boehmeria Jacq. About 100, trop.-subtrop.

excelsa (Bert. ex Steud.) Wedd. Seems to be more nearly related to Pacific
than to American species. Very similar to B. dealbata Cheesem. (Kermadec Is.).

204 C. SKOTTSBERG

Parietaria L.. Some 30 sp.; in all parts of the globe. Several medit.-orient., others
N. and S. Amer., one (P. debzlis Forst.) supposed so be widely distributed.

humifusa Rich. Chile. Until now listed as debz/7s which, in WEDDELL’s mon-
ograph, is a collective species. I have shown (247) that FORSTER’s species from
Australia, New Zealand and Polynesia has little to do with the forms referred to
this but occurring elsewhere and that none of the specimens from S. Amer. seen
by me belong to true dedz/zs. Pending a revision of the genus, Awmzfusa is a cor-
rect name for the Chilean plant.

Loranthaceae.

Phrygtlanthus Eichl. About 30; bicentric: Mex.—Braz. and Chile, Argent., Urug.;
Austral N3-ZealeeN: «Guim Philipp. Is!
Berteroi (Hook. et Arn.) Reiche. Related to Andean sp.

Santalaceae.

Santalum L. 18; Austral—N. Guin. 5, Melan. 2, Polyn. 2, Hawaii 7, Bonin 1, and
the following. See 236 and 270.

fernandezianum F. Phil. Extinct. Belongs to sect. Polynesica Skottsb.; not
as has been said (zzz), related to S. freycinetianum of Hawaii which belongs to a
different section (270).

Chenopodiaceae.

Chenopodium 1. Over 250; world-wide, but essentially temp.

Sanctae Clarae Johow, crusoeanum Skottsb. and nesodendron Skottsb. On
my request Dr. P. AELLEN, who created sect. Skoftsbergia to receive the three
island endemics, sent me the following remarks: “Ich wurde sagen, dass die drei
Arten zu keiner der uns heute bekanntgewordenen Arten irgendwelche verwandt-
schaftliche Ziige und Beziehungen aufweisen. Es sind vollig isolierte Typen, Re-
likte eines im Meere versunkenen Florenreiches.”’ Evidently Dr. AELLEN does
not support the idea that they are related to panzculatum Hook. (N. Amer., Pert—
Chile) or oahuense Meyen (Hawaii), which latter has the same arboreous habit.

Salicornia L. Widely distributed, about 30 sp.
Jruticosa V. (peruviana Kunth). Taken in a wide sense Mediterr., S. Afr., W.
Ind., Polyn.; peruviana, W. coast of S. Amer. to S. Chile.

Aizoaceae.
Tetragonia L. About 25 sp. Strongly represented in S. Afr., several sp. in Chile.
expansa Murr. Coasts and islands of the Pacific; Braz.

Caryophyllaceae.

Spergularia Pres]. A widely scattered genus of about 40 sp.; numerous in Chile.
confertifiora Steud. San Ambrosio. Related to species from Centr. Chile.
masafuerana Skottsb. Related to the former and to S. media (L.) Presl.

DERIVATION OF THE FLORA AND FAUNA 205

Paronychia L. About 40, scattered in temp. and subtrop. regions.
ohilensis OG: Centri—S. Chile, S. Braz., Argent.

Ranunculaceae.

Ranunculus L. Probably over 300; world-wide, particularly boreal, numerous sp.
IN; *Zeal.

caprarum Skottsb. Apparently with distinct relations in New Zealand, not
near boreal or S. American species (229. 125; 173).

Berberidaceae.

Berberis .. About 540 sp. distributed over the N. hemisph. and extending south
along the Andes to Fueg.; also S. Braz.

corymbosa Hook. et Arn. Referred to sect. Corymbosae Schneid. (3, trop.
And.). Not close to Chilean species.

masafuerana Skottsb. Near the former.

Winteraceae.

Drimys Forst. 40; 29 N. Guin., 6 Austral., 1 Borneo—Philipp. Is., 3 Mex.—Fueg.,
and the following.
confertifolia Phil. Near D. Winteri Forst. (Centr. and S. Chile to Fueg.).

Lactoridaceae.

Lactoris fernandeziana Phil. With regard to the position of the family system-
atists disagree, some bringing it to Polycarpicae, others to Prperales. GUNDERSEN
(720) asserts that it belongs to the latter, but it differs from this in very important
characters.

Cruciferae.

Cardamine 1. About 130; world-wide, essentially temperate, many Chilean sp.
chenopoditfolia Pers. Boliv., Braz., Argent., Urug.
flaccida Cham. et Schlechtd. Centr. and S. Amer. to S. Chile.
Kruesselii Johow. Related to C. vulgaris Phil. (S. Chile).

Saxifragaceae.

Escallonia Mutis. About 50, Colomb.—Fueg., Braz., Argent., Urug.
Callcottiae Hook. et Arn. Occupies a rather independent position among
the Chilean sp. (z58).

Rosaceae.

Rubus L. A very large, temp. and trop.-montane genus, richly developed in the
N. hemisph.
geoldes Sm. S. Chile to Fueg., Falkl.

Margyricarpus Ruiz et Pav. 4 or 5, trop. Andes to Patag.

206 C. SKOTTSBERG

digynus (Bitter) Skottsb. Near J7. pzunatus (Lam.) O. K. (Pert, Chile, S.
Braz., Argent.).
Acaena \.. An austral genus’ of over 100 sp., the majority in’ S: Amen sya
Austral.-N. Zeal., single sp. Calif., Hawaii, the Cape, etc.

masafuerana Bitter. Near A. antarctica Hook. f. (Magell., Fueg.).

ovalifolia Ruiz et Pav. subsp. australis Bitter. S. Chile to Fueg., Falkl. Per-
haps accidentally introduced to Juan Fernandez.

Leguminosae.

Sophora V.. 70-80, Old and New World; sect. 7etrapterae (‘“Edwardsia’’) austral-
circump.; 17 closely related species.

fernandexiana (Phil.) Skottsb. and masafuerana (Phil.) Skottsb. Very near
the Chilean “¢etraptera’ (S. macnabiana Phil.), not identical with S. ¢etraptera
Ait. from N. Zeal.

Rutaceae.

Fagara L. Over 200, pantrop.; a large palaeotrop. group well represented in the
S. Pacific (Austral., N. Caled., Polyn.), extending north to Hawaii. The two Juan
Fernandez species F. mayu (Bert., Hook. et Arn.) Engl. and externa Skottsb.,
which are very closely allied, form the sect. M/ayu Engl. Related to W. Pacific sp.

Euphorbiaceae.

Dysopsis Baill. 3; 1 in Ecuador [D. paucidentata (M. Arg.) Skottsb.], 1 in S. Chile
[D. glechomotdes (Rich.) M. Arg. p. p.|, and the following.
hirsuta (M. Arg.) Skottsb.

Callitrichaceae.

Callitriche LL. Widely distributed in the N. hemisph., austral-circump., south to
Austral. and S. Chile but absent from the southern half of Afr.

Lechleri (Hegelm.) Fassett. S. Chile to Magell. (249. 781).

Rhamnaceae.

Colletta Comm. 17, Andean and extratrop. S. Amer.
spartioides Bert. ex Colla. A well-marked species, related to Chilean and
Ofheres, emer norms,

Flacouttiaceae.

Agata Ruizsetwbaves 20-21 .sp.; 18-19 Chile,.1, Braz:, 1, Argent.
fernandeziana Gay. Closely related to A. serrata Ruiz et Pav. (S. Chile).

Myrtaceae.

Ugni Turcz. 15, Mex., Centr. Amer., Andes, south to Chiloé, Venez., Braz.

Selkirkii (Hook. et Arn.) Berg. Probably related to species in Centr. Amer.
and Venez., perhaps also to U. Candollei (Barn.) Berg from Chile; not very close
to U. Molinae Turcz. (S. Chile; KAUSEL in litt.)

DERIVATION OF THE FLORA AND FAUNA 20

sa |

Myrteola Berg. 12, Colomb.—Chile.
nummularia (Poir.) Berg. Chile, Cord. Linares to Fueg., Falkl.
Nothomyrcia \ausel. A monotypical genus, related to Chilean genera.
fernandeziana (Hook. et Arn.) Kausel.
Myrceugenia Berg. About 40; trop. Andes, S. Braz. (many sp.) and about 20 in
Chile, south to Chiloé.
Schulzei Johow. Related to J. planipes (Hook. et Arn.) Berg (S. Chile).

Gunneraceae.

Gunnera 1... About 30 sp. Subgen. Pangue, 16 sp. Costa Rica, Colomb.-S. Chile,
Magell., Braz.; Hawaii 2 sp., but some more, of doubtful taxonomic status, have
been described. Other subgenera in the Andes of Colomb. to Chile, south to
Fueg. and Falkl.; Urug.; S. and Centr. Afr.; Malays.—Philipp. Is.; N. Zeal., Tasm.
peltata Phil. and Masafuerae Skottsb. related to Andean sp.
bracteata Steud. Seems to come closer to the Hawaiian G. kauatens?s Rock
than to S. Amer. species.

Halorrhagidaceae.

FHlalorrhagis Forst. About 80, the great majority Austral—N. Zeal., single sp.
scattered north to Indonesia, Philipp. Is., China and Japan, and east to Rapa
and J. Fern.

asperrima Skottsb., masatierrana Skottsb. and masafuerana Skottsb. belong
to Subsect. Cercodia and are closely related to H. erecta (Murr.) Schindl. (N. Zeal.)
and other Austral. and N. Zeal. sp. All J. Fern. forms were formerly incorrectly
identified with erecta.

Umbelliferae.

Cemieiia 1.29; 19 Afr., Madag., 1 China, 2 Austral., 5 trop.. Amer and, the
widely dispersed C. aszatica (L.) Urb.

trifiora (Ruiz et Pav.) Nannfeldt. Centr. and S. Chile, formerly included under
astatica. Introduced?
Eryngium LL. About 230, in all parts of the world, but with two centres: W. Eur.—
Medit. and trop.—subtrop. Amer. Poorly represented in N. Amer. and Australas.

bupleuroides Hook. et Arn., inaccessum Skottsb. and sarcophyllum Hook.
et Arn. form a special sect. /rufzcosa, but differ mainly in being arborescent from
the species occurring on the mainland.
Apium L. About 30; in all parts of the world.

fernandezianum Johow. A well-marked species, probably not nearly related
to the Chilean species, but showing some affinity to A. prostratum Labill. (Austral.)
or australe Thouars (Tristan da C.)

Ericaceae.

Pernettya Gaud. About 12, 6 or 7 Mex. and Centr. Amer. and along the Andes
to Fueg. and Falkl.; 1 Galapag. Is., 2 N. Zeal., 2 Tasm.

rigida (Bert.) DC. A well-marked sp. (252).

208 C. SKOTTSBERG

Empetraceae.

Empetrum L. A bipolar genus, /. xzgrum L. north, £. rubyum Vahl south, but
according to GOOD the latter is represented in the Subarctic by special forms.
See 270. 781.

rubrum Vahl. Andes of S. Chile and Argent. to Fueg. and Falkl.; Tristan da C.

Convolvulaceae.

Dichondra Forst. 5 trop.-subtrop. Amer., 1 N. Zeal., and the following.

repens Forst. Widely spread over both hemispheres, north to N. Amer. and
China, south to S. Chile and N. Zeal.
Calystegia RK. Br. 10-20; temp.-subtrop. in all parts of the world.

tuguriorum R. Br. S. Chile (Hantelmanni Phil.) and N. Zeal.

Boraginaceae.

Selkirkia Berteroi (Colla) Hemsl. According to JOHNSTON (248) very near Hackelia
Opiz, a genus centering in western N. Amer., with outposts in S. Amer. and
Euras.; //. revoluta (Ruiz et Pav.) Johnst. Peru to Boliv. and Argent.

Verbenaceae.

Rhaphithamnus Miers. 2, one in Centr. and S. Chile. Near C7tharerylon L., a neo-
trop. genus of about 20 sp., Mex. and W. Ind. to Boliv. and Braz.
venustus (Phil.) B. L. Robins. A very distinct species.

Labiatae.

Cuminia Colla. 2. An isolated genus, referred to Praszotdeae by EPLING (go), a
palaeotropical group best developed in Hawaii, but whereas Cumznza has the drupe
of this tribe, it has the corolla of Stachyotdeae-Menthinae, where BRIQUET placed
it; BURGER’S statement that the flowers are “Lippenbliiten” (4z. 23) is erroneous.

fernandezia Colla and eriantha Benth. Perhaps united by intermediate forms.

Solanaceae.

Solanum L. Probably over 1000; in all parts of the world, richly represented in
trop. Amer, many in Chile.

fernandezianum Phil. Related to S. “wderosum L. coll.

masafueranum Bitter et Skottsb. A very well-marked Morella.

robinsonianum Bitter. See above p. 200.
Nicotiana 1. 60; 45 Amer., Calif—Mex., And. S. Amer. (Ecuad.—Pert—Chile, Braz.—
Argent.—Patag.), 15 Austral. (1 N. Caled., etc.).

cordifolia Phil. Belongs to the Austica group, confined to trop. Amer. and
Australas., and related to NW. Ratmondii Macbr. (Andes of S. Pert), solanzfolia
Walp. (N. and Centr. Chile) and panzculata VL. (Pert, N. Chile). Dr. GOODSPEED
expressed his opinion (in litt.) that the Aws/7ca complex originated in the region
now occupied by western Bolivia and Pert and extended to the ‘‘Juan Fernandez
land’, becoming isolated during the final uplift of the Andes or possibly even earlier.

DERIVATION OF THE FLORA AND FAUNA 209

See also r72. F. B. H. BROWN (35. III. 262) suggested an affinity between cordifolia
and fatuhivensis F. B. H. Brown (Marquesas), but they belong to different subgenera
and the latter is referred to V. fragrans Hook. from N. Caled. as a variety (772).

Scrophulariaceae.

Mimulus L. About 130; widely dispersed, predominantly western N. Amer. (80
Calif.), south to S. Chile; 2 Afr., Madag., 4 E. Asia, 5 Austral., Tasm., N. Zeal.
glabratus H.B.K. N. Amer. to Boliv., Argent. and Chile; polymorphous, the
island form very close to var. parviflorus (Lindl.) Grant (zr4).
Euphrasia L. About 100; bor.-circump. with isolated populations in the S. Andes,
south to Fueg. and Falkl., and in Austral—N. Zeal., the gap between E. Asia
and Australia bridged over by trop. mountain stations; see map 7&8. 224.
formosissima Skottsb. The fact that this species is very unlike the 777fidae
of Chile makes it particularly interesting. In 229.169 I emphasized the differ-
ence between /formosisstma and the Australes of N. Zealand and placed it nearer
to the boreal Semzcalcaratae. WETTSTEIN, in his contribution to my paper (I.c. 209),
expressed the opinion that it could be attached to a Japanese group of species,
intermediate between Australes and Semicalcaratae. The question was taken up
by Du RIeETz who thinks, with good reason I believe, that I overrated its rela-
tions to boreal species (77. 533) and that, in most respects, it is more nearly related
to N. Zealand forms.

Plantaginaceae.

Plantago L. About 270; world-wide, essentially temp.

fernandezia Bert. The nearest relative appears to be P. przuceps Cham. et
Schld. of Hawaii. Both belong to sect. Palaecopsyllium, scattered over the S. hemisph.
and extending north to N. Amer., Hawaii and S. Eur.: N. Amer. 5, S. Amer. 1,
S. Eur. 2, Afr. 5, St. Helena 1, Madag. 1, Lord Howe I. 1, Auckl. Is. 1, Rapa 2,
and Hawaii 9.

truncata Cham. Centr. and S. Chile. PILGER distinguished the island form as
a separate subspecies close to ssp. firma Pilger, but I doubt that it deserves the
rank assigned to it, and it is even possible that it is a very late arrival in the
islands.

Rubiaceae.

Fledyotis L. A large pantrop. genus.
thesitfolia St. Hil. Trop.-subtrop. S. Amer., in Chile south to Chiloé. Very
likely not truly indigenous in J. Fern.
Nertera Banks et Sol. 10-12; centering in N. Zeal. (4 endem. sp.), north to Malaya
and Hawaii; Tristan da C.; S. Amer. from Falkl. and Fueg. to Colomb. and Mex,
granadensis (L. fil.) Druce. A polymorphous species, reported from S. Amer.,
Tristan da C., Austral., Tasm., N. Zeal., Java and Hawaii—see 244, where I pointed
out that it is heterogeneous but that the plant from Masafuera seems to be iden-
tical with the form common in Magell. and Falkl.
Coprosma Forst. 90; a western Pacific genus centering in N. Zeal. (39) and distri-
14 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

210 C. SKOTTSBERG

buted from Australia to Malays., Melan. and Polyn. with a secondary centre in
Hawaii (18); entirely absent from Amer.

Hookeri (G. Don) W.R. B. Oliver (792). Forms a separate monotypical section.

pyrifolia (Hook. et Arn.) Skottsb. Belongs to a section of 8 Polyn. sp. (4
Mahiti, 1 Cook Is., 1 Kapa, 1_ Pitcairn, a_.J. Fetn:).
Galium 1. World-wide, essentially boreal. Over 500 have been described, almost
1/, of these Medit.—Orient.; about 50 in S. Amer., mostly along the Andes and
extending south to Fueg., Falkl. and S. Georgia.

masafueranum Skottsb. Related to G. eriocarpum Bartl., DC. and frichocar-
pum DC. (both Coquimbo-Cord. Linares), the fruit as in masafueranum covered
with straight hookless hairs.

Campanulaceae.

Wahlenbergia Schrad. Essentially S. African; about 230 sp. (9 S. Amer., 10 S.
Eur.—Orient., 2 or 3 St. Helena, 20 trop. Afr., 6 Madag.—Mascar., 150 S. Afr., 9 As.,
tN; Goin 3 Austtal., 8 IN. Zeal) 5); Ferm,):

Larrainii (Bert. ex Colla) Skottsb., fernandeziana A. DC. p. p. and Grahamae
Hemsl. are closely related to each other.

Masafuerae (Phil.) Skottsb. approaches the former, but has the large tuber
of the following.

Berteroi Hook. et Arn. occupies a rather independent position.

Most African species are annuals and quite unlike the island species, and
these have little in common with the single Chilean or the other American forms,
nor with those from Australia or N. Zealand, even if there is a superficial likeness
between IV. Masafuerae and a couple of perennial S. African species such as
W. Ecklon Buek and oxyphylla A. DC. On the other hand, the resemblance
between the J. Fernandez and St. Helena species is quite striking, and in spite of
the difference in the number of carpels, the ovary being trimerous in the former
and dimerous in the latter, the possibility of a common origin cannot be dismissed.
The number of carpels in the genus varies between 2 and 5; in most species
they are 3. HEMSLEY (727. 61) regarded the St. Helena species as allied to African
and Juan Fernandez species.

Lobelia L.. 350-370; particularly numerous in Amer. and Afr., less so in As.,
Austral. and Oceania, 2 in Eur.
alata Labill. S. Chile, S. Afr., Austral. A sea-side sp.

Compositae.

Lagenophora Forst. A bicentric austr.-subantarct. genus of 16 sp., most numerous
in Austral.—N. Zeal.; Fiji, Rapa, extending north to Philipp. Is. and Hawaii.
Flariott Franch. Andes of S. Chile to Fueg.

Lrigeron L. A large bor.-temp. genus. Ind. Kew. lists about 700 sp. as valid,
half of them in N. Amer. and about 100 in S. Amer., where the genus is richly
developed along the Andes, south to Patag., Fueg. and Falkl. Not few are known
from trop. mountains in the W. and E. hemispheres; very few reported from

DERIVATION OF THE FLORA AND FAUNA 2251

Australia. One species is found on Rapa, related to the Juan Fernandez species
according to BROWN (35. III. 338).

fruticosus DC. and luteoviridis Skottsb. are related to each other.

Ingae Skottsb., Innocentium Skottsb.' and turricola Skottsb. form another
group.

rupicola Phil. stands apart from all other species.

VIERHAPPER (see 229.182) suggested that the island species are allied to
Andine species, but also that /. fruticosus comes very close to /. lepidotus Less.
of Hawaii; this is, however, now referred to Zetramolopium by SHERFF. There is
no “rzgeron in Hawaii.
Guaphalium L. A large subcosmopol., essentially temp. genus needing revision.

spiciforme Sch. Bip.; comp. 229. 187-188. Patag. Fueg. The assemblage to
which the alpine species of Masafuera belongs is in a state of taxonomic confusion.
The identity with the Magellanian plant may be doubted, but I am convinced
that the island form cannot be referred to either americanum Mill., purpureum L..,
Spicatum Lam. or mucronatum Phil. which, together with coxsanguimeum Gaud., are
regarded as forms of a single polymorphous taxon.

Abrotanella Cass. An austr.-subantarct. genus of 20 sp., the majority in N. Zeal.
with the subantarct. islands (9) and W. Patag._Fueg.—Falkl. (5); of the remainder
rt in Austral., 2 Tasm., 1 N. Guin., 1 Rodriguez I., and the following.
crassipes Skottsb. Very near A. MoseleyZ Skottsb. nom. (W. Patag.); see
229. 189-190.
Robinsonia DC. BENTHAM (20. 460) remarks on Rodivsonia and Rhetinodendron:
“Although their connexion with Eusenecioneae seems greater than with any other
subtribe or tribe, yet in their dioecious capitula, in the presence of small free
anthers without pollen in the female floret and some other points they approach
the subtribe Petrobieae of Helianthoideae.’’ The idea was rather attractive, be-
cause the Petrobieae inhabit St. Helena and S. America. Another suggestion men-
tioned the Hawaiian Duabautza and Razllardia, but they are not dioecious. Re-
cently an undisputable relative was discovered in New Guinea, 4rachionostylum
Mattfeld (278. 27-28):

Von Senecio, dessen pacifische Arten unserer Pflanze recht nahe kommen, unter-
scheidet sich diese Gattung wesentlich durch die Zweihdusigkeit ... und weiter in
Zusammenhang damit durch die Form der Griffelschenkel der Scheibenbliiten, die
der Fegehaare ganz entbehren und vorn nicht gestutzt sondern abgerundet sind. Diese
Merkmale hat sie aber mit den auf Juan Fernandez endemischen Gattungen Rodbinsonia
und PRhetinodendron gemeinsam, von denen sie sich iiberhaupt durch kein generisches
Merkmal unterscheidet. Aber die Ubereinstimmung erstreckt sich sogar auf kleinere
Merkmale, wie den leicht hinfalligen Pappus, die kurzen, aber verhaltnismassig sehr
breiten, etwas vorspringend gestreiften Zungen, die Form der durch ein kleines Be-
cherchen gekr6nten Achaenen, und schliesslich die Wuchsform und Verzweigungsart:
bei beiden schliesst der schopfig beblatterte Stengel mit dem Bliitenstande ab, wahrend
der Fortsetzungsspross aus der Achsel eines der oberen Laubblatter unter dem Blii-
tenstande entspringt und diesen bald zur Seite dringt.... So bleibt als einziger Un-
terschied der durch die sehr verschiedene Blattform bedingte habituelle Eindruck... .

1 Described as a variety of the former and now raised to specific rank.

212 C. SKOTTSBERG

Die Bliitenképfe sind bei Rodimsonia erheblich zahlreicher und kleiner, kurz und breit
glockenf6rmig, bei unserer Pflanze aber schmalglockig.

Wiirden diese Sippen demselben pflanzengeographischen Gebiete angehéren, so
wiirde man sie sicher nicht generisch trennen kénnen. Was hier aber wesentlich dazu
zwingt, ist, dass die Zweihiiusigkeit, die ja hier die wesentliche Ubereinstimmung be-
dingt, bei den Kompositen zu den verschiedensten Malen in den verschiedensten und
auch in denselben Gruppen entwickelt wurde und daher keineswegs als Kriterium
fiir eine generische Verwandtschaft angesehen werden kann.

To this I shall remark that nobody would think of uniting Rodzusonza and
Rhetinodendron, because they differ in essential floral characters. brachionostylum
is, to judge from the description and plate XC: B (only 2 known), closely related
to Robimsonia but not to Rhetinodendron. The mode of growth is not quite the
same, for in Robzmsonia (as well as in Rhetinodendron) 2 or 3 innovations are
developed, and none of them continues the mother axis. Besides, Brachionostylum
has petiolate, penninerved leaves of a type current in dicotyledons and widely
different from the sessile, linear-lanceolate, broadly clasping and parallel-veined
leaves of Rodbznsonta.

gayana Dcne and thurifera Dcne are a pair of related species.

evenia Phil. and Masafuerae Skottsb. form another pair.

gracilis Dcne stands more isolated.

Symphyochaeta (DC.) Skottsb. See 249. 785.
macrocephala (Dcne) Skottsb.
Rhetinodendron Meisn. Berterii (Dcne) Hemsl. See above.

Centaurodendron dracaenoides Johow. Not, as has been supposed, just an ar-
boreous Cendaurea, but differing materially in flower structure (273).

Yunquea Tenzii Skottsb. See 237. 163.

Dendroseris C. Don. An isolated genus, perhaps distantly related to 7hamuno-
servis Phil. from Desventuradas Is., but probably not to /7¢chza Hook. fil. BEN-
THAM stated (20.480) that the achenes of Dendroseris differ from those of the
Cichorioideae in general, and that also the involucre and the habit are different.
With regard to the achenes this is certainly true of Dendroser7s s. str., and also
of Phoentcoseris, whereas they are of a rather normal type in Rea and Hespe-
roserts. All are devoid of paleae, which are present in 7ammnoseris and Fitchza,
a distinctive feature pointed out by BENTHAM, but the awned achenes of /2¢chza
differ from those of all the other genera mentioned. In spite of the profound
discrepances in such important characteristics as inflorescence, involucre, recep-
tacle and stigma Z7hamunoseris has more in common with the Denxdroseris as-
semblage than with other genera.

macrophylla D. Don, macrantha (Bert. ex Decne) Skottsb., marginata (Bert.
ex Decne) Hook. et Arn. and litoralis Skottsb., all lumped together by JOHOW, are
well-marked species; comp. 229. 201-204.
Phoenicoseris Skottsb., 249. 787.

pinnata (Bert. ex Dene) Skottsb. and berteriana (Dcne) Skottsb. (Masatierra)
are closely related but quite distinct; regia Skottsb. (Masafuera) differs from both
in leaf shape, but fertile specimens have not been found.

DERIVATION OF THE FLORA AND FAUNA 213

Rea Bert. ex Dene p.p. See 249. 788.

neriifolia Dcne. Little is known of this, but it is undoubtedly a very good
species; micrantha Bert. ex Decne and pruinata (Johow) Skottsb. are near rela-
tives but good species (comp. 229. 207).

Hesperoseris Skottsb. gigantea (Johow) Skottsb. See 279. 788.

Vicarious species.

VIERHAPPER (276) distinguished between true and false vicarism. True vi-
carists have arisen from a common initial species and become differentiated either
within the limits of the area this once occupied or after penetration into a
different habitat, followed by isolation, whereas in the case of pseudo-vicarism
they have a different origin; a second species may invade the area of the first
and colonize such parts of the area as are unsuitable to the latter. Very often
the term ‘“‘vicarious’’ has been taken in a much wider sense: any two related
species replacing each other in separate areas were called vicarious, and phyto-
geographers used the term to designate two plant species, related or not, that
played corresponding roles in two closely allied plant communities; in this case
Nothomyrcia fernandeziana and Myrceugenta Schulzet, which form the bulk ot
the forest in Masatierra and Masafuera, respectively, are vicarists, although they
belong to different genera. Species not fulfilling the conditions claimed by VIER-
HAPPER were called ‘‘substitute species’.

WULFF (297. 66-67) devotes considerable space to a discussion of vicarism.
He agrees with VIERHAPPER: true vicarism is a result of one taxon breaking
up into two, adapted to different habitat conditions. CAIN (42. 265) expresses
himself in slightly different words, but their meaning is the same: ‘‘closely re-
lated allopatic species which have descended from a common ancestral popula-
tion and attained at least spatial isolation.’ He summarizes statements made
by DRUDE, DIELS, WULFF and SETCHELL, who called species vicarious if they
were only slightly discontinuous morphologically but widely so geographically;
certainly anybody would call them vicarists but use the same term for a pair
of intimately related forms of which one inhabits granite and the other limestone
within the same geographical area.

I am afraid that, in most cases, we know very little or nothing at all of
the ancestry of species we are used to call vicarious, let it be that we have
reason to assume that they have differentiated out of a common population some
time in the past. This is, at least, the situation in Juan Fernandez. Here I
should perhaps refer to what CAIN (l.c. 276) calls polytopic species, 1.e. when
the same taxon occurs in two or more discrete areas, disjunction being the
result either of dispersal from one original centre or of the breaking up of an
area through subsidence or formation of some other kind of barrier; upon a close
investigation it has been shown in many instances that slightly different forms
within the original population happened to become isolated and appear as ex-
amples of true vicarism.

JoHOw (z50. 233) held forth that 6 species endemic in Masatierra correspond
to 6 other species endemic in Masafuera, but he did not use the term vicarious.

214 C. SKOTTSBERG

These were (the species from Masatierra mentioned first): Dexdroserts micrantha-
gigantea (belong to different genera), Waklenbergia Berteroi-tuberosa (= Masa-
fuerae, not very closely related), Myrcengenia fernandeziana—Schulzet (different
genera), Lryngium bupleuroides—sarcophyllum (only distantly related), Cardamiue
alsophila( = flaccida) — Kruesselii(not nearly related), and Urtica glomeruliflora(= fer-
nandeztana)—- Masafuerae (belong to different sections). There is not among them
a single example of either vicarism or pseudo-vicarism in the sense of VIERHAPPER.

In 1914 (227) I distinguished 4 pairs of vicarists in Juan Fernandez: Dendro-
servis micrvantha—gigantea, Myrceugenia fernandeziana—Schulzet, Gunnera peltata-
Masafuerae, and Peperomia margaritifera—Skottsbergi; the flora of Masafuera was
at that time little known. Of these pairs the last two still hold good, the species
replace each other from a taxonomical as well as from a sociological viewpoint;
nevertheless | am not prepared to argue that they arose out of ove initial spe-
cies: several species may have been involved, for they differ in a number of
minor characters and all we can say is that they make the impression of coming
from the same stock. On the basis of our present knowledge of their taxonomy
the following nine pairs are distinguished; the species replace each other, call
them vicarious or substitute: Berberzs corymbosa—masafuerana, Chenopodium cru-
soeanum—nesodendron, Dendroseris macrantha-macrophylla, Fagara mayu—externa,
Gunnera peltata— Masafuerae, Halorrhagis masatierrana—masafuerana, Peperomia
margaritifera—Skottsbergu, Robinsontia evenia— Masafuerae, and Sophora fernan-
deztana—masafuerana. As regards Lerberis, Fagara, Halorrhagis and Sophora,
perhaps also Gunnera, Peperomia and Robinsonia, the two members of a pair are
closely related, but in the case of H/alorrhagis the situation is complicated be-
cause there is a third species, endemic in Masafuera, /7. asperrima; masafuerana
seems, however, to be a better match for masatierrana than asperrima. As we
have seen, there is also a third species of Chenopodium, endemic to Santa Clara,
very likely formerly occurring also on Masatierra, but extinct there. Of the 3
species of Dendroserzs inhabiting Masatierra, margimata and /itoralis form a pair
of one inland and one coast species.

An unbalanced flora.

It has always been argued that an island flora, where the number of species
is small compared with the number of genera and many large and widely distri-
buted families, well developed under different conditions, poorly represented or
altogether absent, furnishes one of the best proofs of the absolute oceanity of its
abode, and Juan Fernandez is no exception from this rule. | have mentioned this
above (p. 194) pointing out that most of these families have numerous genera
and species on the opposite mainland, thus Caryophyllaceae 20 genera, Compo-
sitae some 130, Cruciferae 28, Leguminosae 22, Scrophulariaceae 18, Umbelliferae
about 30, and so forth, and examples of important families in Chile not found
in the islands were also given. This state of things calls for an explanation. JOHOW
paid much attention to it; as he believed that the islands had been isolated from
the very beginning he blamed chance in all cases where, according to the cur-
rent opinion, the diaspores were adapted for dispersal across a wide expanse of

DERIVATION OF THE FLORA AND FAUNA 215

water. He mentions Compositae-Labiatiflorae, Calyceraceae, Valerianaceae, Cacta-
ceae, Orchidaceae and Dioscoreaceae as good examples. In other cases he blames
the diaspores as in Nolanaceae, Leguminosae, Violaceae, Fagaceae, Amary|lidaceae
and Liliaceae.

Also with reference to life-forms the island flora makes an impression of being
unbalanced, a fact already discussed at some length in 257. 825-830; I shall
only repeat that woody species are in overwhelming majority and annuals almost
absent, whereas they are very numerous, also proportionately, on the mainland.
Already in 1914 (227) I argued that climatic differences alone do not offer an
explanation and that historical causes must be taken into account.

Geographical elements.

In his survey of the flora p. 229-232 JOHOW discussed its composition and
distinguished various elements. The endemic genera and a few peculiar endemic
species of non-endemic genera form his first (and most ancient) group; the second
group contains the remainder of endemic species: (a) markedly distinct, (b) nearly
related to continental species. Both groups make up “Continjente A’’. The third
group, Continjente B’’, contains the species found elsewhere, all occurring in South
America except Halorrhagis ‘alata’. America is claimed as the source of the en-
demic element as well: “tanto las especies del primero como las del segundo grupo
pertenecen, con la unica excepcion del Saxtalum, a jéneros representados, si no
en Chile, a lo menos en alguna parte de la costa occidental de Sud-América’’—
this did not, however, apply to the isolated Compositae, to Lacforzs etc., which
JoHow regarded as originated from ancestors to be looked for in the Tertiary
flora of Chile. Halorrhagis, Santalum and Coprosma, mentioned later, were sup-
posed to have immigrated from western Pacific.

It serves no purpose to go into detail; when JOHOW wrote his book our
knowledge of the flora was too defective to allow him to arrive at anything like
safe conclusions. The same may be said of my 1914 paper (227), even if my short
visit to the islands revealed the existence of an up till then unknown element.
Four main groups were distinguished: I. Old Pacific (ENGLER’s Altoceanisches
Element), comprising genera or species supposed to have a long history behind
them within the precincts of the Pacific and lacking near relatives; subdivision
A, Endemic genera, with 11 sp., and Non-endemic genera with 10 sp.; subdivision
B with allied species in Hawaii, Polynesia, Australia and New Zealand, 16 en-
demic species—the genus Dendroseris was also placed here—and 1 non-endemic
(Halorrhagis). Group HU, called Neotropical, contained 6 endemic species; Cumznza
and ¥uania were included here. Group III, called Chilean, was the largest and was
divided into 3 lots: A, very distinct species, 18 (Ochagavia placed here); B, less
well-marked species, 11; C, also found in Chile, 28, but among them were 6 not
now regarded as native. So far the main difference between this arrangement and
JoHow’s lies in the greater number of species with supposed west Pacific connec-
tions. Finally Group IV, Subantarctic-Magellanian, 4 species, was added.

The study of our 1916-1917 collection added many species not known before
and gave rise to a fresh analysis of the vascular plants (239), of which a

216 C. SKOTTSBERG

short summary is given here (translated from French). The ferns are not included
below.

1. Neotropical and Andean element. Species either found in Chile (with one exception, c)
or endemic but allied to Chilean species; 47.

) in Central or S. Chile, 20.

) in the Magellan region, but not of subantarctic character, 3.

)

omm)

in South America, but not in Chile, 1.
endemic species of non-endemic genera, 21.
e) sf endemic mints
2. Element consisting of wide-spread species, also inhabiting Chile, 5.
Neotropical element, not represented in Chile, 7.
a) endemic species of non-endemic genera, 3.
b) species belonging to endemic genera, 4.
4. Magellanian - Old Antarctic element, 27.
a) subantarctic species, 10.
b) austral species, 5.
c) endemic species allied to subantarctic species, 7.
d) endemic species not allied to subantarctic species, 5.
5. Pacific element, 59.
a) isolated endemic species of wide genera, 21.
b) endemic species belonging to genera or sections of west to central Pacific distribu-
tion or, if belonging to wide genera, then more closely related to Pacific species, 17.
c) species of isolated endemic genera with supposed transpacific relations, 18.
d) species of endemic genera with unknown relations, 3.

(ayyelq)

This analysis was based on my memoir on the Phanerogams published in
1922 (229). Since that time several pending questions have been restudied, additional
plant material has come to hand, and recent monographs have been consulted.
The results were communicated above, and we shall now proceed to the following
arrangement of the angiosperms.

I. Andine-Chilean element.—69 (46.9 %).

a. Endemic species (34). In two cases, Ochagavia and Nothomyrcia, also the
genus endemic, but nearly related to Chilean genera. Chaetotropis imberbis, Chus-
quea fernandeziana, Uncinia Douglasii and costata, Hesperogreigia Berteroi, Ocha-
gavia elegans, Luzula masafuerana, Phrygilanthus Berteroi, Spergularia masafue-
rana, Cardamine Kruesselii, Escallonia Callcottiae, Margyricarpus digynus, Sophora
fernandeziana and masafuerana, Dysopsis hirsuta, Colletia spartioides, Azara fer-
nandeziana, Nothomyrcia fernandeziana, Myrceugenia Schulzei, Gunnera peltata
and Masafuerae, Apium fernandezianum, Pernettya rigida, Rhaphithamnus venus-
tus, Solanum fernandezianum, masafueranum and robinsonianum, Galium masa-
fueranum, Erigeron fruticosus, luteoviridis, Ingae, Innocentium, turricola and
rupicola.

6. Known from continental Chile, many also in other parts of S. America or
of still wider distribution (33): Stipa neesiana and laevissima, Piptochaetium bicolor,
Chaetotropis chilensis, Trisetum chromostachyum, Danthonia collina, Koeleria micra-
thera, Cyperus eragrostis and reflexus, Scirpus cernuus and nodosus, Eleocharis
fuscopurpurea, Uncinia phleoides, Juncus imbricatus, capillaceus, procerus, dom-
beyanus and planifolius, Libertia formosa, Salicornia fruticosa, Tetragonia expansa,

DERIVATION OF THE FLORA AND FAUNA 217

Peperomia fernandeziana, Parietaria humifusa, Paronychia chilensis, Cardamine
flaccida, Callitriche Lechleri, Centella triflora, Dichondra repens, Calystegia tugu-
riorum, Mimulus glabratus, Plantago truncata, Hedyotis thesiifolia, Lobelia alata.
c. Also known from San Ambrosio: Spergularia confertiflora.
d. S. American, but not reported from Chile: Cardamine chenopodiifolia.

II. Subantarctic-Magellanian element.—15 (10.2 %).

a. Endemic (4): Agrostis masafuerana, Drimys confertifolia, Acaena masafue-
rana, Abrotanella crassipes.

6. Not endemic, in several cases extending north along the Andes (11): Oreo-
bolus obtusangulus, Uncinia brevicaulis and tenuis, Carex Banksii, Rubus geoides,
Acaena ovalifolia, Myrteola nummularia, Empetrum rubrum, Nertera granadensis,
Lagenophora Harioti, Gnaphalium spiciforme.

III. Neotropical element.—19 (12.9 %).

a. Belonging to endemic genera without or with only distant relations to the
present Andean flora but presumably of neotropical ancestry (5): Megalachne
berteroniana and masafuerana, Podophorus bromoides (only tentatively referred here),
Centaurodendron dracaenoides, Yunquea Tenzii.

6. Belonging to endemic genera of undoubted neotropical affinity (2): Juania
australis, Selkirkia Berteroi.

c. Endemic species belonging to widespread genera and presumably of neo-
tropical ancestry (7): Urtica fernandeziana, Chenopodium Sanctae Clarae, crusoe-
anum and nesodendron, Eryngium bupleuroides, inaccessum and sarcophyllum.

d. Endemic species of unquestionably neotropical parentage (5): Urtica Masa-
fuerae, Berberis corymbosa and masafuerana, Ugni Selkirkii, Nicotiana cordifolia.

IV. Pacific element.— 26 (17.7 %).

Affinities along the route New Zealand—Australia—Melanesia—Polynesia—Hawail;
all endemic.

a. The genus endemic (9): Cuminia fernandezia and eriantha, Robinsonia
gayana, thurifera, evenia, Masafuerae and gracilis, Symphyochaeta macrocephala,
Rhetinodendron Berteril.

6. Only the species endemic (17): Cladium scirpoideum, Carex berteroniana,
Peperomia margaritifera and Skottsbergii, Boehmeria excelsa, Santalum fernande-
zianum, Ranunculus caprarum, Fagara mayu and externa, Gunnera bracteata,
Halorrhagis asperrima, masatierrana and masafuerana, Euphrasia formosissima,
Plantago fernandezia, Coprosma Hookeri and pyrifolia.

V. Atlantic - S. African element.—6 (4.1 %). Endemic.

Peperomia berteroana, Wahlenbergia Larrainii, fernandeziana, Grahamae, Masa-
fuerae and Berteroi.

VI. Eu-Fernandezian element.—12 (8.2 %).

Isolated endemics of unknown parentage, the Cicoriaceous genera forming a
natural group.

218 C. SKOTTSBERG

Lactoris fernandeziana, Dendroseris macrophylla, macrantha, marginata and
litoralis, Phoenicoseris pinnata, berteriana and regia, Rea neriifolia, micrantha and
pruinata, Hesperoseris gigantea.

The differences between this arrangement and the one of 1934 are consider-
able, but partly at least more apparent than real. Group 1 of 1934 corresponds
(if we exclude (b), which from a purely geographical point of view has to go to
the present group II) to I, but Se/kzrkza has now been placed in III, a group
corresponding to the 3d element of 1934. Group 4 was rather heterogeneous and
included, beside Magellanian species, several bicentric ones and some endemics
of austral-circumpolar affinity now referred to I a. Group 5, Pacific element, in-
cluded the new groups IV—VI and part of III.

II. Pteridophyta.

Six families (taken in the old sense) are represented, Ophioglossaceae only
on Masatierra, Lycopodiaceae only on Masafuera. The number of genera is 23;
21 (91.3%) are found on Masatierra and the same number on Masafuera, 3 on
Santa Clara. Two genera are confined to Masatierra (8.7%) and 2 to Masafuera,
19 (82.6%) shared by both islands, 3 of them also known from Santa Clara. A
single genus (7/yrsopteris) is endemic in Juan Fernandez and found on Masatierra
and Masafuera.

Of the 53 species listed 43 occur on Masatierra, 3 on Santa Clara and 45
on Masafuera. Only 8 species are restricted to Masatierra, 15.1% (18.6% of the
total found there) and 10 to Masafuera, 18.9 % (22.2% of the total); 35 (66%)
are found on both.

There are 18 endemic species (34%), of which 16 inhabit Masatierra and 14
Masafuera; no endemic is found on Santa Clara. Of the endemics 4 are confined
to Masatierra and 2 to Masafuera, 12 having been recorded from the two islands;
in percentage: 22.2, 11.1 and 66.7%, respectively.

A high percentage of ferns is to be expected in ‘oceanic’ islands, whether
or not isolated from the beginning, and Juan Fernandez is no exception from this
rule, for of the 200 vascular plants 26.5% are Pteridophytes.

Of the 35 species also found elsewhere 4 are confined to Masatierra, 8 to
Masafuera and 23 found on both islands, 3 of these also on Santa Clara; in per-
centage 11.4, 22.9 and 65.7, respectively. The differences in percentage between
endemic and non-endemic species is due to the occurrence, in the highland of
Masafuera, of a few mountain ferns not found on the other island. But whether
our analysis bears upon families, genera or species the distribution over the archi-
pelago is very much more even than the dispersion of the angiosperms, and this
is of course what we have every reason to expect. Also, the number of species
to a genus is greater, 2.3.

DERIVATION OF THE FLORA AND FAUNA 219

The systematical position of the endemites and the geographical distribution
of the genera and species also found elsewhere.

Hymenophyllaceae.

Trichomanes L. coll. The island species belong to Vandenboschia Copel., a pan-
trop. and circum-austr. genus of about 25 sp., extending north to N. Amer., Engl.
and Japan.

philippianum Sturm. “One of the most distinct species of 77zchomanes ...
its cellular structure quite unique’? (CHRISTENSEN 62. 3). — Also in S. Chile? (see
itr2):

Ingae C. Chr. Belongs to the neotrop. pyazdiferum group.

exsectum Kze. S. Chile, south to Chiloé. Near the neotrop. 7. tenerum Kze.

Serpyllopsis v. d. Bosch. A monotypical genus without near relatives (COPELAND
69. 37).

caespitosa (Gaud.) C. Chr. (var. fernandeztana C. Chr. et Skottsb., slightly
different from the forms described from the mainland). Falkl. and Fueg. to S. Chile.

Flymenoglossum Pres]. Monotypical and without near affinity to any known genus
{COPELAND l.c.).

cruentum (Cav.) Presl. S. Chile, Valdivia to 49° s. lat.
Hymenophyllum Sm. coll.

a. Mecodium Presl. About 100, pantrop. and austral, with several sp. in N.
Zeal.; north to Sakhalin.

cuneatum Kze. S. Chile, Valdiv—W. Patag.; Marquesas Is., Rapa. Near
ff. polyanthes Sw.,a variable pantrop. sp.

caudiculatum Mart. Pert, S. Braz., S. Chile, Valdiv.—qg° s. lat.

fuciforme Sw. S. Chile to W. Patag. A very distinct sp., referred to Meco-
dium with considerable doubt (COPELAND 66.95).

b. Hymenophyllum s. str. About 25, austr.-circump. with outlying stations in
W. Eur. and Japan; 1. peltatum (Poir.) Desv. widely scattered in slightly different
forms.

pectinatum Cav. S. Chile, Valdiv.Fueg. An aberrant species, COPELAND 69. 34.

fatlklandicum Bak. W. Patag., Fueg., Falkl., S. Georgia. Related to peléatum
which, if falklandicum really belongs to Mecodium, where COPELAND placed it
66.94, also must be brought here; felfatum is known from extratropical regions
of both hemispheres and also reported from Chile (perhaps = var. J/enszesz (Pres)
C. Chr. 67.4), by CHRISTENSEN also quoted for Juan Fernandez—he must have
forgotten that in 62.11 all material from there was referred to falklandicum.

rugosum C. Chr. et Skottsb. Related to H. cunbridgense (L.) Sm., W. Ind.,
Venez., Chile, Atl. islands, Mediterr., S. Afr., Austral., Tasm., N. Zeal.

c. Sphacrocionium Copel. A pantrop. and austral genus of about 50 sp., richest
developed in trop. Amer.

ferrugineum Colla. S. Chile, Valdiv.Fueg. Related to the pantropical //. cz-
Ziatum Sw. and belonging to a small austral-tricentric group of closely allied spe-
cies, H. Frankliniae Col. (N. Zeal., probably = ferrugineum, CHRISTENSEN 61. 5),

220

C. SKOTTSBERG

Table 11,

List of species.

Endemics bold-faced,

endemic genera in capital letters.

Adiantum chilense Kaulf.

Asplenium dareoides Desv. .

— obliquum Forst. var.

| — macrosorum Bert. ex Colla
— stellatum Colla

Blechnum auriculatum Cav.

— chilense (Kaulf.) Mett.

— cycadifolium (Colla) Sturm .
— longicauda C. Chr.

— Schottii (Colla) C. Chr. .

— valdiviense C. Chr.
Cystopteris fragilis (L.) Bernh.
Dicksonia berteriana (Colla) Hook. .
— externa Skottsb.

Elaphoglossum Lindenii (Bory) Moore
Gleichenia cf. litoralis (Phil.) C. Chr. .
— pedalis (Kaulf.) Spr.

— quadripartita (Poir.) Moore
Histiopteris incisa (Thunb.) JeSm:
Hymenoglossum cruentum (Cav.) Presl
Hymenophyllum caudiculatum Mart.
— cuneatum Kze .

— falklandicum Bak.

— ferrugineum Colla

— fuciforme Sw.

— pectinatum Cav.

— plicatum Kaulf.

— rugosum C. Chr. et Skottsb.

— secundum Hook. et Grey.

— tortuosum Hook. et Grev. .

Hypolepis rugosula (Labill.) J. Sm.

Lycopodium magellanicum Sw,
— scariosum Forst.
Ophioglossum fernandezianum C. Chr.

| Pellaea chilensis Fée

| Polypodium magellanicum (Desv.)

Arthropteris altescandens (Colla) J. Sm.

Dryopteris inaequalifolia (Colla) C. Chr. .

Lophosoria quadripinnata (Gmel.) C. Chr.

+ +

SC Mf

few."
+ +

DERIVATION OF THE FLORA AND FAUNA 221

Mt SC Mf |

PE NPeEINe MI COllawweener aia deo thas ke) + 6) so els) «sor "ey wr see) el) eaect euets = ane
— lleecallayabinne dees. spehhy c= te Soe Bein Doel Sine A RECra NO NIC a OAC mec Eye oC Ramen i ap = +
— WiagainermG iil a 6 Noe Sa oto) to Mok ceo Hamm mac (cence SC Sur -roa e = | = +
MAC TOMATO’ ES OWia oc rie CUee ne tonas: fos eel ce) side ys (a oa 6 ow wf) se |p ores —
Polystichum berterianum (Colla) C.Chr. ..............{ + =
PROG TTEMUIU( PULSE) ETeES 00. ie sl eae es + es et ee =F — | + |

PUGEG SEO ONIN “gs @ velar 6 tomo uc" «0! ol Bo acolo, BS sarc a hal| ee

=. @OTTeTSTGHIDS A OB sa, cS eS ae A toe eee rine 5 a. _
= scemimadnata ebile . 2... - RMR ee eck ap ah aes eal pete — | +

Ae |
Serpyllopsis caespitosa (Gaud.) C.(\Chr. 2 2 1. ee ee Jae os | +
|

Mio OPDMERUS elegans KZemy se ee es ee we ee aa - +
Tims OMNIES GAGNON TAS 5S <0 Yo 0 By -ol 6 0 Ol io Ge oh GD Oo Go Mole ot a

= lmene C. City se 6 we ame foM Ge & a G20 %0! 6% fol 0 26, “alos OBS Gre) WON = + | = | =
mTaSUSAGI EFI ATUEPENE SCTE (oF. fare oe wee eee re) a ee i Ret ere fee Marie Met | —- | =

aeruginosum (Thouars) Carm. (Tristan da C. and N. Amsterd. I.), and Marlothii
Brause (Cape).

d. Meringium Copel. About 60, S. Chile, Afr., Ceylon, Formosa, N. Guin.,
IN. Zeal:,. Fijt.

secundum Hook. et Grev. S. Chile, Valdiv.—Fueg.

plicatum Kaulf. See 249.763. S. Chile, Valdiv.-W. Patag.

tortuosum Hook. et Grev. S. Chile, Valdiv.—Fueg., Falkl.

Cyatheaceae.

Thyrsopteris elegans Kze. Unanimously regarded as a very ancient type (BOWER,
SEWARD, BERRY, WINKLER, COPELAND, etc.). BERRY (27.88) pointed out that
Cyatheotdes thyrsopteroides Berry is remarkably similar to 7hyrsopteris; it was,
however, found sterile, and he adds that most of the fossils formerly referred to
Thyrsopteris are based on too slender evidence. SEWARD (220. 221) says: “Thyr-
sopteris is very closely allied to certain Jurassic Ferns from the Yorkshire coast
and many other places: geological evidence points to a remote antiquity, and its
present isolation is in all probability the last phase in its history of a direct deriva-
tive of a widely scattered Jurassic type.’ COPELAND called it “a relic from the
time when Décksonia and Cyathea had a common ancestor’ (69. 48) and he thinks
that it is allied to Culcita Presl. He brought 7Ayrsopteris to his large and possibly
very heterogeneous family Pteridaceae, where also Decksonza is placed.

WINKLER’s statement that ZAyrsopteris is found on Masafuera only (287. 472)
is erroneous.

Lophosoria Pres]. One polymorphous species.

guadripinnata (J. F. Gmel.) C. Chr. Trop. and subtrop. Amer., Mex.—Chile and
south to W. Patag., 49° s. lat. The taxonomical status of the various forms re-
mains to be settled. The island form (or forms, for there seems to be some dif-

ZLZ C. SKOTTSBERG

ference between the plants of Masatierra and Masafuera) is certainly unlike the
form inhabiting Chile; see 62. 16.

Dicksonia L’Hérit. 24; 1 Mex., 3 Centr. Amer. to Colomb., 1 Ecuad.—Braz., 2 Peru;
1 St. Helena, 1 Malays., 4 N. Guin., 2 Austra]—Tasm., 3 N. Zeal., 3, Ni @aleam
1 Fiji-Samoa.

berteroana Colla and externa Skottsb. are very near each other and closely
related not to the Andean group but to DY. /anxata Col. from N. Zeal. CHRIST
(59.154) further quotes as relatives D. antarctica Labill. (Austral.), B/amez (Kze)
Moore (Indones., Philipp.), gvazd7s Ros. (N. Guin.) and species from N. Caledonia
and the South Sea islands.

Polypodiaceae.

Cystopteris Bernh. 15; 1 boreal, 10 Ind—China, Japan, N. Zeal., 2 Eur., 1 S. Braz.

fragilis (L.) Bernh. coll. A wide-spread, polymorphous sp., found in all parts
of the world (83° n. lat. to S. Georgia) except Australia. The form occurring on
Masafuera presumably related to a form from the mainland (62. 18).

Dryopteris Adans. Sens u C. Chr. about 1200 sp.; s. str. COPELAND about 150; “an
assemblage of genera that must be segregated in their entirety before of use for
a study of distribution” (COPELAND 69). _

inaequalifolia (Colla) C. Chr. Belongs to a neotrop. group and comes nearer
to tropical forms than to the south Andean D. spectadzlis (Kaulf.) C. Chr., which
extends south to Chile.

Polystichum Roth. About 225; widely spread, numerous in E. As.
berterianum (Colla) C. Chr. Near P. adantiforme Forst., spread through the
5) temp: zone, northivto Wind,
vestitum (Forst.) Presl coll. S. Amer. (also Chile), Austral., N. Zeal.

Arthropteris J. Sm. About 20; palaeotrop., recorded from trop. Afr., north to Arab.;
Madag., Australas., north to Philipp. Is.; New Caled., Fiji, Samoa; best developed
N. Guin., N. Caled. and Madag.

altescandens (Colla) J. Sm. Closely related to Pacific forms (62. 21).

Asplenium . 650-700; cosmopolitan.
obliguum Forst. var. chondrophyllum (Bert.) Mett. Typical od/zguum reported
from S. Chile, Austral. and N. Zeal.
macrosorum Bert. ex Colla. Distantly related to neotrop. sp.; see 62. 23.
stellatum Colla. Belongs to the pantrop. /uzulatum group; see 62. 24.
dareoides Desv. (magellanicum Kaulf.). S. Chile, Fueg., Falkl. Closely related
to A. alvaresense Rudm. Brown of Diego Alvarez (Tristan da C.); see 67. 13.

Blechnum \.. 180-200; essentially southern.

auriculatum Cav. Temp. S. Amer., common in Centr. and S. Chile and related
topbMaustrale le (Sy Atr.).

valdiviense C. Chr. Chile, south to Chiloé. Related to B. /anceolatum (R. Br.)
Sturm from Austral. and N. Zeal. As regards the nomenclature, see 769.54, and
249. 764.

Schottii (Colla) C. Chr. Stands near B. attenuatum (Willd.) C. Chr. (S. Afr.,
E. Austral., Polyn.) and meridense (Kaulf.) C. Chr. (trop. Amer.); see 62. 27.

DERIVATION OF THE FLORA AND FAUNA 223

chilense (Kaulf.) Mett. Centr. Chile to W. Patag.; Falkl. Belongs to the capense
group (S. Afr., Indomal.), see 62.27 and 169. 47.

cycadifolium (Colla) Sturm. Almost too near 2B. magellanicum (Desv.) Mett.
(S. Chile to Fueg. and Falkl.); both related to 2. zabulare (Thunb.) Kuhn (S. and
E. Afr., Madag., Mascaren.).

longicauda C. Chr. Very near 4. Sprucez C. Chr. from trop. S. Amer. (Ecuad.,
Boliv., Braz.), see 250. HICKEN records it from Tucuman in Argent. (733.246),
a statement overlooked also by CHRISTENSEN.
Pellaea Link. About 80, the majority in S. Amer. (south to Chile), S. Afr. and
islands: N. Amer. north to Canada; N. Zeal.

chilensis Fée. Very close to P. xivea (Poir.) Prantl (Ariz.—Chile), see 62. 30.
Hypolepis Bernh. A pantrop. genus of 45-50 sp., half of them trop. Amer., several
Afr. and surrounding islands, others N. Guin—E. As. and Austral.—Polyn.

rugosula (Labill.) J. Sm. Centr. and S. Chile; the typ. sp. Austral., N. Zeal.,
var. villosoviscida (Thouars) C. Chr. (67. 6) on Tristan da C.; Polypodium viscidum
Roxb. from St. Helena very likely is another variety.

Adiantum LL. 200-225; widely distributed but most numerous in S. Amer.
chilense Kaulf. Pert—Chile, Patag., Falkl.

Pteris L.. 270-280, mainly trop. but extending north to S. Eur. and south to S. Afr.,
Tasm. and N. Zeal.

chilensis Desv. Near P. leptophylla Sw. (S. Braz.).

semtadnata Phil. S. Chile, Valdiv—Huafo I. A distinct sp., possibly related
to P. pulchra Schlechtd. from Mexico.

berteroana Ag. Related to P. comans Forst. (Austral., Tasm., N. Zeal.) and
particularly to P. exdlicheriana Ag. (Norfolk 1.).

Histiopteris (Ag.) J. Sm. An assemblage of trop. and austr.-circump. forms which
according to CHRISTENSEN cannot claim to be regarded as different species
(62. 36), whereas COPELAND (69.60) speaks of local, derived species (a concentra-
tion in Indomal.—Polyn.) differing sufficiently from the following polymorphous taxon.

tncisa (Thunb.) J. Sm. Chile, S. Afr., Tristan da C., Austral., Tasm., N. Zeal.,
Polynes.

Polypodium 1. coll. World-wide; Ind. Fil. registers more than 1100 sp. The genus
is now generally broken up, most radically by COPELAND, who segregates numer-
ous genera partly restored from synonymy.

a. Grammitis Sw. About 150; an essentially southern genus, reaching north
to West Indies and Bonin Is. and richest developed in New Guinea.

magellanica Desv. S. Chile, Valdiv._Fueg., And. Patag. (Rio Negro); Tristan
da C., Marion I. In 62.36 as P. Billardieri (Willd.) C. Chr. var. magellanicum
(Desv.) C. Chr., but later restored to specific rank by CHRISTENSEN 67.18. Distri-
bution of Bzllardiert: Austral., Tasm., N. Zeal., Auckl. and Campb. Is., Lord
Howe and Norfolk Is., St. Paul and N. Amsterd. Is., Kerguel.

b. Syxammia Presl. According to COPELAND 69.184 only one species, S.
Feuillet (Bert.) Copel. (Polypodium trilobum Cav.), but I fail to see how we could
exclude the island species.

224 C. SKOTTSBERG

intermedium Colla |translucens (Kze) Fée]. JoHow united P. rzlobum Cav.
and californicum Kaulf. with translucens, and they seem to belong to the same
small group, to which also P. Espznosae Weatherby (Chile, Atacama) must be
referred.

c. Polypodium s. str., about 75 sp., mainly N. hemisph. and neotrop.

Masafuerae Phil. S. Pert to N. Chile (Antofagasta), Argent.; comp. 749.14
and 268.33. Very clese to P. pycnocarpum C. Chr. (Mex.-N. Chile).

d. Xzphopteris Kaulf. About 50, pantrop.

trichomanoides Sw. Trop. Amer. A puzzling record (see 249. 766).

e. Pleopeltis Humb. et Bonpl. About 40, pantrop.; the following sp. wide-
spread.

lanceolatum \.. Mex. and W. Ind. to subtrop. S. Amer. (also in Chile), St.
Helena, Tristan da C., Afr., Madag., Indomal., Hawail.

Elaphoglossum Schott. A pantrop. genus of more than 400 sp., very numerous
in And. S. Amer., 4 sp. on Tristan da C., and many sp. in Polyn. and Hawaii.
Lindentt (Bory) Moore. Mex. to Ecuad. and Braz.

Gleicheniaceae.

Gleichenta Sm. About 130 sp., the majority of them referred to Stcherus (Kaulf.)
Ching, to which also the species in J. Fern. belong; about a dozen sp. spread
over the austral zone; 5 Chile, one extending to Falkl.; 1 5S. Afr., 1 Madag.,
Masearen. and Seych. Is., 1 Tasm., 1 Tasm. and,N: Zeal:, 2 N. Zeal

quadripartita (Poir.) Moore. S. Chile to Fueg.

pedalis (Kaulf.) Spreng. S. Chile, south to Chonos Is.

cf. Zztoral’s (Phil.) C. Chr. Hardly identical with this little known species and
perhaps only a form of G. fedalis.

Ophioglossaceae.
Ophioglossum \.. 28 sp. recognized by. CLAUSEN (64); the genus “scattered with
remarkable uniformity over the habitable globe’? (COPELAND 69. 12).
fernandezianum C. Chr. Not very near O. ypanense Mart. (Colomb., Braz.)
as CHRISTENSEN thought, but close to O. scarzosum Clausen from Pert, Dept. Junin.

Lycopodiaceae.
Lycopodium L. A large world-wide genus, badly ill-treated by NESSEL, who listed
several undistinguishable species in J. Fern.; see 249. 766.
magellanicum Sw. S. Chile to Fueg., Falkl., S. Georgia, Marion I., Kerguel.
scartosum Forst. Chile, Valdiv.-Guaytecas Is. (ZL. gayanum Remy), N. Zeal.;
Trop. Andes, Braz. (L. Fusszeuz Desv.). See |.c. 767.

In his treatment of the geographical groups distinguished by him, JOHOW
included the ferns—no Fern Allies were at that time known from Juan Fernandez.
All were classified as American except 3, Dicksonia berteroana, said to be similar
to a species from Fiji, Preris berteroana (identified with comans), and “Asplenium
longissimum Blume’, the species now known as Blechnum longicauda. In 1914
(227) I went a step further; to my ‘“‘Altpazifisches Element” 5 species were referred,
to the Trop. American 5, and all the rest to the Chilean. In 1934 I attempted
a more detailed subdivision (239, here translated from French).

DERIVATION OF THE FLORA AND FAUNA 225

1. Neotropical and Andean element. Species either found in Centr. and S. Chile or
endemic but allied to Chilean species; 28.
a) in Chile, 22.
b) in the Magellan. region, but not of subantarctic character. 3.
d) endemic species of non-endemic genera, 3.
Element consisting of wide-spread species, also inhabiting Chile, 2.
3. Neotropical element, not represented in Chile, 7.

a) non-endemic species, 1.

b) endemic species of non-endemic genera, 6.
4. Magellanian-Old Antarctic element, 9.

a) non-endemic Magellanian species, 2.

By o%55 * Old Antarctic but not Magellanian species, 4.
c) endemic species allied to subantarctic species, 2.
d) & E. not allied to subantarctic species, 1.

Peace element, 5.
a) isolated endemic species of wide genera, 1.
b) endemic species belonging to genera or sections of west to central Pacific distribu-
tion, 3.
d) species of endemic genus with unknown relations, 1.

A modified arrangement is attempted below, corresponding to the analysis
of the angiosperms.

I. Andine-Chilean element.—34 (64.1 %).

a. Endemic species (5, 9.4%): Hymenophyllum rugosum, Polystichum berteri-
anum, Blechnum cycadifolium, Pellaea chilensis, Polypodium intermedium.

6. Known from continental Chile, many also in other parts of S. America
or of still wider distribution (29, 54.7%): Trichomanes exsectum, Hymenoglossum
cruentum, Hymenophyllum cuneatum, caudiculatum, fuciforme, pectinatum, ferru-
gineum, secundum, plicatum and tortuosum, Lophosoria quadripinnata, Cystopteris
fragilis, Polystichum vestitum, Asplenium obliquum, Blechnum auriculatum, valdi-
viense and chilense, Hypolepis rugosula, Adiantum chilense, Pteris chilensis and
semiadnata, Histiopteris incisa, Polypodium magellanicum, Masafuerae and lanceo-
latum, Gleichenia quadripartita, pedalis and cf. litoralis, Lycopodium scariosum.

II. Subantarctic-Magellanian element.— 4 (7.5 %).

Not endemic, extending north but not beyond Centr. Chile: Serpyllopsis caes-
pitosa, Hymenophyllum falklandicum, Asplenium dareoides, Lycopodium magel-
lanicum.

Ill. Neotropical element.—g9 (17 %).

a. Endemic species (7, 13.1%): Trichomanes philippianum and Ingae, Dryo-
pteris inaequalifolia, Asplenium macrosorum and stellatum, Blechnum longicauda,
Ophioglossum fernandezianum.

6. Not endemic (2, 3.8 %): Polypodium trichomanoides, Elaphoglossum Lindenii.

IV. West Pacific element.—5 (9.4 %).

Affinities in Australia, New Zealand and Oceania, not in America. All en-
demic: Dicksonia berteroana and externa, Arthropteris altescandens, Blechnum
Schottii, Pteris berteroana.

15 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

226 C. SKOTTSBERG

V. Eu-Fernandezian element.—1 (1.9 %).

Endemic genus without affinities among the living ferns: Thyrsopteris elegans.

As could be expected, the Chilean element is stronger among the ferns than
among the angiosperms, followed in great distance by the neotropical, which also
is comparatively stronger, whereas the Magellanian and the Pacific are less im-
portant, especially the latter; long-distance advocates would expect the opposite,
because ferns ought to travel across the ocean much more readily than flowering
plants.

Ill. Musci.

The following list is based on BROTHERUS’ paper in vol. II (34) and on his
treatment of the group in the 2nd editicn of Natirl. Pflanzenfam. In many cases
the figure for the number of species (in brackets) is too low, as numerous mosses
have been described in later years, but I have refrained from searching the litera-
ture. Much information was obtained from HERZOG’s work (729) and in pt. 2 of
IRMSCHER’s book (743). Iam indebted to Dr. HERMAN PERSSON for kind assistance.

Mt = Masatierra, Mf = Masafuera.

Ditrichaceae.

Pleuridium Brid. (about 30). Wide-spread, mainly temperate.
Robinsonii (Mont.) Mitt. Chile, Urug. Belongs to a S. Amer. group.—Mt.

Ditrichum Vimm (about 50). In all parts of the world.

affine (C.M.) Hampe. S. Chile and Patag., E. Austral., N. Zeal., Auckl. Is.
Not listed in 270, where D. elongatum (H. f. et W.) Mitt. is quoted for Chile, Austral.,
Tasm. and N. Zeal.—Mt, Mf.

longisetum (Hampe) Jaeg. S. Chile to Fueg.—Mf.

Ceratodon Brid. (2, one trop.).
purpureus (L.) Brid. Cosmopol.—Mt, Mf.

Pottiaceae.

Hymenostomum R. Br. (50-60). All over the world.
kunzeanum (C.M.) Broth. S. Chile.-—Mt.

Gymnostomum Hedw. (10). All over the world.

calcareum Bryol. germ. Eur., Afr., As., N. Amer., S. Amer.: Ecuad.—Chile;
Austral., Tasm., N. Zeal.—Mt, Mf.
Trichostomum Hedw. (81). Cosmopolitan.

brachydontium Bruch. Eur., Caucas., N. Afr., Macaron., Mascar., Japan, N.
Zeal., but not mentioned in 2z70.—Mf.

Dicranaceae.

Amphidium Nees (10). Very widely distributed.
cyathicarpum (Mont.) Broth. Ecuad.—-Chile, S. Georgia, Afr., E. Austral., Tasm.,
N. Zeal.—Mt, Mf.

DERIVATION OF THE FLORA AND FAUNA 227

Dicranella Schimp. (60). All continents, numerous S. Amer. and As.
costata Broth.—Mt, Mf.

Oncophorus Brid. (8). Southern S. Amer., Eur., E. As., Ceylon.
fuegianus Card. Patag—Fueg.—Mf.

Dicranoloma Ren. (76). Almost exclusively S. hemisph., austr.-subantarct., 1 S. Afr.
fernandezianum Broth. Near PD. Duseniz Broth. (S. Chile).—Mt.
capillifolium Broth. S. Chile-Fueg.—Mt.
capillifolioides Broth. Near the former.—Mf.

Menzies (Tayl.) Par. S. Chile, E. Austral., Tasm., Norfolk I., N. Zeal.,

Chatham Is., Auckl. Is.—Mt.

Billardiert (Schwaegr.) Par. Pert-Fueg., Falkl., S. Afr., Austral., Tasm., N.

Zeal., Auckl. and Campb. Is.—Mt, Mf.
nigricaule Angstr. S. Chile, south to Fueg.—Mf.

Campylopus Brid. (about 500). All over the world, mainly trop.-subtrop.
introfiexus (Hedw.) Mitt. N. Amer., S. Amer., Ecuad. to Urug. and Fueg.,

islands of W. and E. Afr., Ascens., St. Helena, Tristan da C., Marion I., Austral.,

Tasm., N. Zeal., Auckl., Campb. and Antipodes Is., Pacific islands.—Mt.
punacams ©. M. Chile-—Mt.
polytrichoides De Not. W. and S. Eur., N. Afr., Mascar.—Mt.
aberrans Broth. A very peculiar sp.—Mt, Mf.
areodictyon (C.M.) Mitt. Centr. and S. Amer., Venez.—Boliv.—Mf.
subareodictyon Broth. Related to the former.—Mf.
blindioides Broth.—Mf.

Thysanomitrium Schwaegr. (31). Trop.-subtrop., especially Old World, extending

south to the austr.-subantarct. zone.

Richardt Schwaegr. Centr. and S. Amer. to Chile.—Mt, Mf.
leptodus (Mitt.) Broth. [Campylopus clavatus (R. Br.) H. f. et W.| Ecuad., Chile,
Austral., Tasm., N. Zeal., Auckl. and Campb. Is—Mt, Mf.

Dicnemonaceae.

Eucamptodon Mont. (7). Chile (1), Austral. (1), N. Caled. (4), N. Zeal. +N. Caled. (1).
perichaetialis Mont. S. Chile to Magell.—Mf.

Fissidentaceae.

Fissidens Hedw. (over 700). All continents, very numerous in trop. zones.

fernandezianus Broth.—Mt, Mf.

crassicuspes Broth. Related to F. crassipes Wils. (Eur. Mediterr., Madeira).—Mt.

rigidulus Hook. f. et Wils. Ecuad._W. Patag., Austral., Tasm., N. Zeal.—
Mt, Mf.

leptochaete Dus. Chile.—Mt.

maschalanthus Mont. Chile, south to W. Patag.—Mt, Mf.

pycnotylus Broth. Very near the former.—Mt.

asplenioides (Sw.) Hedw. Centr. Amer. to Pert, S. Chile and Braz., W. Afr.,
Macaron., Tristan da C., Indomal., Queensl., N. Zeal.—Mt.

228 C. SKOTTSBERG

Leptodontium Hampe (80). Most numerous in Amer., 40% And.
fernandezianum Broth. Related to L. luteum (Tayl.) Mitt. (trop. And.).—Mf.

Didymodon Hedw. (91). Subcosmopol., mainly temp., essentially Amer.: Mex.—
Centr. Amer.—Pert and Chile; 1 Antarct.

calymperidictyon Broth.—Mt.

linearis Broth.—Mf.

Tortula Hedw. (220). Subcosmopol., mainly temp.
scabrinervis (C.M.) Mitt. Chile-—Mt.
flagellaris (Schimp.) Mont. Chile.— Mt.

Grimmia Ehrh. (230). Subcosmopol. but rare in the trop. zones.
phyllorhizans Broth.-——Mt.

Rhacomitrium Brid. (80). As Grimmia.

subnigritum (C.M.) Par. W. Patag.—Fueg.—Mf.

symphyodontum (C.M.) Jaeg. S. Chile to Fueg. and Falkl., S. Afr., Kerguel.,
Tasm., N. Zeal. Not listed for N. Zeal. in 270.—Mt, Mf.

striatipilum Card. S. Chile to Fueg., S. Georgia; N. Zeal. (2z0).—Mf.

lanuginosum (Hedw.) Brid. Cosmopol.—Mf.

loriforme Dus. W. Patag.—Mf.

convolutum Mont. S. Chile to W. Patag.—Mf.

Ptychomitriaceae.

Ptychomitrium (Bruch) Fiirnr. (62). Of wide distribution in temp. zones.
fernandezianum (Mitt.) Jaeg.—Mt, Mf.

Orthotrichaceae.

Zygodon Hook. f. et Tayl. (about 100). N. and Centr. Amer. (6), trop. S. Amer.
(40), S. Chile and Argent. (20), Eur. (4), trop. Afr. (2), trop. As. (6), E. Austral.—
N. Zeal. (10).

intermedius B. et S. S. Amer., Afr., Monsoon reg., Austral., Tasm., N. Zeal.
—Mt. (264). Z. obovalis Mitt., a doubtful species (373. 139), is supposed to be iden-
tical with zz~fermedius.

Menzies (Schwaegr.) W. Arn. S. Chile, Austral., Tasm., N. Zeal.—Mt.
Stenomitrium (Mitt.) Broth. (2). Peru, Chile.

pentastichum (Mont.) Broth. Peri-W. Patag.—Mf.
Ulota Mohr (43). N. and S. temp. zones, 1/3 Chile, some N. Zeal.

fernandeziana Malta (373). Near U. rufula (Mitt.) Jaeg. (Chile-W. Patag.,
Argent., N. Zeal.).—Mf.
Macromitrium Brid.(415). Trop.-subtrop., special groups Chile—Patag. and E. Austral.—
Tasm—N. Zeal.

hymenostomum Mont. S. Chile-Fueg.—Mt.

saxatile Mitt.—Mt, Mf.

fernandezianum Broth. Related to I/. asperulum Mitt. (Tasm., N. Zeal.; not in
210).—Mt.

Masafuerae Broth. Near the former.—Mf.

DERIVATION OF THE FLORA AND FAUNA 229

Funariaceae.
Funaria Schreb. (200). A world-wide genus.
hygrometrica (L.) Sibth. Cosmopol.—Mt, Mf.

Bryaceae.
Mielichhoferia Hornsch. (100). Widely distributed, centering in the Andes.
longiseta C. M. Ecuador.—Mt.
Bryum Dill. (about 800). All over the world.
Lechlert C. M. Chile.—Mf.
fernandezianum Broth. Related to B. Cruegert Hampe (neotrop.).—Mt, Mf.

Leptostomaceae.

Leptostomum R. Br.(11). S. Chile (2), Indomal., Austral., N. Zeal., Norfolk I., Campb. I.
Menziesti (Hook.) R. Br. S. Chile to W. Patag. and Fueg.— Mf.

Eustichiaceae.

Eustichia (Brid.) Mitt. (8). 6 Mex.-S. Chile, 1 S. Afr., 1 islands of E. Afr.
Poeppigtt (C. M.) Par. S. Chile-Magell.—Mt, Mf.

Rhizogoniaceae.
Rhizogonium Brid. (27). Austr.-circump. (S. Amer., S. Afr., Austral., N. Zeal., Polyn.),
extending into the N. hemisph.; 1 pantrop.
Novae Hollandiae Brid. var. patagonicum Card. et Broth. W. Patag.; the typical
speeustral., Lasm., N. Zeal:-—Mt.
mntotdes (Hook.) Schimp. Colomb.-Fueg., E. Austral., Tasm., N. Zeal.—Mf.

Bartramiaceae.

Anacolia Schimp. (7). W. N. Amer.-trop. Andes (5), 1 Medit., 1 Ethiop.
subsessilis (Yayl.) Broth. Mex._Ecuad.—Mt, Mf.
Bartramia Hedw. (110). Subcosmopol., a few sp. subantarct.-circump.
aristata Schimp. Chile.—Mt, Mf.
patens Brid. W. Patag.—Fueg., Falkl., S. Georgia, Kerguel., E. Austral., Tasm.,
N. Zeal.—Mf. Not mentioned in 270, where B. halleriana Hedw., widely distributed
in the N. hemisphere, is credited to Patag., Fueg., Austral., Tasm. and N. Zeal.—Mf.
fernandeziana Card.—Mt.
Philonotis Brid. (174). Subcosmopol., two monotyp. sections in S. Amer.
krauseana (C. M.) Jaeg. Centr. Chile to W. Patag.—Mt, Mf.
glabrata Broth. Related to the former.—Mt.
scabrifolia (Hook. f. et Wils.) Broth. Circump.-subantarct., forming a separate
section. Ecuad—Fueg., Falkl., S. Georgia, S. Afr., Marion I., Kerguel., Austral.,
iiasni.,..N:. Zeal,, Avekl and -Campb: is:—Mt; Mf,
vagans (Hook. f. et Wils.) Mitt. Forms a separate section. Chile-Fueg., 5.
Georgia.—Mf (an aberrant form).
Breutelia Schimp. (104). Best developed trop. mountains, rare N. Amer., numerous
austr.-subantarct.
Masafuerae Broth.—Mf.

230 C. SKOTTSBERG

Hedwigiaceae.

Rhacocarpus Lindb. (24). Centr. Andes and Braz. mountains etc.; almost confined
to the S. hemisph.

Humboldt (Hook.) Lindb. Mex., W. Ind., Colomb—And. Patag., Fueg., Falkl.,
Centr. Afr., Madag., Réunion, Austral., Tasm., N. Zeal. A polymorphous sp.—Mf.

Cryphaeaceae.

Dendrocryphaea Par. et Schimp. (4). Austr.-subantarct., 2 Chile, 1 Argent.—Patag.,
1 Tasm.-N. Zeal.

cuspidata (Sull.) Broth. S. Chile.—Mf.
Cyptodon Par. et Schimp. (6). 2 Austral—N. Zeal., 1 N. Caled., 2 Fiji-Samoa—Tonga,
La) epeiern.

crassinervis Broth. Close to C. (2z0 under Cryphaea) dilatatus (Hook. f. et
Wils.) Par. (E. Austral., N. Zeal.).—Mf.

Lepyrodontaceae.

Lepyrodon Hampe (6-8). 1 bicentric (Chile-Fueg., E. Austral., Tasm., N. Zeal.,
Campb, I.), 2 or 3 trop. Andes to Braz. and Argent., 2 or 3 Chile sieNeewear
parvulus Mitt. Centr. and S. Chile.—Mt, Mf.
tomentosus (Hook.) Mitt. Colomb.—Peru, Braz., Argent., W. Patag.—Mt.

zmplexus (Kze) Par. Chile, south to Fueg.—Mt.

Ptychomniaceae.

Ptychomnium Hook. f. et Wils. (9, or 8 if evyguisetum = aciculare). Austral-subant-
arct., extending north to Braz. and Hawaii; Argent. and Chile to Fueg., E. Austral.,
Tasm., N. Zeal. and subant. islands, Lord Howe I., N. Caled., Polyn.

subaciculare Besch. Chile, south to Fueg.—Mt.

falcatulum Broth. Related to the former.—Mt.

ptychocarpum (Schwaegr.) Mitt. Chile, south to W. Patag.—Mf.

Neckeraceae.

Weymouthia Broth. (3-4). Austr.-circump., Hawaii.
mollis (Hedw.) Broth. Centr. and S. Chile to Magell., E. Austral., Tasm.,
N. Zeal.—Mt, Me.

Leptodon Mohr (4). Afr. and islands (3), and the following.
Smithii (Dicks.) Mohr. Argent., Chile; S. Eur., Caucas., Afr., Macaron.; E.
Austral., N. Zeal.—Mt, Mf.

Neckera Hedw. (127). Widely distributed trop.-subtrop., extending south to Chile
and N. Zeal.—Auckl. I.
rotundata Broth. A very distinct sp.——Mf.

Porothamnium Fleisch. (51). 1 W. N. Amer., 41 Centr. and S. Amer., extending
southito!Patasy,..7..trop. sAfcy1 10 Geylon,yim N.. Zeal:

fasciculatum (Sw.) Fleisch. W. Ind., Colomb.—Pert, Braz.—Mf.

arbusculans (C. M.) Broth. Chile, Patag.—Mf.

DERIVATION OF THE FLORA AND FAUNA 231

Thamnium Bryol. eur. (33). N. Amer., W. and S. Eur., Macaron., S. Afr., N. and
E. As., Indomal., Austral., N. Zeal., Melan. The Juan Fernandez species belong
to a southern section.

rigidum (Mitt.) Broth.—Mt.

latinerve (Mitt.) Broth.——Mt.

Caroli Broth. Related to rz7g7dum.—Mt.

Ingae Broth. Very near the former.—Mt.

crassinervium (Mitt.) Broth.—Mt.

proboscideum Broth. Near the former.—Mt.

assimile Broth. Very near the former.—Mt.

confertum (Mitt.) Broth.—Mt.
Pinnatella (C. M.) Fleisch. (37). Pantrop.

macrosticta Broth. A very distinct sp.—Mf.

Hookeriaceae.

Distichophyllum Dozy et Molk. (93). Essentially austral, extending south to Austral.,
Tasm., N. Zeal.; well represented Indomal. region.

subelimbatum Broth.—Mt.

assimile Broth. Near the former.—Mt.

fernandezianum Broth. Related to D. rotundifolium (Hook. f. et Wils.) Broth.
(Chile, Patag., E. Austral., Tasm., N. Zeal.).—Mf.
Pterygophyllum Brid. (32). Mainly S. hemisph., 6 S. Chile-Fueg., 25 Austral—Tasm.—
Ne Zeal. 2 of these also S.. Amer.

anomalum (Schwaegr.) Mitt. Fueg.—Mt. Perhaps only an extreme water form
of the following (z7z. 85)

obscurum (Mont.) Mitt. S. Chile to W. Patag., Falkl., Tasm.—Mf.

tenuinerve Broth. Very near the former.—Mt.

denticulatum (Hook. f. et Wils.) Mitt. W. Patag.—Fueg., Falkl., E. Austral.,
Tasm., N. Zeal., Auckl. and Campbell Is.—Mt. Not included in 2z0, where
P. dentatum (Hook. f. et Wils.) Mitt. is supposed to occur in Chile and Fuegia.
Eriopus (Brid.) C. M. (25). Some trop. Andes, 1 S. Afr., numerous N. Zeal.

leptoloma Broth. Related to £&. apiculatus (Hook. f. et Wils.) Mitt. (S. Chile-
Fueg., Austral., Tasm., N. Zeal.).—Mt, Mf.

grandiretis Broth. Near the former.—Mf.
Lamprophylum Schimp. Monotypical; more or less distantly related to neotrop.
genera.

splendidissimum (Mont.) Schimp. S. Chile and W. Patag.—Mf.

Hypopterygiaceae.

Lopidium Hook. f. et Wils. (16). 14 paleotrop., 2 S. Amer. (Braz., Chile).
conciunum (Hook.) Fleisch. Chile, south to W. Patag., Austral., Tasm., N. Zeal.,

Auckl. Is.—Mt.

Hypopterygium Brid. (61). Widely distributed N. and S. Amer., Afr., As., Indomal.,

one small group austr.-bicentr.

232 C. SKOTTSBERG

Thouini (Schwaegr.) Mont. S. Chile-Fueg. Belongs to a group of 4 sp. (3 Chile,
1 N. Zeal.).—Mf.

Rhacopilaceae.

Rhacopilum Palis. (51). Pantrop., mainly southern, south to N. Zeal.
fernandezianum Card. S. Chile. Related to R. tomentosum (Sw.) Brid. (trop.—
subtrop. Amer.).—Mt, Mf.

Thuidiaceae.

Thuidium Bryol. eur. (161). All over the world, especially in humid mountain climates.
Masafuerae Broth. Related to 7. fulvastrum (Mitt.) Jaeg. (Tristan da C.,
N. Zeal. but not included in 270) and to the following sp.—Mf.
Valdiviae Broth. S. Chile.—Mt, Mf.

Amblystegiaceae.

Sciaromium Mitt. (22). Mainly austral, the majority southern S. Amer., 1 Cape, 2
China, 2 E. Austral., 1 N. Zeal.; the single N. Amer. sp. forms a separate section.
Sect. Aloma 6 (1 Boliv., 2 Fueg., 2 E. Austral., 1 N. Zeal.).

pachyloma (Mont.) Par. S. Chile-W. Patag.—Mt, Mf.

Hypnaceae.
Stereodon Mitt. (7). 2 Mex., 1 Chile, 4 mountains of Asia.
Lechlert (C. M.) Mitt. Centr. Chile to W. Patag.—Mt.
Isopterygium Mitt. (170). Widely distributed, preponderately trop.—subtrop.
fernandezianum Broth. Related to /. cexerum (Sw.) Mitt. (neotrop.).—Mt.

Sematophyllaceae.

Rhaphidostegium (Bryol. eur.) De Not. (106). Subcosmopol., trop. and temp. The
Chilean species sometimes regarded as belonging to a separate genus Xhaphido-
rhynchiume.

Masafuerae Broth. Similar to 2. cyparisstoides (Hornsch.) Besch. (Braz.).—Mf.

aberrans Broth. Similar to A. callidum (Mont.) Jaeg. (S. Chile to W. Patag.).—
Mt, Mf.

caespitosum (Sw.) Jaeg. W. Ind., trop.—subtrop. S. Amer.—Mt, Mf.

caespitosoides Broth. Related to the former.—Mf.

brachycladulum Broth.—Mt.

Rigodium Kze. (19). Trop.-subtrop. Amer., 2 Afr.
toxarium (Schwaegr.) Schimp. Trop. S. Amer. to W. Patag.—Mt, Mf.
arborescens (C. M.) Broth. S. Chile-W. Patag.—Mt, Mf.
Aylocomioides Card. et Broth. Patag.—Mf.
robustum Broth.—Mt.
Looseri Thér. Related to R. gracile Rén. et Card. (Costa Rica).—Mt.
tamarix C. M. (elegantulum Card.). S. Chile-W. Patag.—Mt.
Rhynchostegium Bryol. eur. (130). Temp.-subtrop., almost cosmopol.
complanum (Mitt.) Jaeg. Centr. Chile—Mt, Mf.
tenuifolium (Hedw.) Jaeg. Urug., S. Chile, E. Austral., Tasm., N. Zeal.—Mt.

DERIVATION OF THE FLORA AND FAUNA 233

Catagoniopsis Broth. Monotypical, related to Catagoninm (S. hemisph.)
berteroana (Mont.) Broth. Centr. Chile.—Mt.

Hypnodendraceae.

Hypnodendron Lindb. (28). Chile (2), Indomal., Austral., Polyn., Hawaii.
Mf.

microstictum Mitt. Chile.

Polytrichaceae.

Oligotrichum Lam. et DC. (13). Widely scattered: 2 W. N. Amer., 4 Chile; single
Sieoraz., trop. Andes, Arct.-alp., Eur., bor.-cireump.

canaliculatum (Hook.) Mitt. var. The typical sp. in S. Chile.—Mf.
Psilopilum Brid. (17). A bipolar genus, tricentric in the S. hemisph.

antarcticum C. M. Boliv., Fueg., Falkl., S. Georgia, Kerguel.—Mf.
Polytrichadelphus (C. M.) Mitt. (22). Centering in the Andes (17), 1 W. N. Amer.,
1 Braz., 2 Magell., 1 subant.-bicentric.

magellanicus (L.) Mitt. Patag., Fueg., Falkl., E. Austral., Tasm., N. Zeal., Auckl.
and Campb. Is.—Mf.
Dendroligotrichum (C. M.) Broth. Monotypical.

dendroides (Brid.) Broth. Peru, S. Chile south to Fueg., N. Zeal.—Mf.

The geographical distribution of the mosses is less well known than of the
vascular plants, and some of the species now considered to be endemic in Juan
Fernandez will perhaps be discovered on the mainland. No specialist ever visited
the islands, where most likely further species, endemic or known from elsewhere,
will be found. Species now only recorded from Masatierra may be found on Masafuera,
and vice versa.

The 131 species of mosses hitherto recorded from Juan Fernandez belong to 65
genera (2:1), all known from elsewhere; 48 species (36.6 %) are endemic, a high
figure in a spore-bearing group, higher than for the Pteridophytes (34%). On Masa-
tierra 84! were collected, 50 of these (38.1%) not found on Masafuera, where 81
were recorded, of which 47 (35.9 %) are restricted to this island; 34 (26 %) are known
from both islands. No mosses have been reported from Santa Clara.

The mosses are less evenly distributed than the ferns, presenting a higher
degree of local endemism, but this may be due to insufficient knowledge of their
distribution. Endemics are more local than non-endemics; of the former (48), 24
(50 %) are only known from Masatierra, and 16 (33.3 %) only from Masafuera, 8 (16.7 %)
being found on both islands; of the non-endemics (83), 26 inhabit Masatierra (31.3 %),
31 Masafuera (37.4%) and 26 (31.3 %) both islands. Endemics are proportionately
more numerous on Masatierra, where 32 species or 38 % are endemic; the figures for
Masafuera are 24 and 29.6. It is remarkable that all the 8 species of 7amnium
are endemic on Masatierra.

For two reasons it is a difficult task to segregate with sufficient accuracy the

1 BROTHERUS p. 420 records Rhacomitrium subnigritum from both islands, but the locality
quoted for Masatierra is situated on Masafuera.

234 C. SKOTTSBERG

various geographical elements of which the flora is made up: many of the non-
endemic species have a more or less wide distribution, and the accurate systematic
position of the endemic ones is, in many cases, uncertain or quite unknown; where
no information was given by their author and nobody has studied them after him,
I have not ventured to find a place for them. Thus 11 species had to be left out,
bringing the number down to 120, on which the percentages have been calculated.
Even so it stands to reason that the arrangement below cannot be definite because
many doubtful points remain to be cleared by the bryologist.

As to the 83 non-endemic species the main thing is whether, regardless of
their total distribution, they have been found in S. America or not; in consequence
of the geographical position of Juan Fernandez they must be referred to one of
the American groups I-III. No less than 25 % are austral or subantarctic and
bicentric; their origin, if antarctic or not, will not concern us here.

The geographical elements.
I. Andine-Chilean element.—77 (64.2%).

a. Endemic species (15): Dicranoloma fernandezianum and capillifolioides, Fissi-
dens pycnotylus, Didymodon calymperidictyon and linearis, Ulota fernandeziana,
Philonotis glabrata, Ptychomnium falcatulum, Distichophyllum fernandezianum,
Pterygophyllum tenuinerve, Eriopus leptoloma and grandiretis, Thuidium Masa-
fuerae, Rhaphidostegium aberrans, Rigodium robustum.

6. Also known from Chile, rarely extending to the extreme south (57):
Pleuridium Robinsonii, Ditrichum affine and longisetum, Hymenostomum kunzeanum,
Amphidium cyathicarpum, Dicranoloma capillifolium, Menziesii and nigricaule,
Campylopus truncatus, Thysanomitrium Richardi and leptodus, Eucamptodon
perichaetialis, Fissidens rigidulus, leptochaete, maschalanthus and asplenioides,
Tortula scabrinervis and flagellaris, Rhacomitrium loriforme and convolutum, Zygodon
intermedius and Menziesii, Stenomitrium pentastichum, Macromitrium hymeno-
stomum, Bryum Lechleri, Leptostomum Menziesii, Eustichia Poeppigii, Rhizogonium
mnioides, Bartramia aristata, Philonotis krauseana and vagans, Rhacocarpus Hum-
boldtii, Dendrocryphaea cuspidata, Lepyrodon parvulus and implexus, Ptychomnium
subaciculare and ptychocarpum, Weymouthia mollis, Leptodon Smithii, Porotham-
nium arbusculans, Lamprophyllum splendidissimum, Lopidium concinnum, Hypo-
pterygium Thouini, Rhacopilum fernandezianum, Thuidium Valdiviae, Sciaromium
pachyloma, Stereodon Lechleri, Rigodium toxarium, arborescens, hylocomioides
and tamarix, Rhynchostegium complanum and tenuifolium, Catagoniopsis berte-
roana, Hypnodendron microstictum, Oligotrichum canaliculatum, Dendroligotrichum
dendroides.

c. Cosmopolitan (5): Ceratodon purpureus, Gymnostomum calcareum, Cam-
pylopus introflexus, Rhacomitrium lanuginosum, Funaria hygrometrica.

II. Subantarctic-Magellanian element.—13 (10.8 %).
All non-endemic, found in the far south, many going north to the latitude
of Valdivia or even farther; several occur on the Falkland Is., South Georgia or
other subantarctic islands, and not few reappear in New Zealand, etc.

DERIVATION OF THE FLORA AND FAUNA 235

Oncophorus fuegianus, Dicranoloma Billardieri, Rhacomitrium subnigritum,
symphyodontum and striatipilum, Rhizogonium Novae Hollandiae, Bartramia patens,
Philonotis scabrifolia, Pterygophyllum anomalum, obscurum and denticulatum,
Psilopilum antarcticum, Polytrichadelphus magellanicus.

III. Neotropical element.—14 (11.7 %).

Recorded from tropical America but not from Chile, or related to tropical
species.

a. Endemic (8): Campylopus subareodictyon, Leptodontium fernandezianum,
Bryum fernandezianum, Pinnatella macrosticta, Isopterygium fernandezianum, Rha-
phidostegium Masafuerae and caespitosoides, Rigodium Looseri.

6. Non-endemic (6): Campylopus areodictyon, Mielichhoferia longiseta, Anacolia
subsessilis, Lepyrodon tomentosus, Porothamnium fasciculatum, Rhaphidostegium
caespitosum.

IV. West Pacific element.—13 (10.8 %).

Endemic species, allied to S.W. Pacific species (Australia, Tasmania, New Zea-
land etc.) but, as far as known, not to S. American species: Macromitrium fer-
nandezianum and Masafuerae, Cyptodon crassinervis, Distichophyllum subelimbatum
and assimile, Thamnium rigidum, latinerve, Caroli, Ingae, crassinervium, probosci-
deum, assimile and confertum.

V. Atlantic element.-—3 (2.5 %).

a. Endemic: Fissidens crassicuspes.
6. Not endemic: Trichostomum brachydontium, Campylopus polytrichoides.

Endemic species of unknown position: Dicranella costata, Campylopus aberrans
and blindioides, Fissidens fernandezianus, Grimmia phyllorhizans, Ptychomitrium
fernandezianum, Macromitrium saxatile, Bartramia fernandeziana, Breutelia Masa-
fuerae, Neckera rotundata, Rhaphidostegium brachycladulum.

The dominance of a South American element is self-evident; groups I to III
make up 86.7 %. It is hard to draw a line between I and II; the Magellanian species
are supposed to have come from the far south, and theirs is a more southern area,
but many of the species referred to 16 may have had the same history though,
at present, they do not reach so far south.

The Atlantic element is artificial. /Zss7dens crassicuspes is, if BROTHERUS is right,
related to a species that has its nearest station on Madeira, but the genus is a very
large one, and another connection may be found. Campylopus polytrichotdes is an
Atlantic species with its nearest locality on Madeira; its presence on Masatierra
is indeed surprising. 77r7chostomum brachydontium is scattered over half the globe,
with its nearest stations in Macaronesia, but it extends not only to and beyond
the Mediterranean region, but turns up on the island of Réunion, in Japan and
on New Zealand. Have we to do with isolated remnants of a once more continuous
area, or is it still to be discovered in other places? Is it a bipolar species?

236 C. SKOTTSBERG

Of the species left aside for the present, Décranella costata (many in S. Amer.),
Ptychomitrium fernandeszianum (other sp. in the Andes), Wacromitrium saxatile (many
in S. Amer.), Bartramia fernandeziana and Breutelia Masafuerae (many austral sp.)
may turn out to belong to an American element. The occurrence in Masatierra
of 8 endemic species of 7/amnzum is astonishing, for not one is quoted for South
America, whereas related species are found in Australia-New Zealand, Oceania and
Malaysia; for this reason I have referred the species in Masatierra to the Pacific
element.

IV. Hepaticae.

No handbook equal to BROTHERUS’ comparatively modern account of the mos-
ses exists of the Hepaticae. SCHIFFNER’s treatment of this group in the Ist edition of
Naturl. PAanzenfamilien is too antiquated to be of much use. Much important infor-
mation is, however, found in HERZOG’s work (729) as well as in DOMIN’s paper (76),
and also in this case Dr. PERSSON kindly helped me, but to search the voluminous
special literature of the last thirty years was not to be thought of. The number of
species given 1s, in many cases at least, too low, but I don’t think this matters
very much.

The following list is based on EVANS’ and HERZOG’s papers (93, 730) with
a few alterations (73z). Several species credited to Juan Fernandez by STEPHANI
(333) but not mentioned by HERZOG are included here; some of them are, perhaps,
identical with other species.

Marchantiales.

Plagiochasma WLehm. et Lindenb. (about 20). Mostly trop.—subtrop.
vupestre (Forst.) St. Cosmopol., also Chile-—Mt, Mf.

Reboulta Raddi (1).

hemisphaerica (L.) Raddi. Cosmopol., also Chile.-—Mt, Mf.
Lunularia (Mich.) Adans. (1).

cructata (L.) Dumort. S. Amer., also Chile, Medit. Eur., Atl. islands, Afr.,
Austral.—Mt, Mf.

Marchantia L. (about 50). Subcosmopol., numerous trop.

polymorpha . Cosmopol., also Chile.—Mt, Mf.

berteroana Lehm. et Lindenb. Chile south to Fueg., Falkl., St. Helena, S. Afr.,
Marion I., Kerguel., Austral., Tasm., N. Zeal.—Mt, Mf.

foliacea Mitt. S. Chile, Tasm., N. Zeal.—Mt, Mf.

Metzgeriales.

Riccardia S.F. Gray (140-150, but much higher figures are given). N. temp. 6,
trop. Amer. 43, Afr. 14, trop. As.—Oceania 53, austr.-subantarct. 35.

Juegiensis Massal. S. Chile to Fueg.—Mf.

breviramosa (St.) Evans. Falkl.—Mt, Mf.

adglutinata Evans.—Mt, Mf.

zmsular7s Schiffn. St. Paul and New Amsterd. Is.—Mt.

variabilis Evans. S. Chile.—Mt.

DERIVATION OF THE FLORA AND FAUNA 237

leptostachya Evans. Related to the former.—Mt.

nudimitra (St.) Evans. S. Chile to W. Patag.—Mf.
Metzgeria Raddi (about 50). Well developed trop., some widely spread N. hemisph.
in oceanic climates.

decrescens St. S. Chile to W. Patag.—Mf.

decipiens (Massal.) Schiffn. et Gottsche. Centr. Chile to W. Patag., Austral.,
N. Zeal., Antipodes Is.—Mt, Mf.

multiformis Evans. Closely related to the former.—Mt, Mf.

violacea (Ach.) Dumort. Centr. and S. Chile, Fueg., Argent., W. and S. Afr. N.
Zeal., Antip. Is. Brought to W/. deczpzens as a variety by HODGSON (736. 278).—Mf.
Hymenophytum Dumort. (5). S. Amer., Ind., Austral., Tasm., N. Zeal., Melanes.

flabellatum (Labill.) Dumort. Colomb., Chile, Austral., Tasm., N. Zeal., N. Caled.,
Fiji,.—_<Mf.
Symphyogyna Nees et Mont. (39). Trop. and austral.

circinata Nees et Mont. Centr. and S. Chile-—Mt, Mf.

Hochstetterzt Nees et Mont. Falkl.—Mt, Mf.

hymenophyllum (Hook.) Nees et Mont. Trop. Amer.?, Chile, Austral., Tasm.,
N. Zeal.—Mt, Mf. Probably identical with S. podophylla (Thunb.) Mont. et Nees
from Centr. and S. Africa (S. ARNELL, ms.).
Pallavicinia S. F. Gray (about 25). Trop.-subtrop., Colomb., Indomal.; Kerguel.,
IND Zeal:

aiphoides (Tayl.) St. N. Zeal.—Mf.
Monocloa Hook. (2, 1 trop. Amer.).

Forstert Hook. S. Chile-W. Patag., N. Zeal.—Mt, Mf.
Androcryphaea Nees (1; Noteroclada Tayl.).

confluens (Tayl.) Nees. Mex.; Colomb.—Chile, south to Fueg., Braz., Argent.,
Falkl., Kerguel.—Mf.
Fossombronia Raddi (26). Trop.-subtrop., extending into cooler regions.

fernandeziensis St.—Mt.

Anthocerotales.

Anthoceros \.. (about 140). Mainly trop.

Skottsbergii (St. ex p.) Evans.—Mt, Mf.
Megaceros Campb. (about 40). Mex., W. Ind., S. Amer. south to Chile, Réunion,
Indomal.—N. Zeal., Oceania.

Juegiensis St. W. Patag.—Fueg. Related to neotrop. sp.—Mt, Mf.

Jungermanniales.

Solenostoma Mitt. (30). Subcosmop., but now partly referred to 4p/ozza Dum., partly
to Plectocolea.

crassulum (Mont.) St. S. Chile.—Mf.

obtusiflorum St.—Mt.

rostratum St.—Mf.
Famesoniella Spruce (20). Mainly trop.-subtrop. (1 Engl.) but also important in the
austral zone.

238 C. SKOTTSBERG

colorata (Lehm.) Spruce. Centr. Amer., S. Amer. south to Fueg., Falkl.; Tristan
da C., S. Afr., Marion I., Kerguel., E. Austral., Tasm., N. Zeal., Auckl., Campb. and
Antipodes Is.—Mt, Mf.

maluina St. Falkl.—Mt.

oenops St. S. Chile, south to Fueg., S. Georgia.—Mt.

grandifiora (Lehm. et Gottsche) Spruce. Colomb.—Chile, south to Fueg.; Tristan
da Cao Marion ils Dasm:—Me
Anastrophyllum Nees (31). Very widely distributed.

leucocephalum (Yayl.) Spruce. Venez., Peri, Fueg.—Mf.
Acrobolous Nees (11). Essentially austral: 2 S. Amer., 1 subantarct., 1 Himal., 7
Austral., N. Zeal., N. Caled.

excisus (Mitt.) Schiffn. W. Patag., Fueg., Kerguel.—Mf.
Anastrepta (Lindb.) Schiffn. (4). 2 southern S. Amer., 1 Alaska—Eur.—E. As.-Hawaii,
1 Himal.

bifida St. S. Chile: Valdivia, Magell.—Mt, Mf.

Plagiochila Dumort. (about 1000). Very few bor. zone; trop., particularly neotrop.,
but well represented southern S. Amer.

gayana Gottsche. S. Chile.—Mt.

Jfasciata St. Chiloé.—Mt, Mf.

hyadestana Besch. et Massal. S. Chile, Fueg.—Mt, Mf.

deformifolia St. W. Patag.—Mf.

chiloénsis St. S. Chile to W. Patag.—Mf.

rectangulata St. W. Patag.—Mf.

remotidens St. S. Chile, Magell.—Mf.

fuscobrunnea St. Almost too closely related to P. rudescens Lehm. et Lindenb
(Syn. P. chilens?s St. according to HERZOG 130).—Mt, Mf.

pudetensis St. Chiloé.—Mt.

homomalla St. W. Patag.—Mf.

neestana Lindenb. S. Chile south to Magell.—Mt.

riparia St. W. Patag.—Mt, Mf.

squarrosa St. W. Patag.—Mt, Mf.

robusta St. W. and E. Patag., Fueg.—Mf.

elata Hook. f. et Tayl. Fueg.—Mf.

Notarisit Lehm. Falkl.—Mt, Mf.

Tylunanthus Mitt. (25). S. hemisph., best represented Australia; 1 W. Ind.
limbatus St. W. Patag.—Mt.
silvaticus St.—Mf.
densiretis Herz.—Mf.
bilobatus St.—Mt.
Mylia S. F. Gray (40-50). The majority neotrop. mountains (Mex., W. Ind., Centr.
and S. Amer. south to Magell.), 4 Afr., 1 S. Georgia.
repens (Mitt.) Herz. Magell., N. Zeal.—Mf.
fuscovirens (Tayl.) St. See 130.714. Centr. and S. Chile to Fueg.—Mf.
ligulata (St.) Herz. W. Patag.—Mt.

DERIVATION OF THE FLORA AND FAUNA 239

Lophocolea Dumort. (250-300). World-wide; main concentrations subantarct. 5S.
Amer. (60), S. As—Oceania (35), and Austral—N. Zeal. (70).

rotundifolia St. W. Patag., Fueg., Falkl.—Mf.

Jernandestana St. Centr. Chile (Coquimbo).—Mt, Mf.

pallidevirens (Tayl.) St. Centr. (Coquimbo) and S. Chile to W. Patag., Fueg.,
Falkl., Marion I.—Mt, Mf.

papulosa St.—Mt.

attenuata St. S. Chile to W. Patag.—Mt, Mf.

terxtilis Tayl. W. Patag.—Fueg., Falkl.—Mf.

divergenticiliata St. S. Chile, W. Patag., Fueg.—Mt, Mf.

muricata Nees. W. Ind., S. Braz., Centr. and S. Chile to W. Patag., S. Afr.,
Réunion, Java, N. Guin., E. Austral., N. Zeal.—Mf.

angulata St.—Mt.

chilensis St. Chile.—Mf.

submuricata Herz. Near muricata, but also very close to L. fragrans Tay).
(atl—mediterr., 730. 720).—Mt, Mf.
Chiloscyphus Corda (150-200). Trop—subtrop., best developed S. hemisph. (about
50 Austral—Tasm—N. Zeal., 8 subantarct. Amer-.).

integrifolius Lehm. et Lindenb. Centr. Chile to Fueg.—Mt, Mf.

lobatus St. W. Patag., Fueg.—Mf.
Saccogyna Dumort. (9-10). Braz. 2, Chile 1, Eur—Macaron. 1, Indomal—Polyn. 2,
Tasm.-N. Zeal. 3; Hawaii?

squarristipula Herz. S. Chile to W. Patag.—Mt.
Marsupidium Mitt. (about 40?). Austr.-circump., subantarct. Amer., Austral.—N.
Zeal.

puliferum St. S. Chile, E. Austral., N. Caled.—Mt, Mf.
Adelanthus Mitt. (7). Widely scattered: W. Ind., Ecuad., Chile, Eur., St. Helena.

sphalerus Hook. f. et Tayl. W. Patag—Fueg.—Mt.
Bazzania S. F. Gray (230). World-wide.

cerina (St.) Fulford. S. Chile, W. Patag., Falkl., Tristan d. C.—Mt, Mf.

peruviana (Lehm. et Lindenb.) Trev. Pert, Chile to W. Patag.—Mt.
Lepidozia Dumort. (about 200). World-wide, but few north temp., most numerous
trop.subtrop., many austr.-subantarct. (70-80).

sejuncta (Angstr.) St. Mex., W. Ind., S. Braz., Azor., W. and S. Afr., Tasm.
—Mf.

bicuspidata Massal. Centr. Chile, W. Patag., Fueg.—Mt.

pseudozoopsis Herz. S. Chile to W. Patag. (z3z. 51).—Mt.

fernandeziensis St. S. Chile (230. 725). Near L. plumulosa and Lindenbergi St.
(N. Zeal.).—Mt.

plumulosa Lehm. et Lindenb. S. Chile to W. Patag. and Fueg., Falkl., N.
Zeal., Auckl. Is., Antipodes Is.—Mt.

fragillima Herz. Related to Chilean species.—Mt.

disticha St.—Mt.

Facquemontit St. Centr. Chile-Magell., Fueg.—Mt, Mf.

240 Cc. SKOTTSBERG

Herberta S. F. Gray (15). Widely scattered.

runcinata (Tayl.) Herz. S. Chile to W. Patag.—Mf.
Lepicolea Dumort. (5). Scattered S. hemisph.

ochroleuca (Spreng.) Spruce. S. Braz., S. Chile to Fueg., Cape.—Mt, Mf.
Lepidolaena Dumort. (12). S. temp. and cold zones.

magellanica (Lam.) Schiffn. S. Chile to Fueg., E. Austral., Tasm., N. Zeal.
——Mf.
Trichocolea Dumort. (32). 1 Bor. zone, 19 trop. Amer., 7 trop. As.—Oceania, 5
Austral.-N. Zeal.

opposita St. Near 7. australis St. (N. Zeal.).—Mt.

verticillata St. S. Chile to W. and E. Patag.—Mt, Mf.

Schistochila Dumort. (83). Essentially southern: austral-subantarct. (20), Afr. (7),
Indomal.—Oceania (35), Austral-N. Zeal. (21).

berteroana (Hook.) St. S. Chile.—Mt, Mf.

Skottsbergii St. Related to S. stratosa (Mont.) St. (S. Chile to Fueg.).—
Mt, Mf.

pachyla (Tayl.) St. W. Patag—Magell.—Mf.

splachnophylla (Tayl.) St. W. Patag.—Fueg., N. Zeal.—Mf.
Balantiopsis (Nees) St. (17). 1 Braz., 1 Queensl., the remainder austr.-subantarct.

cancellata (Nees) St. S. Chile to W. Patag.—Mf.

chilensis St. 5. Chile te: W. Patag.— Mi.

purpurata Mitt. S. Chile. Determination uncertain.—Mf.

hiians Herz Met.

lancifolia St.—Mt.
Radula Dumort. (
subtrop. Amer. (6
antarct. (15).

Masidiaust. 5S. Chile to Pues:— Mf,

microloba St. S. Chile-—Mt, Mf.

Dusen sts S. Chile ‘to. W> Patagi——Mf

Mittenat St. Falkl.—Mt, Mf. 2
Madotheca Dumort. (153). Trop-subtrop. Amer. (45), trop. As.-Oceania (67),
Austral._N. Zeal. etc. (12).

chilensis Lehm. et Lindenb. var. fernandesiensis Herz. Centr. Chile (Co-
quimbo; main sp. S. Chile to W. Patag.).—Mt, Mf.

subsquarrosa Nees et Mont. S. Chile to Fueg.—Mt.

220). Mainly trop.—subtrop. Bor. zone (Eur., N. Amer., 7), trop.—
6), Afr. (37), Indomal.—Oceania (69), Austral—-N. Zeal. (29), sub-

Frullania Raddi (500-600). World-wide, but mainly trop. and southern: N. Amer.
(35).

Ecklonz Spr. (crassa Herz.). Centr. Chile to W. Patag.; W. and S. Afr.—Mt.

chilensiss St. Se (Ehileto’ W: Patag:—Mf.

lobulata Hook. f. et Wils. W. Patag., Fueg.—Mf.

magellanica (Spreng.) Web. et Nees. Centr. and S. Chile to Fueg., Tristan
da (G2 Wasme,, Campb. Mt VE.

stipatiloba St. Centr. Chile to W. Patag.—Mt.

DERIVATION OF THE FLORA AND FAUNA 241

Lopholejeunea Spruce (74). Trop. and austral; trop. Amer. (13), trop. Afr. (17),
trop. As.Oceania (38), Austral.-N. Zeal. (6).

spinosa St.—Mt, Mf.
Brachiolejeunea Spruce (65). Trop. Amer. (26), Chile (1), Afr. (10), trop. As. —Oce-
ania (22), Austral._N. Zeal. (6).

spruceana (Massal.) St. S. Chile-Magell.—Mf.
Harpalejeunea Spruce (57). Trop. Amer. (36), Chile (2), Afr. (2), As—Oceania (8),
Austral.-N. Zeal. (9).

oxyota(Mont.) St. Centr. (Coquimbo)and S. Chile to W. Patag, Tristan da C.—Mf.

setifera (St.) Herz. Magell.—Mf.
Strepsilejeunea Spruce (47). Trop. Amer., south to Chile (19), Afr. (6), As. (7),
Austral._N. Zeal. (7), subantarct. (8).

acuminata (Lehm. et Lindenb.) St. S. Chile —Mt.

squarrosula Herz.—Mf.

macroloba Herz.—Mf.

Siphonolejeunea Herz. (1).
nudicalycina Herz. Centr. Chile (Coquimbo, 237. 65).—Mt, Mf.

Lejeunea Lib. (190). Pantrop., scarce toward the south.
reticulata Herz. Related. to Chilean sp.-—Mt, Mf.

Aphanolejeunea Evans (13). Trop.-subtrop.
asperrima St. S. Chile.—Mt.
diaphana Herz. Centr. Chile (Coquimbo, S. ARNELL ms.).—Mt.

Cololejeunea Spruce (about 80). Mainly trop.
Skottsbergii Herz. Nearly related to a species from N. Zealand.—Mt, Mf.

Colura Dumort. (30). Trop. to subantarct.; Amer. (8), Magell. (2), Eur. (1), Afr.
(2), Indomal. (16).
bulbosa Herz. W. Patag. (137. 66).—Mf.

The 124 Hepaticae, of which 25 (20.2 %) are endemic, belong to 47 genera
(2.6:1); 27 are thallose, 97 foliose. None have been reported from Santa Clara.

Of the 27 ¢hallose species 21 have been recorded from Masatierra and 23
from Masafuera; 4 (15 %) are known from Masatierra only, 7 (25 %) restricted to
Masafuera, 16 common to the two islands. Of the 5 endemics 2 have been found
on Masatierra only, 3 on both islands. Endemism is stronger on Masatierra (25 %)
than on Masafuera (13 %). ,

Of the 97 foliose species 59 occur on Masatierra and 67 on Masafuera; 30
(30.9 %) are restricted to Masatierra, 38 (39.2 %) to Masafuera, 29 (29.9 %) are found
on both islands. Endemic species 20 (21.6%), of these g only on Masatierra, 5
only on Masafuera and 6 found on both islands.

Of the 59 species recorded for Masatierra 15 are endemic in the islands (25° Aca)
the corresponding figures for Masafuera are 67, 11 and 16.4 %. After the discovery
of Szphonolejeunea on the mainland of Chile there is no endemic genus in Juan
Fernandez.

Of the ¢ofal number of species, 124, 80 occur on Masatierra and 90 on Masa-

16 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

242 C. SKOTTSBERG

fuera; of these 20 (25 %) and 14 (16.5 %), respectively, belong to the endemic
element; 34 species are restricted to Masatierra (27.4 %), 44 to Masafuera (35.5 %)
and 46 (37.1%) shared by the two islands. The corresponding figures for the
endemic species are: Masatierra 11 (44%), Masafuera 5 (20%), both islands 9
(36 %), and for the non-endemics 23 (23.2 %), 39 (39.4 %) and 37 (37.4 %). Of the 99
species also found elsewhere Masatierra has 60 and Masafuera 76. This island is
richer in non-endemic species, many of which belong to higher altitudes.

Geographical elements.
I. Andine-Chilean element.—97 (78.3 %).

a. Endemic (20): Riccardia adglutinata and leptostachya, Metzgeria multifor-
mis, Solenostoma obtusiflorum and rostratum, Plagiochila fuscobrunnea, Tylunan-
thus silvaticus, bilobatus and densiretis, Lophocolea papulosa, angulata and sub-
muricata, Lepidozia fragillima and disticha, Schistochila Skottsbergii, Balantiopsis
hians and lancifolia, Strepsilejeunea squarrosula and macroloba, Lejeunea reticulata.

6. Also known from Chile and not restricted to the extreme south (73): Mar-
chantia berteroana and foliacea, Riccardia fuegiensis, variabilis and nudimitra,
Metzgeria decrescens, decipiens and violacea, Hymenophytum flabellatum, Sym-
phyogyna circinata and hymenophyllum, Monoclea Forsteri, Androcryphaea con-
fluens, Solenostoma crassulum, Jamesoniella colorata and grandiflora, Anastrophyl-
lum leucocephalum, Anastrepta bifida, Plagiochila gayana, fasciata, hyadesiana,
deformifolia, chiloénsis, rectangulata, remotidens, pudetensis, homomalla, neesiana,
riparia, squarrosa and robusta, Tylunanthus limbatus, Mylia repens, fuscovirens
and ligulata, Lophocolea fernandeziana, attenuata, divergenticiliata, chilensis and
muricata, Chiloscyphus integrifolius, Saccogyna squarristipula, Marsupidium piliferum,
Bazzania cerina and peruviana, Lepidozia bicuspidata, pseudozoopsis, fernandeziensis,
plumulosa and Jacquemontii, Herberta runcinata, Lepicolea ochroleuca, Lepidolaena
magellanica, Trichocolea verticillata, Schistochila berteroana, Balantiopsis cancellata,
chilensis and purpurata, Radula hastata, microloba and Dusenii, Madotheca chi-
lensis and subsquarrosa, Frullania Ecklonii, chilensis, magellanica and stipatiloba,
Brachiolejeunea spruceana, Harpalejeunea oxyota, Strepsilejeunea acuminata, Sipho-
nolejeunea nudicalycina, Aphanolejeunea asperrima, Colura bulbosa.

c. Cosmopolitan (4): Plagiochasma rupestre, Reboulia hemisphaerica, Lunu-
laria cruciata, Marchantia polymorpha.

II. Subantarctic-Magellanian element.—18 (14.5 %).

All non-endemic: Riccardia breviramosa, Symphyogyna Hochstetterf, Mega-
ceros fuegiensis, Jamesoniella maluina and oenops, Acrobolbus excisus, Plagiochila
elata and Notarisii, Lophocolea rotundifolia, pallidevirens and textilis, Chiloscyphus
lobatus, Adelanthus sphalerus, Schistochila pachyla and splachnophylla, Radula
Mittenii, Frullania lobulata, Harpalejeunea setifera.

Ill. Neotropical element.—5 (4.0 %).
a. Endemic (4): Fossombronia fernandeziensis, Anthoceros Skottsbergii, Lo-
pholejeunea spinosa, Aphanolejeunea diaphana.
6. Not endemic: Lepidozia sejuncta.

DERIVATION OF THE FLORA AND FAUNA 243

IV. West Pacific element.— 3 (2.4 %).
a. Endemic (2): Trichocolea opposita, Cololejeunea Skottsbergii.
6. Not endemic: Pallavicinia xiphoides.
V. Known from St. Paul and New Amsterdam Is.—1r (0.8 %).

Riccardia insularis.

2 0/

The American element is in absolute dominance; 120 species, 96.8 %, belong
to elements I-III. The Pacific group is very small, but as we shall find later, many
species are austral and bicentric.

Table IV.
Comparison between the Angiosperms, Pteridophytes and Bryophytes. Figures in %.

A, Andine-Chilean; M, Magellanian; N, Neotropical; P, West Pacific.

A M N 1B
Leaks: 575 iG oo Go 5 ZACKS) 10.2 12.9 WET
Ptenidapliytesmm-ispcm cn eens) BOARD 7-5 17.0 9.4
IMIGSSCSmee wean ory a fan a eeeen, or) OAR 10.8 Taliny/ 10.8
IG DARIGS. bepis we vagts, Sana eee EONS 14.5 4.0 Dr

The lower percentage of Mosses in A, as compared with the Hepatics, may
be due to imperfect knowledge of the distribution and to the fact that 11 moss
species had to be left out of consideration.

V. Lichenes.

What I have said above when dealing with the Bryophytes holds good of
the Lichens in a still higher degree: no lichenologist ever visited Juan Fernandez,
and a non-specialist is bound to pass over many species; collecting crustaceous
lichens growing on hard basalt is not easy and generally time-absorbing. Our
collection is listed after ZAHLBRUCKNER (296, 297), where also the species found
by other collectors, but not found by us, are included. Some changes had to be
made; for instance, ZAHLBRUCKNER did not distinguish between S7zcta and Pseu-
docyphellaria; on my request, Dr. R. SANTESSON kindly revised the nomenclature.
Some determinations in these and other genera were corrected by him (see vol.
II. 886). Later he went over the proof-sheets of this paper and added further
corrections. | thank him for generous assistance.

It is to be regretted that ZAHLBRUCKNER did not indicate the distribution
of the non-endemic species, and the statements in his Catalogus are too general.
I have tried to collect further data from a number of papers on the flora of Chile
and the Subantarctic and Antarctic regions (z, 72, 73, 139, 176, 177, 182, 154, 295,
327-329) but it was beyond my possibilities to search the entire literature. Dr.
A. H. MaGnusson, who kindly took the trouble to go over my list, supplied much
useful information.

SC = Santa. Clara:

244 C. SKOTTSBERG

Verrucariaceae.
Verrucaria Schrad. (about 270). All parts of the world.
microspora Nyl. N. Amer., Chile, Eur., Jap.—Mt.
Microglaena Koerb. (37). Eur., except 1 Braz., 1 Socotra, and the following.
fernandeziana Zbr.—Mt, SC.

Dermatocarpaceae.

Normandina Nyl. (1).

pulchella Ny\. N. and S. Amer., south to Fueg., Eur., Afr., St. Paul’s 1, As,
Hawaii. Zeal.— vite

Pyrenulaceae.

Arthopyrenta Mass. (150). Trop.—subtrop.

Cinchonae M. Arg. Widely distributed in the trop. zone; also Hawaii.—Mt.

adnexa M. Arg. var. leptosperma Zbr. The typ. sp. Braz.—Mf.

planorbis M. Arg. Trop.-subtrop., also Hawaii.—Mt.
Porina M. Arg. (235). All over the globe, south to N. Caled. and N. Zeal.

fernandeziana Zbr. Belongs in the vicinity of P. chlorotica (Ach.) M. Arg.,
a cosmop. sp.—Mt.

rufocarpella Zbr.—Mt.

depressula Zbr. Possibly related to P. exserta M. Arg. (Braz.).—Mt, Mf.
Pyrenula Ach. (170). Widely distributed in the trop. zone.

aspistea Ach. Trop., also Hawaii.—Mt.

mammillana Trev. Trop.; Hawaii, 5. Chile.—Mt.
Kunthii Fée. Trop.—Mt, Mf.

Astrotheliaceae.
Pyrenastrum Eschw. (18). Trop.
chilense Mont. Chile.—Mt.

Sphaerophoraceae.
Sphaerophorus Pers. (8). Of wide distribution, centering 5S. hemisph.
melanocarpus (Sw.) DC. Almost cosmop.; Chile, south to Fueg., Falkl.; N. Zeal.,
Auckl. and Campb. Is.—Mt, Mf.

Arthoniaceae.
Arthonia Ach. (about 370). The majority trop.-subtrop.
Cytisi Massal. var. meridionalis Zbr. The typ. sp. Eur.—Mt.
subnebulosa Zbr. Related to A. ephelodes Ny). (N. Caled.) and scztuda Krmph.
(Braz.).—Mt.
berberina Zbr. Related to A. varia (Ach.) Nyl. (trop. Amer.).—Mt.
complanata Fée. Pantrop., also Hawaii; Chile.—Mt.

Graphidaceae.
Graphis Adans. (about 280). The majority trop.subtrop.
intricata Fée. Widely distributed trop. zone.—Mt.
Dumasti Spreng. Trop.—Mt.

DERIVATION OF THE FLORA AND FAUNA 245

Phaeographina M. Arg. (80). Trop.-subtrop.
scalpturata M. Arg. Trop.-subtrop. S. Amer., N. Zeal.—Mt.

Chiodectonaceae.

Enterostigma M. Arg. (2). 1 trop. Amer., and the following.
Skottsbergii Zbr.—Mf.

Dirinaceae.

Dirina Fr. (12). Of wide distribution.
limitata Nyl. Chile.—Mt.

Lecanactidaceae.

Schismatomma Fw. et Koerb. (80). Mainly warmer countries.
accedens (Nyl.) Zbr. Chile.—Mt.

Chrysotrichaceae.

Byssocaulon Mont. (5). Austral—Oceania.
niveum Mont. Subtrop., north to Japan, south to Chile and N. Zeal.—Mt.

Thelotremaceae.
Ocellularia Spreng. (over 100). Mainly warmer regions.
subdenticulata Zbr.—Mf.

Thelotrema (Ach.) M. Arg. (over 100). Mainly warmer regions.
lepadinum Ach. Widely distributed; also Hawaii, Chile, south to Fueg.,
N. Zeal., Auckl. Is.—Mt, Mf.

Diploschistaceae.

Diploschistes Norm. (30). Cold-temp., trop.-alp.
actinostomus (Pers.) Zbr. N. and S. temp.; Hawaii.—Mt.
scruposus (Schreb.) Norm. N. and S. temp.; S. Chile, south to Fuegia.—Mt.

Gyalectaceae.
Dimerella Trev. (39). Mostly trop.
lutea (Dicks.) Trev. Widely distributed, also S. Chile, Patag.; Haw.—Mt, Mf.

Gyalecta Ach. (60-70). Mainly colder climates.
jenensis (Batsch) Zbr. N. Amer., Eur.; N. Zeal.—Mt.

Pachyphiale Loennr. (4). 2 S.W. Eur., 1 bor.-temp., and the following.
cornea (With.) Poetsch et Schiederm. Eur.; Chile.—Mt.

Coenogoniaceae.
Coenogonium Ehrenb. (30). Centr. and trop. S. Amer., Afr., trop. As., Austral.
velutinum Zbr. S. Chile (comm. by R. SANTESSON).—Mt.

Racodium Fr. (1).
rupestre Pers. N. and S. Amer. (Staten I., comm. by R. SANTESSON); Eur.—Mt.

246 C. SKOTTSBERG

Collemaceae.

Lemmopsis (Vain.) Zbr. (4). 1 temp. N. Amer. + Eur., 2 S.W. Eur., and the
following.

polychidioides Zbr.—Mt.
Physma Massal. (10). W. Ind., trop. As., Japan, Austral., N. Caled.

chilense Hue. Chile.—Mt.

Leptogium (Ach.) S. F. Gray (over 100). All over the world.

moluccanum (Pers.) Vain. Widely scattered; Hawaii, also Chile.—Mt, Mf.

tremelloides (L. fil.) S. F. Gray. Trop.temp., in Chile south to W. Patag.,
5; Georeia.—Mt.

cyanescens (Ach.) Koerb. Temp.-subtrop., e.g. Hawaii, S. Amer.—Mt, Mf.

phyllocarpum (Pers.) Mont. Trop.-subtrop., S. Amer., Chile, S. Afr., Philipp.,
Australia.—Mt.

Menziestt (Sm.) Mont. Mount. of trop. Amer., Hawaii, Chile south to Fueg.,
Falkl., S. Georgia.—Mf.

callithamnion (Tayl.) Nyl. Trop. Amer.—Mt.

Pannariaceae.

Parmeliella M. Arg. (40). Widely scattered, warm and cold climates.

nigrocincta (Mont.) M. Arg. Chile, south to Fueg., W. Afr. islands, Hawaii,
Austral., N. Zeal.—Mt, Mf.

symptychia (Tuck.) Zbr.—Mt.

pycnophora (Nyl.) R. Sant. var. subdivisa (Zbr.) R. Sant. W. Patag. (329). The
tygonsps N- Zeal:-—Me:
Pannaria Del. (about 60). Widely spread over the world.

Juegiensis Zbr. Fueg.—Mt.

hilaris Zbr.—Mt.

rubiginosa Del. Scattered; e.g. Hawaii, Chile, Falkl., St. Helena, Campb. I.—Mt.

rubiginosa var. vulcanica Zbr. Perhaps specifically distinct.—Mt.
Massalongia Koerb. (2, 1 E. As.).

carnosa (Dicks.) Koerb. Mount. of N. Amer. and Eur., Falkl., N. Zeal.—Mf.
Psoroma Nyl. (about 60). Mainly cold and temp., centering in N. Zeal. etc.

vulcanicum Zbr.—Mf.

cephalodinum Zbr.—Mt.

pholidotum (Mont.) M. Arg. Chile, south to Fueg.—Mt.

sphinctrinum (Mont.) Nyl. S. Amer., also Chile, south to Fueg.; N. Zeal.—
Mt, Mf.

dasycladum Zbr.—Mt.

angustisectum Zbr.—Mt.

Stictaceae.

Lobarta Schreb. (70). Mainly warmer countries.
crenulata (Del.) Trev. Trop. Amer., south to Chile, Hawaii, Austral.—Oceania.
—Wihe:

DERIVATION OF THE FLORA AND FAUNA 247

Pseudocyphellaria Vain. (with Sticta about 200). Humid trop. to temp. climates.

argyracea (Del.) Vain. Chile, S. Afr., Madag., Mascar., Malays., Hawaii,
N. Zeal., Polyn.—Mt, Mf.

intricata (Del.) Vain. S. Amer.; Chile south to Fueg., Falkl.; Ireland, Macaron.
Mascar., Cape, Tristan da C., Hawaii.—Mt.

Jragillima (Bab.). S. Amer., Austral., N. Zeal., Auckl. and Campb. Is.—Mt.

subvariabilis (Nyl.) Vain. Philipp., Austral., N. Zeal.—Mt.

chloroleuca (Hook. f. et Tayl.) Du Rietz. S. Chile to Fueg., N. Zeal.—Mf.

cinnamomea (Rich.) Vain. S. Chile to Fueg., Austral., Tasm., Philipp.—Mt, Mf.

berteroana (Mont.).—Mt.

hirsuta (Mont.) Malme. S. Amer.; Chile south to Fueg.—Mt.

Guillemini (Mont.). S. Chile-Fueg.— Mt.

gilva (Ach.) Malme. S. Chile to Fueg., Falkl., S. Afr., Australia —Mt.

mougeottana (Del.) Vain. Warmer countries; Hawaii, also Chile-—Mt, Mf.

aurata (Ach.) Vain. Trop.-subtrop.; N. Amer., Chile, W. Eur., St. Helena,
Hawaii, Austral.—Mt, Mf.

nitida (Tayl.) Malme. S. Chile to W. Patag., Fueg.—Mf.

endochrysea (Del.) Vain. S. Chile to Fueg., Falkl., S. Georgia; Austral., N. Zeal.,
Auckl. and Campb. Is.—Mt, Mf.

Durvillet (Del.) Vain. S. Chile to Fueg., Falkl., N. Zeal.—Mt.

flavicans (Hook. f. et Tayl.) Vain. W. Patag.—Fueg.; Philipp., Hawaii, Australia.
—Mt.

Freycinetit (Del.) Malme. S. Chile to Fueg., Falkl., S. Georgia, Austral., Tasm.,
N. Zeal., Campb. and Antipodes Is.—Mt, Mf.

Richardi (Mont.) Raes. W. Patag.Fueg., N. Zeal., Auckl. Is.—Mt.
Sticta Schreb.

Weigel (Ach.) Vain. Widespread, also Hawaii, S. Chile to Fueg.; Austral.,
N. Zeal.—Mt.

lineariloba (Mont.) Nyl. S. Amer., south to Magell.—Mt.

latifrons A. Rich. Chile, N. Zeal.—Mt.

laciniata (Huds.) Zbr. S. Amer.—Mt.
Nephroma Ach. (27). N. and S. temp.

plumbeum Mont. Chile.—Mt.

cellulosum (Sm.) Ach. S. Chile to Fueg., Tasm., N. Zeal.—Mt.

antarcticum (Wulf.) Nyl. S. Chile to Fueg., Tasm.—Mt, Mf.

australe A. Rich. S. Chile, N. Zeal.—Mt.
Peltigera Pers. (20). World-wide.

rufescens (Neck.) Humb. Cosmop.; Patag., Fuegia, Falkl., S. Georgia.—Mt.

polydactyla (Neck.) Hoffm. Cosmop.; Chile, south to Fueg., Falkl., Tristan
da C., Marion I., Kerguel., Auckl. and Campb. Is.—Mt, Mf.

Lecideaceae.

Lecidea (Ach.) Th. Fr. (1000 or more?). Preferably cold or temp. climates.
avium Zbr. Related to L. aeruginosa Nyl. (Chile).—Mt.

248 C. SKOTTSBERG

inactiva Zbr.—Mt.

cyanosarca Zbr.—Mt, Mf.

leucoplaca M. Arg. Chile.—Mt.

leucozonata Zbr.—Mt.

enteroleuca Nyl. Temp. Eur., etc. St. Helena, etc.—Mt, Mf.

latypea Ach. Temp. widely distributed.—Mt.

viridans Lamy. Eur.—Mt.

mutabtlis Fée. N. and S. Amer., also Chile, W. Eur.—Mt.

icterica Tayl. S. Amer., also Chile.—Mf.
Catillaria (Ach.) Th. Fr. (about 150). Widespread N. and S. hemisph.

intermixta Arn. Widespread, south to N. Zeal.—Mt.

melastegia (Nyl.) Zbr. S. Amer., Chile south to Fueg., N. Zeal.—Mt, Mf.

endochroma (Fée) Zbr. N. and S. Amer.—Mt.

leucochlora (Mont.) Zbr. Chile.—Mt, Mf.

theobromina Zbr.—Mt.
Megalospora Mey. et Flot. (about 50). Warmer regions.

versicolor (Fée) Zbr. var. microcarpa Zbr. The typ. sp. S. Amer., N. Zeal.—Mt.
Bacidia (De Not.) Zbr. (at least 200). World-wide.

endoleuca Kickx. Almost cosmop., also S. Amer., N. Zeal.—Mt, Mf.

arceutina (Ach.) Arn. var. hyposcotina Zbr. The typ. sp. Eur.—Mt.

delapsans Zbr. Hawaii.—Mt.

subluteola (Nyl.) A. Zbr. Braz.—Mf.
Toninia Th. Fr. (about 80). Mainly temp.

bullata (Mey. et Flot.) Zbr. Ecuador: Chimborazo.—Mf.
Lopadium Koerb. (about 50). Almost cosmop.

leucoxanthum (Spreng.) Zbr. var. albidius Zbr. The typ. sp. subtrop., south
to Austral. and N. Zeal.; Hawaii.—Mt.

sp., different from the former, described as M/yxodictyon lopadioides by ZAHL-
BRUCKNER p. 383. Chile.—Mt.
Rhizocarpon DC. (about go). Cosmopol., cold to temp. climates.

geographicum (1...) DC. Cold and temp., both hemisph.; Patag., Fueg., Falkl.,
Kerguel., W. Antarct.—Mf.

microspermum Zbr. Similar to the following sp.—Mf.

obscuratum (Ach.) Massal. var. deminutum Zbr. The typ. sp. cold and temp.
climates.—Mf.

Phyllopsoraceae.

Phyllopsora M. Arg. (25). Trop.—subtrop.

parvifolia (Pers.) M. Arg. Widespread, also Chile, Hawaii and N. Zeal.—Mt.

Cladoniaceae.

Baeomyces Pers. (34). Majority trop.; 7 N. Zeal.
chilensts (Mont.) Cromb. Chile.—Mt, Mf.
Cladonia (Hill) Vain. (about 280). All over the world.
pycnoclada (Pers.) Nyl. Boliv., S. Chile to Fueg., Falkl.; Tristan da C.?-—Mt,

DERIVATION OF THE FLORA AND FAUNA 249

Mf. To this belongs C. a/pestris of ZAHLBRUCKNER 296. 370, a species not occur-
ring in the south hemisphere (SANTESSON 327 and in letter).

bacillaris (S. F. Gray) Nyl. Very widespread; Chile, Fueg., Falkl., Campb. I.
—Mt.

didyma Vain. Mex.—Fueg., Austral., N. Caled., Hawaii.—Mt.

coccifera (L.) Willd. Very widespread; Chile, Patag., Fueg., Falkl., Tristan
da C., W. Antarct.—Mt, Mf.

aggregata (Sw.) Ach. N. and S. Amer., Chile to Fueg., Falkl.; St. Helena,
S. Afr., Madag., Auckl. and Campb. Is., Macquarie I.; Asia; Hawaii.—Mt, Mf.

furcata (Huds.) Schrad. Cosmopol.; Chile to Fueg., Falkl., S. Georgia, W.
Antarct., Antipodes Is.; Hawaii.—Mt.

gracilis (L.) Willd. Cosmopol.; Chile to Fueg., Falkl., S. Georgia, Tristan
da C.; Hawaii; Kerguel., W. Antarct.—Mt, Mf.

pyxidata (L.) Fr. Cosmopol.; Chile to Fueg., Falkl., Tristan da C., Kerguel.,
St. Paul’s I.; Hawaii.—Mf.

jimbriata (L.) Fr. Cosmopol.; Chile to Fueg., Falkl., Marion I., Kerguel., W.
Antarct., Auckl. and Campb. Is.; Hawaii.—Mt, Mf.

pityrea (Flk.) Fr. Cosmopol.; Falkl., Tristan da C.; Hawaii.—Mt, Mf.
Stereocaulon Schreb. (about 90). N. and S. temp., trop—subtrop. mountains.

patagonicum M. Lamb. S. Chile and Patag. to Fueg., Falkl.—Mt, Mf.

ramulosum (Sw.) Raensch. N. and S. Amer., Chile south to Magell., N. Zeal.
—Mt, Mf.

wmplexum Th. Fr. S. Amer., south to Fueg.; N. Zeal.—Mt, Mf (f. compactius
(Zbr.) M. Lamb.).

Acarosporaceae.
Acarospora Massal. (over 200). World-wide.

smaragdula (Wahlenb.) Massal. var. N. temp.—Mf.
aanthophana (Nyl.) Jatta. S. Amer. mountains.—SC.

Pertusariaceae.
Coccotrema M. Arg. (1).
granulatum (Hook. f. et Tayl.) R. Sant. n. comb. (C. curbitula M. Arg., Porina
granulata Hook. f. et Tayl.). Chile: Valdiv—Fueg., Ceyl., Jap., Philipp., N. Zeal.
—— Mt:
Pertusaria DC. (about 200). World-wide.
letoplaca (Ach.) Schaer. Cosmop.—Mf.
polycarpa Krph. var. monospora Zbr. The typ. sp. Braz.—Mf.
hadrocarpa Zbr. Similar to A. cerebrinula Zbr. (Falkl.).—Mt, Mf.
Skottsbergii Zbr. Related sp. in Chile-—Mt, SC.
Melanaria Erichs. (15). N. and S. Amer., Eur., S. Afr., As., N. Zeal.
melanospora (Nyl.) Erichs. S. Amer., also Chile.—Mt, SC, Mf.

Lecanoraceae.

Lecanora Ach. (over 200). World-wide.
masafuerensis Zbr. Close to L. subimmersa (Fée) Vain. (Braz.).—Mf.

250 C. SKOTTSBERG

coarctata (Sm.) Ach. Widespread temp.; Chile.—Mf.

atra (Huds.) Ach. Cosmop.; Chile, Falkl.; Hawaii.—Mt, Mf.

Ingae Zbr.—Mt, Mf.

albellina M. Arg. var. validior Zbr. The typ. sp. Fueg.—Mf.

dispersa (Pers.) Flk. Widespread N. hemisph.—Mt, Mf.

polytropa (Ehrh.) Ach. Cosmop.; Chile, Falkl., W. Antarct.; Hawaii.—Mf.

chrysoleuca (Sm.) Ach. W. Arct., Eur. mountains, Antarct., etc.—Mt.

saxicola (Poll.) Ach. N. Amer., Eur.; ?Chile.—Mt.
Placopsis Nyl. (31). Widely distrib., but mainly austral, 18 southern S. Amer.

chilena M. Lamb. Chile.—Mf.

Juscidula M. Lamb, S. Chile to Fueg., Tristan’ da C.—Mt,’ Mf.

parellina (Nyl.) M. Lamb. Boliv. And., Chile to Fueg., Java, S. Austral.,
N. Zeal., Hawaii.—Mt.

gelida (L.) Ach. Circumpol. Arct. and temp. oceanic; Chile, Tristan da C.,
Kerguel., Java, N. Zeal., Hawaii.—Mf (var. subreagens M. Lamb, but identity
doubtful).
Candelariella M. Arg. (27). N. and S. Amer., Eur.

vitellina M. Arg. Widespread; Chile, Falkl., Hawaii.—Mt, Mf.
Myxodictyon Massal. (3, 1 Australia).

chrysostictum (Tayl.) Mass. Chile, N. Zeal.—Mt. J7. lopadioides Zbr. 296. 383
is a species of Lopfadium (SANTESSON in letter).

Parmeliaceae.

Parmelia (Ach.) De Not. (about 400). World-wide.

laevigata (Sm.) Ach. Widespread temp. and trop.; also Hawaii and Chile.
NE,

laevigatula Ny\. Braz.—Mt.

revoluta F\k. Widespread temp.-trop. Also Hawaii.—Mf.

cetrata Ach. Very widespread temp. and trop., also Hawaii.—Mt, SC, Mf.

_ saxatilis (L.) Ach. Widespread, in Chile south to Fueg., Falkl.; W. Antarct.

= Mit.

conspersa (Ehrh.) Ach. Cosmop., also Hawaii, Falkl.—Mt, SC, Mf.

abstrusa Vain. Braz.—Mf.

perlata Ach. Widespread; also Hawaii and S. Chile.—Mt.

nilgherrensis Ny\. Trop.-subtrop.—Mf.

puosella Hue. N. Amer. to Mex., Eur.—Mt, Mf.

piloselloides Zbr.—Mt.

cetrarioides Del. Very widespread.—Mf.

microstictta M. Arg. Trop. Amer.—Mt.

caperata (L.) Ach. Temp. zones; Chile, Hawaii.—Mt, Mf.

soredica Nyl. Calif., Mex.—Mf.
Menegazzia Mass. (30). Few north hemisph., majority S. Amer. (11) and Austral.-
Tasm.-N. Zeal. (14).

sanguinascens (Raes.) R. Sant. (328. 11, Parmelia pertusa (Schrad.) Schaer. in
290. 389), Valdiv. to Fueg.—Mt, Mf.

DERIVATION OF THE FLORA AND FAUNA 2

On
_

Ramalina Ach. (about 100). World-wide.
linearis (Sw.) Ach. Warmer regions; Chile, Fueg., Falkl., N. Zeal.—Mt.
usnea (L.) Howe Jr. Trop.temp., N. Amer., Chile.—Mf.
Usnea Wigg. (about 100). Cosmop.
dasypogoides Ny\. Rodriguez I.—Mt, Mf.
florida (L.) Hoffm. Very widespread, also Chile.—Mf.
subtorulosa (Zbr.) Motyka. Easter I.—Mf.
angulata Ach. N. Amer., south to Mex.; S. Amer. to Chile.—Mf.

Caloplacaceae.

Blastenia (Massal.) Th. Fr. (about 60). World-wide.

fernandeziana Zbr.—Mt, SC.

ferruginea (Huds.) Massal.—Mt. Determination probably incorrect acc. to Dr.
MAGNUSSON.
Bombyliospora De Not. (25). Mostly trop.-subtrop.

dolichospora (Nyl.) Zbr. Chile.—Mt.
Caloplaca Th. Fr. (over 100). World-wide.

clandestina Zbr.—Mf.

Selkirkii Zbr.—Mt, Mf.

rubina Zbr. Easter I1.—Mt, SC, Mf.

isidioclada Zbr.—Mf.

subcerina (Nyl.) Zbr. var. aurantiaca Zbr. Trop.—Mf.

elegans (Link) Th. Fr. var. australis Zbr. Chile, W. Patag. The typ. sp. cold
tomremp. N: and S: hemisph.—Mt, SC, Mf.

orthoclada Zbr. In the vicinity of C. Felipone? Zbr. (Urug.).—Mf.

Teloschistaceae.

Teloschistes Norm. (12). World-wide.

flavicans (Sw.) M. Arg. Widespread trop.-subtrop., in 5. Amer. south to
muess Falkl-—Mt-.

Buelliaceae.

Buellia De Not. (about 200). World-wide.

concinna (Stzbgr.) Th. Fr. var. oceanica Zbr. The typ. sp. Eur.—SC.

siphoniatula Zbr. Similar to B. posthabita (Nyl.) Zbr. (Colomb.) and fa/k-
landica Darb. (Falkl.).—Mt.

stellulata (Tayl.) Mudd. Cosmop. Chile; Kerguel.—Mt, SC, Mf.

halophila M. Arg. Australia.—Mt.

halophiloides Zbr. Easter 1.—Mt.

Jernandeziana Zbr. Easter I.—Mt.

masafuerana Zbr.—Mf.

barrilensis Zbr.—Mf.

Physciaceae.
Pyxine (Fr.) Nyl. (16). Warmer regions.
curvatula Zbr.—Mt.

252 C. SKOTTSBERG

Physcia (Schreb.) Vain. (100-150). Cosmop., most numerous temp. regions.
picta (Sw.) Nyl. Widespread trop.-subtrop. Also Hawaii.—Mt.
Anaptychia Koerb. (30). Widespread, mainly warmer regions.
hypoleuca (Muhlb.) Massal. Widespread, also Hawaii.—Mt.
pectinata (Zbr.) R. Sant. Patag. (Nahuelhuapi), Fueg.—Mt.

Hymenolichenes.

Cora Fr. (8). Trop.—subtrop.
pavonta (Sw.) Fr. Mex.—Chile, south to Fueg.; St. Helena.—Mt, Mf.

The list includes 194 species; 103 (53 %) of these are restricted to Masatierra,
including 3 also found on Santa Clara, 2 (1 %) have only been encountered on
this islet, 39 (20 %) only on Masafuera; 50 (25.7 %) are listed for both islands,
6 of them also found on Santa Clara. The number of endemic species is 36 (18.5 %),
a number likely to be reduced when the lichen flora of South America becomes
better known; some species described by ZAHLBRUCKNER as endemic in Juan
Fernandez have later been found on the mainland or on Hawaii. New discoveries
will, on the other hand, be made in the islands.

Masatierra has 153 species, of which 26 (17%) are endemic in Juan Fer-
nandez; the corresponding figures for Santa Clara are 11 and 3 (27.3 %) and for
Masafuera 89 and 15 (16.8%). Masatierra is richer in lichen species than Masa-
fuera; this may have something to do with the greater variety of substratum of-
fered by the numerous species of trees and shrubs inhabiting only Masatierra.
Of 100 corticolous species 62 are restricted to Masatierra, 16 to Masafuera and
22 occur on both islands; thus 84 species have been collected on the former and
only 38 on the latter.

The greater wealth of Masatierra is also shown in the number of endemic spe-
cies found only on Masatierra, 21 (of 36) or 58.3%; 3 were found also on Santa
Clara. The figure for Masafuera is 10 = 27.7%; of these 5 belong to the highland
above 1000 m. Only 5 species (14%) have been found on both islands. I suppose
that the superiority of Masatierra depends on the greater variation of habitat.
Future research will, I suppose, yield numerous additional species, but I do not
expect that the relation between the islands will be much altered.

It is difficult to arrive at a geographical classification of the lichens because
in too many cases only very general information is given: “‘in the tropics’, “in
warmer regions’, ‘in temperate regions” and so forth, and “‘cosmopolitan” is used
too generously. Statements suggesting the most surprising disjunctions are not
uncommon; some are probably due to wrongly named specimens. Unfortunately
the lichen flora of Chile is not very well known. I have tried to find out if a
species called cosmopolitan has been recorded for Chile. Many world-wide lichens
have been found in Juan Fernandez; if also found in Chile they were referred
to the Chilean element.

DERIVATION OF THE FLORA AND FAUNA 2

n
eS)

I. Andine-Chilean element.—106 (62.4%).

a. Endemic species supposed to be most nearly related to Chilean species (7):
Psoroma vulcanicum, cephalodinum, dasycladum and angustisectum, Pseudocyphel-
laria berteroana, Lecidea avium, Pertusaria Skottsbergii.

6. Also known from Chile (88): Verrucaria microspora, Normandina pulchella,
Pyrenula mammillana, Pyrenastrum chilense, Sphaerophorus melanocarpus, Arthonia
complanata, Dirina limitata, Schismatomma accedens, Byssocaulon niveum, Thelo-
trema lepadinum, Diploschistes scruposus, Dimerella lutea, Pachyphiale cornea,
Coenogonium velutinum, Racodium rupestre, Physma chilense, Leptogium molucca-
num, tremelloides, cyanescens, phyllocarpum and Menziesii, Parmeliella nigrocincta
and pycnophora var., Pannaria rubiginosa, Psoroma pholidotum and sphinctrinum,
Lobaria crenulata, Pseudocyphellaria argyracea, intricata, cinnamomea, hirsuta,
Guilleminii, gilva, mougeotiana and aurata, Sticta Weigelii, lineariloba and lati-
frons, Nephroma plumbeum and australe, Peltigera rufescens and polydactyla,
Lecidea leucoplaca, mutabilis and icterica, Catillaria melastegia and leucochlora,
Rhizocarpon geographicum, Phyllopsora parvifolia, Baeomyces chilensis, Cladonia
pycnoclada, bacillaris, didyma, coccifera, aggregata, furcata, gracilis, pyxidata,
fimbriata and pityrea, Stereocaulon ramulosum and implexum, Coccotrema granu-
latum, Melanaria melanospora, Lecanora coarctata, atra and polytropa, Placop-
sis chilena, fuscidula, parellina and gelida, Candelariella vitellina, Myxodictyon
chrysostictum, Parmelia saxatilis, laevigata, conspersa, perlata and caperata, Mene-
gazzia sanguinascens, Ramalina linearis and usnea, Usnea florida and angulata,
Bombyliospora dolichospora, Caloplaca elegans, Buellia stellulata, Theloschistes
flavicans, Cora pavonia.

c. Widespread to cosmopolitan species expected to occur on the mainland
of Chile (11): Diploschistes actinostomus, Pseudocyphellaria fragillima, Sticta laci-
niata, Catillaria intermixta and endochroma, Bacidia endoleuca, Pertusaria leio-
placa, Parmelia revoluta, cetrata and cetrarioides, Physcia picta.

II. Subantarctic-Magellanian element.—1 4 (8.2 %).

a. Endemic: Pertusaria hadrocarpa.

6. Also in W. Patagonia, Fuegia etc. (13): Pannaria fuegiensis, Pseudocyphel-
laria chloroleuca, nitida, endochrysea, Durvillei, flavicans, Freycinetii and Richardi,
Nephroma cellulosum and antarcticum, Stereocaulon patagonicum, Lecanora albel-
lina (endem. var.), Anaptychia pectinata.

III. Neotropical element.—26 (15.3 %).

Endemic, or found or expected in South America, but not recorded for Chile.

a. Endemic (5): Arthonia subnebulosa and berberina, Enterostigma Skotts-
bergii, Lecanora masafuerensis, Caloplaca orthoclada.

6. Not endemic, some perhaps to be expected in Chile (14): Arthopyrenia
Cinchonae, adnexa and planorbis, Phaeographina scalpturata, Leptogium calli-
thamnion, Megalospora versicolor (endem. var.), Bacidia subluteola, Toninia bullata,
Acarospora xanthophana, Pertusaria polycarpa (endem. var.), Parmelia laevigatula,
abstrusa and microsticta, Anaptychia hypoleuca.

254 C. SKOTTSBERG

c. Widespread tropical-subtropical species, probably also occurring in South
America (7): Pyrenula aspistea and Kunthii, Graphis intricata and Dumastii, Lo-
padium leucoxanthum (endem. var.), Parmelia nilgherrensis, Caloplaca subcerina

(endem. var.).
IV. Pacific element.—7 (4.1 %).

Pseudocyphellaria subvariabilis, Bacidia delapsans, Usnea subtorulosa, Calo-
placa rubina, Buellia halophila, fernandeziana and _ halophiloides.

V. Boreal element.—16 (9.4 %).

a. Endemic: Lemmopsis polychidioides.

6. Not endemic (15): Arthonia cytisi (endem. var.), Gyalecta jenensis, Massa-
longia carnosa, Lecidea enteroleuca, latypea and viridans, Bacidia arceutina
(endem. var.), Rhizocarpon obscuratum (endem. var.), Acarospora smaragdula,
Lecanora dispersa, chrysoleuca and saxicola, Parmelia pilosella and soredica, Buel-

lia concinna (endem. var.).

VI. Only reported from Juan Fernandez and Rodriguez I.

Usnea dasypogoides.

The following 22 endemic species had to be left out, their taxonomic rela-
tions being unknown: Microglaena fernandeziana, Porina depressula, fernandeziana
and rufocarpella, Ocellularia subdenticulata, Parmeliella symptychia, Pannaria hilaris,
Lecidea inactiva, cyanosarca and leucozonata, Catillaria theobromina, Rhizocarpon
microspermum, Lecanora Ingae, Parmelia piloselloides, Blastenia fernandeziana,
Caloplaca clandestina, Selkirkii and isidioclada, Buellia siphoniatula, masafuerana
and barrilensis, Pyxine curvatula—further, Lopadium sp, and the dubious Blastenia

sp. have been excluded. The percentages were calculated with 170 as a total.

VI. Fungi.

Our knowledge of the fungus flora is very limited, and time did not permit
us to pay due attention to this group. Of Baszdiomycetes about 40 species were
identified by ROMELL (207), including those enumerated by JONOW (several spe-
cies doubtful). Endemic species few. KEISSLER’s list of Ascomycetes (159, 160),
with additions by ARWIDSSON (377), short as it is—only 31 species—gives some
idea of the relation between the geographical elements. Of the 20 named spe-
cies, 9 were known before from S. America (mostly Chile) and 1 from N. Amer-
ica; 5 are widespread and 5 endemic, one of these belonging to an endemic
genus. Limacinia scoriadea (Berk.) Keissl. is also known from Chile, Java and New
Zealand. Three of the endemic genera of Compositae have their special rusts,
the endemic Luphrasza is attacked by the same Uvedo that is found on two species
of sect. 77zfidae in Chile, Azara fernandeztana by the rust known from A. ixtegrifolia
in Chile, and Rudus geoides is accompanied by the same parasite as in Fuegia (z59).

Of Gasteromycetes only 2 named species were reported; of these //eodictyon
gracile Berk. is of geographical interest: S. America, S. Africa, Australia and New
Zealand, the second species was known before from N. America and Samoa (z02).

DERIVATION OF THE FLORA AND FAUNA

nN
UW
wn

The Myxophyta, 18 species, are more or less cosmopolitan (zo7).

I have tried above to indicate where the native plants have their nearest sta-
tions outside the islands or, if endemic, where their closest relatives occur. Sta-
tistics like this serve to assign to a local flora its position within a certain floristic
region and, when dealing with an oceanic island, to trace the sources from where
its living world is likely to have been derived. The position generally assigned
to Juan Fernandez is that of an outpost from South America. In ENGLER's Syl-
labus the islands form the “Gebiet von Juan Fernandez und Masafuera’’ under
“Zentral- und siidamerikanisches Florenreich’’; it is characterized thus: “Gattungen
vorzugsweise verwandt mit denen der chilenischen Ubergangsprovinz”, i.e. Cen-
tral Chile and the Valdivian forest region, ENGLER’s transitional belt between the

Andean and Magellanian provinces. GOOD (zog) distinguished a ‘“‘Region of Juan
Fernandez’ under his “Neotropical Kingdom’, giving it the same rank as the
“Amazone Region’, the ““Andean Region’, etc. We have seen that there is an
unmistakable floristic agreement between Juan Fernandez and South Chile, but
also that it is far from complete, and both ENGLER and GOOD were well aware
of the presence of elements that had little to do with the flora of the mainland.
They were barely recognized by JOHOW (see above p. 215) who, with his faith
in unlimited transoceanic dispersal, paid little attention to them; they were too
few to disturb the Chilean picture. Not until the flora had become better known
did its strange features stand out in a clearer light. It is surprising that GUILLAU-
MIN who knew and quoted the synopsis published in vol. II of this work (229)
failed to recognize them. In his paper on the floristic divisions of the Pacific he
states that Juan Fernandez lies outside Oceania and he includes it in his discus-
sion only for the sake of comparison. The flora is characterized as follows (778.
931): “Sur 142 Phanérogames indigénes, la moitié sont endémiques mais appa-
rentees aux espéces chiliennes; les autres sont cosmopolites ou Sudaméricaines,
surtout chiliennes.” It is not easy to understand how he arrived at this conclusion.
On the other hand, some authors were led astray by the dzfference between the
islands and the continent. ERMEL, who got his impression from a short visit to
Masatierra, wrote (gz. 48):

.. muss die Zusammengehdrigkeit dieser Insel zum amerikanischen Festlande, zu
welcher Ansicht deren geringe geographische Entfernung von selbst hinleiten kénnte, aufs
nachdriicklichste in Abrede gestellt werden, weil die beiderseitige Flora and Fauna zu
grosse Verschiedenheiten aufweist, wo wir an geeigneter Stelle die notigen Beweise bei-
bringen werden. Die Gestaltung der Flora ordnet die Inselgruppe vielmehr dem australischen
Weltteile zu, von dem der grésste Teil heruntergegangen ist.

It is hardly necessary to mention that his proofs were based on his ignorance
of the composition of the island flora and very likely also of the floras of Australia
and Chile.

256 C. SKOTTSBERG

Chapter II.

Sources of the island flora as judged by the total distribution of the
geographical elements distinguished, with special reference to the
composition of the Chilean flora.

I have attempted above to describe the distribution of the non-endemic spe-
cies, to state where the endemics have their relatives, if any, and to distinguish
a number of geographical elements. Now I shall proceed a little further and look
at the matter from a wider horizon. A species was called Chilean because it is
found also in Chile or has its relatives in the south Andean flora, or it was referred
to a Magellanian group because it occurs only in the farthest south of the conti-
nent, and so forth; but in order to know something of the genesis and history
of each group we cannot stop here. We shall find that our “Chilean element’,
Chilean from our insular viewpoint, consists of several types, each with its own
distribution pattern. To speak with WULFF (297. 203), until now we occupied our-
selves with the geographic elements, now we shall try to trace the gezefzc ones,
“species grouped according to their region of origin, thus reflecting the genesis
of a given flora”. He very properly adds: ‘To determine the region of origin of
a species’’—and indeed also of a genus or family—“‘is often a very difficult mat-
ter, requiring a monographic study... .’ With regard to Juan Fernandez, the ge-
netic elements are congruent to WULFF’s “migration elements”.

I. Angiospermae. ;

Of the two species of S#pa, xeeszana is distributed from Mexico through the
tropical Andes to Central Chile and east to Brazil, Argentina and Uruguay, /ae-
vissima (Nassella) a typical Andean species; the former is neotropic, the latter
Chilean, extending north into Pert and east into Argentina. Almost the same
area is occupied by Piptochaetium bicolor, a genus limited to extratropical South
America.

Podophorus is a unique anomaly without known neotropical affinities, as it
were, a far-travelled member of an Arcto-tertiary flora, if its affinity with Lrachy-
elythrum hits the mark; in PILGER’s opinion it sides with A/egalachne.

The species of Chaefotropis were referred to the neotropical group— tropical”
not to be taken in a purely climatic sense, because a species included under this
heading may just as well be subtropical and even extend into a temperate zone.

Agrostis masafuerana and the bicentric A. magellanica were linked together,
but only provisionally, because in a large and world-wide genus like Agvosézs the
relationships cannot be safely judged without a thorough taxonomic-genetic study
of the whole genus. Assuming that PILGER was right, Antarctica becomes in-
volved, and the two species—and probably others as well—should be classified as
“Old Antarctic’, or, as I now prefer to term them, Antarcto-tertiary (correspond-

DERIVATION OF THE FLORA AND FAUNA 257

ing to Arcto-tertiary). An Antarcto-tertiary taxon is not necessarily of Antarctic
ancestry; strictly spoken it certainly should be, but there are numerous cases
where the point of origin was either the South American or the Australasian—
New Zealand centre, and Antarctica only served as a trans-continental route of
migration.

Trisetum chromostachyum is Chilean, but the genus is both N. and S. temper-
ate, perhaps originally Arcto-tertiary? Danthonia collina and Koeleria micrathera
also are Chilean, but the distribution pattern of the genera indicates that Antarc-
tica eventually was involved in their history.

Megalachne, temporarily referred to Bromus but once more stated to be an
independent genus, is a relict type with unknown history, eventually a remnant
of a pre-Andean flora (comp. above p. 217).

The genus Chusquea belongs to the neotropical element in the flora of Chile,
and the same is true of the species of Cyperus. Scirpus nodosus and cernuus are
circumpolar seaside plants and most likely thalassochorous; if Antarctica had a
share in their earlier history is impossible to say, but it is not improbable. //eo-
charis is another large world-wide genus, perhaps too wide-spread to allow us to
locate its place of origin.

Oreobolus. The Antarcto-tertiary character of Oveobolus can hardly be dis-
puted; it is often referred to as a classical example of an Antarctic genus. Of the
6 species recognized by KUKENTHAL, I (with 2 varieties) is found in S. and E.
Australia, Tasmania and New Zealand, 2 in N.E. New Guinea, one of them
extending to Borneo, 1 in N. Sumatra and Malacca, 1 in Hawaii, and O. ob/usangu
fus in Chile from the cordilleras of Valdivia to Fuegia, the Falkland Is. and Juan
Fernandez. [A seventh species, O. pfezfferianus Barros, was identified by KUKEN-
THAL (764. VIII) with pumzlio var. fectinatus.|

Cladium (164. Xi1) is wide-ranging but it is not cosmopolitan in spite of the
large areas occupied by C. marzscus L. and its varieties, among which jamazcense
is circumpolar and distributed also south of the equator. The main distribution
of subg. Machaerina (incl. Vincentia), where C. scérpotdeum of Juan Fernandez
belongs, is palaeotropical with 11 species (Madagascar, Mascarene Is., Australia,
Lord Howe I., New Guinea, Indonesia, Oceania); 5 are neotropical (W. Indies,
Brazil). Its austral character is clear enough. The closest relative of scirpozdeum
is not an American species but C. anxgustifolium (Gaud.) Benth. et Hook. fil. (New
Guinea, Tahiti, Hawaii). Subg. Bauwmea, with 29 scattered from Australia, where
there are 18 species, over the Indomalayan region west to Ceylon, north to Hong-
kong and Japan, east to Melanesia and Hawaii, has 15 in Australia—lasmania—
New Zealand and 1 in the region of Madagascar and Mascarene Is. Together the
two subgenera cover the 3 sectors, the African, the Australian-Malaysian and the
American. In the centre of this vast area lies Antarctica or, as it were, Gond-
wana Land. The history of Cladim may well lead back to the Mesozoic, and
it seems natural to refer the genus to the Antarcto-tertiary element. The same
applies to Unxcznia, one of the generally recognized Antarctic genera, represented
in Juan Fernandez by 5 species, 2 of them endemic. Within the South American
sector are 13 species distributed along the Andes with a concentration toward the

17 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

258 C. SKOTTSBERG

south, one species going north as far as Mexico and the West Indies, 1 extending
east from Fuegia to the Falkland Is. and Tristan da Cunha. On the opposite side
of Antarctica are 18 species, 14 of these indigenous in New Zealand, where 8 are
endemic, the remainder scattered along the route Macquarie I—Auckland and Camp-
bell Is.-Tasmania—Australia-New Zealand—Lord Howe and Norfolk Is.; one New
Zealand species reappears in Hawaii, another on Kerguelen and New Amster-
dam I., and Marion I. has an endemic species. Finally, 2 species are found in New
Guinea, one of them also reported from Borneo, and the Philippines have 1 en-
demic species. Both sections of subg. Lu-Uncinia are represented in America, only
Stenandrae in the opposite sector. The monotypical subg. Pseudocarex is Magel-
lanian.

It is not easy to find one’s way through the labyrinth of the enormous and
still. growing genus Carex, world-wide but unbalanced as the tropics are poor in
species in comparison with the temperate and cold zones. Our two island species,
the endemic C. derteroniana and the south Andean Baxksz7, belong to different
sections and different geographic groups, the former to sect. Echinochlaenae: of
20 species 16 are endemic in New Zealand (one with a variety on Norfolk I.),
1 in Australia, 1 in Tasmania; the little known Chilean C. /amprocarpa Phil. and
C. berteroniana are far-flung outposts, but it lies near at hand to assume that they
or their ancestors migrated across Antarctica. C. Banksia belongs to Frigidae-
Fuliginosae, a boreal group centering in Eurasia with one species in Pacific North
America, but I cannot tell if Baxkszz comes near this species.

The systematic position of F¥wanza australis was briefly discussed in 229.
109 and above p. 202, but whether we bring it to Morenieae or Iriarteae or let it
form a separate subtribe it remains a member of the neotropical element. HUTCHIN-
SON followed BENTHAM and HOOKER in placing it next to Ceroxy/on, but this
genus is polygamo-monoecious and the stigma becomes basal in fruit. To me
CROIZAT’s opinion lacks foundation; he solves what he calls “‘a hopeless conflict
among taxonomists” (77.85) by deriving ¥wanza from ‘‘a massive center of origin
of angiospermy at the Mascarenes”’ (p. 103).

Ochagavia and Hesperogrezgia belong to an Andean-neotropical assemblage
of genera, with close relatives in Chile.

To judge from BUCHENAU’s monograph of the Juncaceae in Pflanzenreich
Luzula masafuerana must be referred to a group of Andean species, L. racemosa
Desv. (Mex.-S. Chile), exce/sa Buch. (Boliv.), Hzeronym7 Buch. (Argent.), Leydoldiz
Buch. (Chile), and chz/ens7s Nees et Mey. (Chile, south to Fuegia), but in the same
group we find ZL. sfzcata (L.) DC. (Arct.-circump. and Alpine) and adyssznica Parl.
(Ethiop., Brit. E. Afr.), and the possibility that the Andine species are of boreal
origin should be considered. On the other hand much speaks in favour of a south-
ern origin of Juncaceae: the subantarctic genera Marsippospermum and Rostkovia,
the isolated Przoxium in South Africa, endemic Andine genera like Oxychloe and
Patosia, well-marked endemic species of Luzu/a in New Zealand and the Magel-
lanian region, and the subantarctic-bicentric Funcus scheuchzerioides group. It is
true that 4 of the 5 Yuzcus species reported from Juan Fernandez—some of them
perhaps not native—inhabit the Andes, 3 going east to Brazil, Argentina and Uru-

DERIVATION OF THE FLORA AND FAUNA 259

guay, the fifth, ¥. planzfolmus, shared by Chile and Australia~Tasmania—New Zea-
land, but it is also true that even if we look for the origin of the family in
Antarctica, a possibility pointed out by WEIMARCK (287), we must count with
secondary centres of evolution in the boreal zone where the overwhelming major-
ity of sections and species are found. Some 50 species are in the South hemi-
sphere with main centres in South Africa (20) and Australia (17).

Libertia with 3 species in Chile and 2 in New Guinea—Australia-New Zealand
telis a story of an Antarctic past, and the group Sisyrinchineae has a stronghold
in the South, where the genera concentrate; Szsyrizchinm itself centres in South
America, where 4 small genera are endemic (Symphyostemon, Chamaelum, Sole-
nomelus and Tapetnia); South Africa is another stronghold (Avzstea, Witsenia,
Bobartia, Klattia and Cleanthe), and 3 (Orthosanthus, Diplarrhena and Patersonia)
are found in the East Australian -Indomalayan region, but this does not entitle us
to derive Iridaceae from the far south.

Of the 4 island species of Peperomia only P. fernandeziana (Chile) is neo-
tropical, while the other species appear to be more nearly allied to palaeotropical
ones (Java, Australia, Oceania); P. derteroana occupies a unique geographical posi-
tion (p. 203). The genus is of tropical origin and centres in America, but if we
try to understand the history of the endemic species of Juan Fernandez, the possi-
bility of Antarctica as a migration route should be considered.

Whereas Urtica masafuerana, one of the few indigenous annuals, is related
to a species from Ecuador, U. fernandeziana with the habit of a miniature tree
(229. 862) appears to lack near relatives. The family is, I suppose, of tropical
origin, but the actual centre of Uyézca is in the north temperate and subtropical
zones; there are 14 species in Eurasia, including the Mediterranean region, and
7 in North and Central America with a secondary centre in the Andes (Colomb.—
Fueg. 6); 3 species are tropical (Braz—Urug., Ethiopia, Java). There are a few
species in the south hemisphere, 2 S. Afr., 1 Australia, 2 N. Zeal. and 1 Auckl. Is.
U. fernandeziana seems to represent an ancient type.

Boehmeria excelsa was described by BURGER (47) as an elegant shrub which came
from S. America; it is a clumsy tree and not related to neotropical species. It
comes nearest to B. dealbata (Kermadec Is.) and points west, not east.

Parietaria humifusa (Chile); see above p. 204. Belongs to a neotropical group
which has not been cleared up (247) but is also closely related to the Australian-
Polynesian debz/zs of FORSTER. WEDDELL’s dedz/7s is a mixture of varieties scat-
tered over the globe and most likely consists of several good species.

Phrygilanthus Berteroz. The family Loranthaceae is tropical, but Phrygzlanthus
is of Antarctic origin. Of the 7 sections, 4 (about 20 species) are American and
range from Lower California to S. Chile, 1 with 2 species belongs to Australia
and New Zealand, 1 (1 species) to New Zealand, and 1 has one species in Aus-
tralia, another in New Guinea and a third in the Philippines—a typically austral-
bicentric genus.

The genus Saxtalum ranges from Australia and Melanesia to Micronesia and
Malaysia and east to Polynesia and Hawaii with a distant outlying station in
Juan Fernandez; see TUYAMA’s map (270). Of the 4 sections distinguished by

260 C. SKOTTSBERG

TuyaMA (who divided my section Lusaxfa/um in 2 and referred the Hawaiian
Eusantala to a separate section So/enanthus), Eusantalum s. str. is the largest with 9
species, Solenanthus and sect. Hawaitensza (together 8) confined to Hawaii (incl. Lay-
san), and sect. Polynesica (2, with varieties) to Polynesia; in this section the extinct
S. fernandesianum occupies a rather independent position. “Aber woher kam der
fremdartige Eremit Santalum?’’ BURGER (47. 22) exclaims, “zweifelsohne fern aus
ostindischen Meeren, wo die Wiege seines altberihmten Geschlechtes steht’’—
nothing could give a wronger idea of the history of the sandalwoods, because
everything points toward Antarctica as their cradle. There are related endemic
genera in Australia, J/zda in New Zealand, Exocarpus ranges from Australia across
Polynesia to Hawaii, and several endemic genera are at home in temperate S.
America (Arjona, lodina, Ovidia, Quinchamalium, and Nanodea in the extreme
south and in the Falkland Is.}; one is tempted to regard the family as of Antarctic
ancestry, but with Zheszuwm in mind it might be safer to speak of a special
Antarcto-tertiary centre.

The 3 endemic species of Chenopodium were commented upon above (p. 204).
In general appearance they are very like the Hawaiian cahuense but the special-
ist opines that they are not nearly related to this nor to other species. I can find
no better place for them than in a neotropical group.

Salicornia fruticosa, taken in a wide sense, is a thalassochorous plant scattered
along tropical and subtropical coasts; the same is the case with Jetragonia ex-
pansa in the S. hemisphere, but while Sa/zcornza is a world-wide genus, Jefra-
gonia has a stronghold in South Africa and several endemic species in Chile.

Spergularia is a wide-ranging, but mainly boreal genus with a vigorous branch
in Andean America but absent from Australia, New Zealand and Oceania. The
2 island species, of which S. conxfertifiora is also found on San Ambrosio, are
closely related to Chilean species. Paronychia has about the same distribution
pattern as Spergularia, but is poorly represented in S. America.

Ranunculus caprarum. A very large essentially boreal-temperate genus with
well-stocked branches in S. America and on New Zealand. The Masafueran en-
demic stands apart from its American congeners and approaches certain New Zea-
land species, perhaps also the Hawaiian ones. An Antarctic migration route seems
probable.

Berberts corymbosa and masafuerana belong to a small section confined to
the tropical Andes and not extending to Chile, where we have many other spe-
cies. The present area of the Berberidaceae testifies to its Arcto-tertiary charac-
ter; it centres in E. and S. Asia and ranges far south only in America, where 3
species reach Fuegia.

Of the six genera of Winteraceae (253), Bubbza has 2 species in Australia,
1 on Lord Howe I., 8 in New Caledonia and 19 in New Guinea, Belliolum 4
in New Caledonia and 4 in the Solomon Is., Pseudowintera 2 in New Zealand,
Exospermum 2 and Zygogynum 6 in New Caledonia, and Drimys 6 in Australia
{1 also in Tasmania), 29 in New Guinea and 1 on Borneo, Celebes and in the
Philippines, all these belonging to sect. Zasmania; the other section, Ludrzmys,
is American with 4 species, D. confert7folia endemic in Juan Fernandez. This sec-_

DERIVATION OF THE FLORA AND FAUNA 261

tion is a distant branch of an Australasian family and ranges from the uttermost
south along the mountains to Mexico and to Roraima in Brazil. That Antarctica
once played a role in the history of Drzmys is proved by the Tertiary fossils
discovered in West Antarctica.

Lactorts is generally looked upon as a primitive member of the Ranales and
claimed to belong to the most primitive element in the island flora. It has no
relatives in America. It is no typical member of the Magnoliales, an Arcto-tertiary
order, which it needs a CROIZAT to derive from the Antarctic. Geographically,
Lactoris is a parallel to Degeneria of Fiji, but the affinities of the latter are not
questionable and they have little in common. It lies near at hand to think of the
small family Lardizabalaceae, endemic in Chile. HUTCHINSON regards Lactorida-
ceae as ‘‘closely related to the Winteraceae, of which it is probably a reduced
derivative’ (zzo. II. 85)—but is the perfect trimery a result of reduction? And are not
the never quite closed carpels an indication of primitiveness? I daresay most sys-
tematists agree that the Polycarpicae, whether regarded as ove order or split up, are
among the oldest living angiosperms. The distribution of Winteraceae and the
occurrence in the South Hemisphere of small isolated families as Degeneriaceae,
Lactoridaceae and Lardizabalaceae suggests that the Antarctic continent was one
of the centres of evolution.

Cardamine is a world-wide, mainly temperate and essentially boreal genus,
extending into the tropics and south to Fuegia and New Zealand. The 3 island
species, one of them endemic, were commented on above (p. 205).

Among the numerous Chilean species of Fscallonia, £. Callcottiae stands a
little apart from the rest (758); the genus is spread along the Andes and extends
to Brazil and Uruguay. The subfamily Escallonioideae is austral-circumpolar: 772-
beles (1 S. Chile-Fueg.), Valdivia (1 S. Chile), Forgesia (1) and Berenice (1),
Réunion, Azopterus (2, Tasm., E. Austral.), Cuéts7a (1 E. Austral.), Argophyllum
(10 E. Austral., N. Caled.), Colmeiroa (1 Lord Howe I.), Carpodetus (1 N. Zeal.,
N. Guin.), Quéntinéa (15 Austral.-N. Guin.-N. Caled., Philipp.), Pottingerta (1 N.E.
Ind.)—a distribution suggesting an Antarctic origin.

Rubus geoides has a single near relative, R. radicans, in S. Chile. They differ
very much from the numerous north temperate and tropical montane species and
form their own section or subgenus, and their resemblance to the Tasmanian
R. gunnianus Hook. Icon. Plant. II is no proof of affinity. They seem to represent
an isolated offshoot from the north which has become cut off from its source of
origin and found its way into the subantarctic zone. ;

Acaena. Nobody if not CroIzAT would argue that Rosaceae are a southern
family, but this cannot prevent us from assuming that the Saxzguzsorba assemblage
of genera has gone through part of its evolution at least in the far south. BITTER
(3z) distinguished 10 sections: I, 13 species, S. Amer.; II, 1, The Cape; ELE, 1.2
tS, Amer., 1 Hawaii;-IV, 1, S: Amer.; V, 8, 6 S. Amer., 1 J. Fern., 1 Tasm:;
VIy 1, S: Amer.; VII; 28; 26 S. Amer.,:1 Galif., 1 Tasm:; VII, 64; 58°S. Amer,
1 Tristan da C., 1 N. Amsterd. I., 3 N. Zeal., 1 N. Zeal—Tasm.—Austral.—N. Guin.;
IX, 1, N. Zeal.; and X, 2, N. Zeal. Two Magellanian species occur on S. Georgia
and 1 on Kerguelen. The circumpolar distribution shows no gaps. About 70 %

262 C. SKOTTSBERG

of the species inhabit extratropical S. America, but many of them are so closely
related that they are little more than microspecies and in not few cases based
on one or two specimens from a single locality, and the number of separable
taxa will perhaps be reduced when more material becomes available. Be this as
it may, 7 sections are represented in America, 5 in the Australian—N. Zealand
area, 3 are common to both, one of them ranging to the African sector, where
another section is endemic. As regards dA. masafuerana see p. 206 above. I sup-
pose we can draw no other conclusion from this distribution than that Acaena is
an Antarcto-tertiary genus, 2 sections having developed numerous species in the
Andes and Patagonia.

Margyricarpus, a small Andean genus, is so closely allied to Acaena that they
have produced a bigeneric hybrid in Juan Fernandez. Two more genera are found
in the Andes, 7etraglochin and Polylepis. In S. Africa we have the large genus
Cliffortia. The remaining genera Sanguzsorba, Poteritum and Bencomia (Macarone-
sia) belong to the N. hemisphere.

Sophora sect. Tetrapterae is austral-circumpolar: New Zealand (3 species),
Chatham I. (1), Lord Howe I. (1), Austral Is. (1), Rapa (1), Marquesas (1), Hawaii
(1), Easter I. (1), Juan Fernandez (2), Chile (2; S. sacrocarpa, however, rather unlike
all the others), Diego Alvarez (1), and Réunion (1). With the exception of macrocarpa
and the Hawaiian chrysophyla the remaining species used to be united under
tetraptera Ait., otherwise endemic in New Zealand. They are very closely related,
but distinct; it is of minor importance if we call them species or geographical
subspecies. Unless we believe that S. zedvaptera was carried by water from island
to island and was transformed into a new species wherever it landed, we must
look upon Antarctica as a one-time centre of a polymorphous population, which
radiated in various directions; we shall not discuss here how this may have hap-
pened. We have not to do with litoral but with inland plants; the pods are
adapted to float, assisted by the four narrow wings, JOHOW says, but some of the
forms have no wings at all, and even if they have, the pods open on the tree and
discharge their seeds.

Fagara mayu and externa form their own section. When BURGER said (47. 19)
that /agara had migrated to Juan Fernandez from the primeval forests of Pert
and Colombia he overlooked that the affinity is with palaeotropical rather than
with neotropical species; there are numerous species scattered from Australia and
New Caledonia to Polynesia and Hawaii, where many are endemic. Rutaceae were
perhaps represented in the Antarctic in Tertiary times, and we have too look for
a route across to the American sector.

The family uphorbiaceae is pantropical, let alone that Lwphorbia has at-
tained a world-wide distribution and flourishes also in temperate climates. Dysopszs
is Andean, Se7delza (2) and Lezdesza (1) South African, the fourth genus of the
Mercurialis group, Mercurialis (8), ranges from North Europe to the Mediterranean
and is found in E. Asia. The southern genera seem to be more closely con-
nected mutually than with J/ercuralis. The disjunctions are interesting and difficult
to explain, unless we can find good reason to look for a common source in the
Antarctic.

DERIVATION OF THE FLORA AND FAUNA 263

The systematical position of Ca//itriche has been discussed many times, but
we know nothing of its history or where it started. The genus is world-wide, but
many species are not wide-spread and some are quite local, among them C. Lech-
feri, which may have been carried to Juan Fernandez from the mainland by
accident.

The tribe Colletieae of the otherwise wide-ranging Rhamnaceae is called
austral-antarctic by SUSSENGUT (Natiirl. Pflanzenfam. 2nd ed.): except Adolphia
(Mexico-U.S.A.) the genera are distributed over Andean and extratropical S. Amer-
ica, centering in Chile, and Colletta spartioides finds its place with the Andean
element. It should be mentioned that Dzscarza (11) extends south to Fuegia and
reappears in Australia (1) and New Zealand (1), suggesting transantarctic migra-
tion from America.

The family +/7/acourtiaceae is tropical; Azara is neotropical with about 19
species in Chile, 1 in Brazil and 1 in Argentina. The tribe to which it belongs
is well developed in the south hemisphere: tropical America and Africa, Mada-
gascar and neighbouring islands, Asia, New Guinea and (Ay/osma) Oceania to
Hawaii, but Antarctica may not at all be involved in its history.

Myrtaceae. A very large world-wide and tropical-subtropical family. Of the
subfamily Myrtoideae, some 2400 species, 75 % are American, the remainder scat-
tered over Asia, Africa, Australia and Oceania. The Leptospermoideae, some 850,
are restricted to Australasia with the single exception of 7epwa/ia, monotypical
and endemic in the Chilean rain forests, south to West Patagonia, a most inter-
esting case of disjunction. BERRY (26) regards the family to be of American ori-
gin and to have attained its present distribution before the close of the Creta-
ceous. Basing his conclusions on fossil evidence he thinks that during the cooling
down of the climate during late Tertiary, the ancestral stock of Myrtoideae with-
drew from North America to the neotropical zone; the Australian Leptospermo-
ideae represent the remnants of the Cretaceous radiation during which numerous
new types became evolved. Some of these eventually invaded Antarctica and
Tepualia survives on Chilean soil.

The Juan Fernandez Myrtoideae are closely linked to Andean types. For Ugwz
Selkirkit and Myrteola nummularia see p. 206. Nothomyrcia, now restricted to Masa-
tierra, may or may not have inhabited a larger area. Iyrceugenia has about 20
species in Chile.

Gunneraceae (often placed as a subfamily under Halorrhagidaceae) is a classi-
cal example of a tricentric Antarcto-tertiary type. Its long and complicated history
is reflected in its taxonomic differentiation; Gwuera is composed of 6 subgenera.
Pangue is the largest with 10 species ranging from Costa Rica to S. Chile, 1 in
Brazil, 3 in Juan Fernandez and 2 in Hawaii. Perpensum is monotypical with
separate varieties in S. Africa, British E. Africa and Madagascar, Ostenza an aber-
rant monotype endemic in Uruguay. MW/rsandra includes 3 species, one extending
from Colombia to Fuegia and Falkland, one restricted to the S. Chilean Andes
and one to subantarctic America; MWi//igania has 8 species in New Zealand and
1 in Tasmania. The monotypical Pseudogunnera inhabits New Guinea, Java, Su-
matra and the Philippines. No subgenus is found in more than one sector. It is

264 Cc. SKOTTSBERG

surprising that the geographically isolated Hawaiian species are closely related to
the species of Juan Fernandez. We know other examples of this connection, but
to construct a route between Hawaii and Juan Fernandez meets with serious obsta-
cles, and Pangue may have reached Hawaii along a quite different route. The
evolution and differentiation of the subgenera very likely took place in Antarctica.

SCHINDLER (Pflanzenreich) expressed his opinion of the history of Hador-
rhagidaceae thus: ““Aus der geographischen Verbreitung der urspringlichsten Halor-
rhagaceen, namlich der Gattung Halorrhagis, ist mit Sicherheit zu folgern, dass die
Familie antarktischen Ursprungs ist.’ It is not easy to see how he arrived at this
conclusion, though I think it is correct, because at that time the endemic species
of Juan Fernandez were unknown; they passed as //. erecta, a New Zealand en-
demic, and this, incorrectly attributed also to Juan Fernandez and Chile, had been
carried there on purpose: “Der Standort auf Juan Fernandez und in Chile ist kein
urspriinglicher, sondern durch die auch in Neuseeland erfolgende Verwendung der
Pflanze als Futterpflanze erklart.’’ But there was no //a/orrhagis on the mainland,
and the species indigenous in Juan Fernandez were not used as forage, let it be
that the introduced animals eat them. H/alorrhag7s, eminently Australian (59 out
of 80 species) and with 7 species in New Zealand, extends north to Indomalaya,
Micronesia, S.E. China and Japan and east to Rapa (not known to SCHINDLER)
and Juan Fernandez—see TUYAMA’s map of distribution (277). The genus “well
illustrates the not infrequent extension of an Australasian group far north of the
equator, and the much rarer condition of occurrence in Juan Fernandez but not
in continental America’ (zog. 108). Halorrhagis shows the same distribution pat-
tern as Santalum.

Centella is essentially African, see above p. 207; the distribution is tricentric
with some remote stations. The widespread C. aszatica is scattered over a broad
belt but not reported from America. Very likely the Hydrocotyloideae, a sepa-
rate family according to some authors, are of Antarctic origin, but C. ¢v7flora
may well have reached Juan Fernandez with the traffic from Chile.

The peculiar endemic species of Axvyxgzum differ so much in habit from all
other species of this large and widespread genus that they have been referred
to a separate genus, a rank to which they are not entitled. Exyxgzum concen-
trates in the Mediterranean region and in tropical South America, where also the
island species belong, in spite of their arborescent habit; to quote TURMEL (269.
130): “Se rattachant a la Région chilienne, on cite les espéces de Juan Fernandez
£. bupleuroides, sarcophyllum et tmaccessum..., plantes, du moins pour les deux
premicres, arbustives s’opposant radicalement aux autres especes des territoires
voisins —but 7zzaccessum also is a dwarf tree, more so I would say than sarco-
Phyllum. | doubt that they descend from herbaceous continental forms; they be-
long to an ancient type and find their proper place with the neotropical element.

The tribe Apioideae-Ammineae is very widely spread, with a concentration
in the North hemisphere, Afzwm distributed also in the south temperate zone, and
A. graveolens L. is frequently regarded as bipolar species. I don’t believe that
any of the southern forms should be included; they are, however, in need of
revision. A. fernandezianum is a well-marked species, related, but not very closely,

DERIVATION OF THE FLORA AND FAUNA 265

to a series of forms, probably good species, reported from subantarctic America,
Falkland, Tristan da Cunha, Australia, etc. and possibly descending from an
old Antarctic stock.

Pernettya: 13 species, 8 Mexico and Centr. America to Chile, south to Fuegia
and Falkland, 1 Galapagos Is., 2 in Tasmania and 2 in New Zealand; P. rigida
is linked to Andean species, but very distinct (252). The genus is more diversi-
fied in the American sector, and this seems to be where it originated, having
reached New Zealand across the Antarctic, if not with Gau/therza, well developed
in New Zealand, descending from a common Antarcto-tertiary stock.

Empetrum is a bipolar genus, the family most likely of boreal origin. Con-
cerning /. rubrum see 249.781.

Of the 7 Drchondra species 5 are neotropical, 1 endemic in New Zealand
and PD. repens (incl. servzcea) spread round the world. It is common on the coast
of Chile and possibly adventitious in Juan Fernandez. The occurrence of an en-
demic species in New Zealand suggests that Antarctica witnessed part of its
history.

Calystegia. About 25 species have been described, scattered over the globe,
C. sepium sensu lat. reported from America, Eurasia, N. Africa, Australia, New
Zealand, Easter I., etc. and evidently very easily naturalized. The plant found
on Masafuera and also on the mainland was described as C. Hantelmanni Phil.
and later identified with tgurzorum from New Zealand; see 249. 783. If this is
correct, C. tuguriorum offers one of the very few cases of a species restricted to
Chile and New Zealand, but even if they are kept apart, they present a remark-
able case of disjunction.

Selkirkia Berterot, the only representative of Boraginaceae, so richly devel-
oped on the mainland, was regarded as an isolated, Old Pacific type (227. 31, 224.
593) until JOHNSTON (748) showed that it comes close to //ackelia and differs from
this principally by its arboreous habit. With Uréica fernandeszana, the species of
Eryngium etc. I refer it to the neotropical element.

Rhaphithamnus venustus is of neotropical ancestry (see above p. 208); the
second species is common in Centr. and S. Chile.

BRIQUET placed Cumznia in the Stachyoideae-Menthinae next to Oreosphacus
Phil., a shrub of the high Cordillera in the boundary region between Chile and
Argentina, but this genus has a schizocarp of four nuts. It has also been com-
pared with Bystropogon L) Hérit. (Canary Is., Colomb.—Pert, different sections). Ep-
LING (using the fancy name Yohow7a) referred Cuminia to Prasioideae, an ancient
group showing great disjunctions: Praszwm Mediterranean, Stexogyne, Phyllostegia
and Haplostachys in Hawaii, Bostrychanthus and Gomphostemma in Asia; Cuminia
differs in the shape of the corolla (see above p. 208), but even so it seems to repre-
sent a palaeotropic element in the island flora.

Solanum fernandezianum is a distinct species of indubitable neotropical and
Andean ancestry. With regard to S. vobzxsonianum, see above p. 200.

Nicotiana cordifolia has, according to GOODSPEED (z72. 347), its closest resem-
blance, in flower structure as well as in general habit, to V. Ratmondi. Crosses

266 C. SKOTTSBERG

with this species and with solanifolia gave evidence of fundamental affinities between
the island endemic and species of the mainland to the north (l.c.).

Mimulus is essentially a western N. American genus with few species else-
where: the somewhat polymorphous J. glabratus ranges from N. America to
Bolivia, Chile and Argentina.

The different opinions on the systematic position of Luphrasia formosissima
have already been referred to above. Within the area of the geographically isolated
Chilean-Magellanian 77zfidae occurs the semicalcarate /. perpuszlla Phil. (S. Chile).
Both would indicate a road from the Australian-New Zealand area across the Ant-
arctic to America, just as the tropical mountains of Malaysia served as a road
between Asia and Australia, as DU RIEYZz thinks (77. 536): “The Euphrasia popu-
lation of Juan Fernandez may therefor very well have formed the northern end
of a population so far south that the lack of close relationships between £. for-
mosissima and the species of Middle Chile is fully explained, and the semical-.
carate anthers of the more southern L. perpuszlla may be the last South American
remnant of this old connection.’ Another question, not yet answered, is this: are
the very well-marked 777fidae, reaching from subantarctic America to the Andes
of S. Chile, likewise descendants from a remote southern population or did they
originate in Chile? .

Plantago fernandezia is another arborescent member of a herbaceous world-
wide genus and often cited as an example of a mysterious connection between
Hawaii and Juan Fernandez, but if we look at the total distribution of sect. Padlaeo-
psyllium (see above p. 209) its character of an austral group is revealed, even if
it extends north to N. America and S. Europe. In the south it is circumpolar,
and the route from New Zealand (Auckland Is.) via Rapa to Hawaii can be traced
—a radiation from Antarctica seems not unthinkable.

Plantago truncata, represented in Juan Fernandez by an endemic (?) variety,
belongs to the large section Novorbzs and needs no further comments.

The neotropical Hedyotis thestifolia is a recent addition to the flora and may
have been accidentally introduced.

Nertera is an austral-circumpolar genus allied to Coprosma and best devel-
oped in New Zealand where 5 species occur, 4 of them endemic; the fifth is
N. granadensis (depressa), claimed to be very widely spread. One species is en-
demic in Tristan da Cunha. Recent observations tend to show that granadensis
includes taxonomically distinct forms; the Malaysian plant is not identical with
the Andean, and other forms will perhaps become distinguished after a critical
revision. Be this as it may, the genus is Antarcto-tertiary and, if we link Tristan
da Cunha with Africa, tricentric.

Coprosma is a parallel to Halorrhagis but differs in being present in Hawaii;
another difference is that the species of Juan Fernandez do not have their closest
relatives in New Zealand or Australia—Hookerz forms its own section, pyrifolia
is of Polynesian affinity. As the genus is absent from America as well as from
Africa it should lie near at hand to refer it to a West Pacific element, a position
favoured by the relationships of pyrzfolza, and to the believers in transoceanic
migration combined with evolution of local endemics wherever Coprosma happened

DERIVATION OF THE FLORA AND FAUNA 267

to land this is the only course to take, even if the sudden appearance of an iso-
lated type like H/ookerz becomes somewhat embarrassing. With an extension north
and east of an Antarctic borderland Cofrvosma could pass as of Antarcto-tertiary
origin.

C. pyrifolia offers a good example of an incorrect taxonomic position leading
to false conclusions. It was described as Psychotria and referred to a neotropical
group: it came, as BURGER writes (gz. 20), “‘von den Urwaldern Perus und Co-
lombias’’.

Of the more than 500 species described under Gadium about 400 are distri-
buted over the boreal zone and some 50 or 60 are known from South America,
extending from the tropical Andes to Fuegia, the Falkland Is. and South Georgia;
about 30 are African, the remaining species being divided between India, Malaysia,
Australia, New Zealand, etc. Gal/ium is a boreal genus with a strong represen-
tation along the Andes, a not uncommon case. G. masafueranum is allied to
species from the mountains of Central Chile, probably also to species found farther
north. Nothing indicates that Antarctica ever had a share in the history of this
genus.

Whether Campanulaceae-Lobelioideae evolved in the tropics or in the far South
will not be discussed here. The remarkable concentration of arboreous genera,
two of them large, in Hawaii, has given rise to much speculation, and the occur-
rence of other endemic genera in Polynesia as well as the Australian affinity of
Brighamza of Hawaii have led some authors to look for the origin of the sub-
family (or family) in the Antarctic. In our special case we can leave this question
aside because Lobelia alata is a widely distributed seaside species, tricentric in
the South Hemisphere.

The distribution of Wahlenbergia, a large and widespread genus, is inter-
esting. It is essentially southern, of the about 230 species described 150 are South
African and 20 tropical African, 6 are reported from Madagascar and the Masca-
rene islands. North it extends to the Mediterranean, the Orient and Asia, together
some 25 species. Scattered species are known from New Guinea (1), Australia (7),
New Zealand (7), Lord Howe I. (2) and St. Helena (3). Species are few in America
{North Amer. 1, South Amer. 8); a single species, WV. gracilz7s, ranges over the
south hemisphere.

If we look at the related genera, some 13 in number, the dominant position
of South Africa becomes still more conspicuous: 7 are exclusively or preponder-
antly South African, 1 is from tropical Africa, E. Indies and Brazil, 4 Asiatic
and 1 from S.E. Europe, all according to the old synopsis in Natirl. Pflanzenfam.
Our island species are, as we have seen, not matched in Africa, nor in America
or other parts of the world except on St. Helena, see above p. 210. All known
facts suggest that the genus had a long and complicated history in the far south
and that S. Africa is a secondary centre where evolution has been progressive
and prolific in species formation. It is important to remember that our island forms
are not a group of closely related taxa but that Il’. Wasafuerae and particularly
Berteroi stand apart from the rest. The distribution of the genus can hardly be
explained without admitting Antarctica into the picture.

268 C. SKOTTSBERG

The distribution of Lagenxophora clearly testifies to its Antarctic origin: 3
Magellanian species, one of them also on Tristan da Cunha and one on Masafuera,
7 in New Zealand, 2 in Australia, one of these north to the Philippines, 1 in Fiji
and 3 in Hawaii, the four Pacific ones connected with ZL. Bl/ardieri Cass. (Austral.),
whereas the Magellanian species point toward L. pumila Cheesem. and fetiolata
Hook. fil. (New Zealand).

In Evigeron we meet again with a large boreal genus strongly represented
along the Andes and ranging south to Patagonia, Fuegia and Falkland; 135 species
are reported from Eurasia, incl. India, 345 from North America, 35 from Centr.
America and the W. Indies, 95 from S. America, 10 from Africa, 8 from Australia
and a single species from Oceania (Rapa), /. rapenszs F. B. H. Brown, and this is
compared with our island species. To judge from the description it has the same
habit as these, a shrub about 3 dm tall with the leaves in terminal rosettes. In
Hawaii the genus is represented by Zetramolopium, which is very close to Erigeron;
VIERHAPPER (229. 182) suggested that the Juan Fernandez species came near the
Hawaiian &. lepidotus Less., which is now referred to 7etramolopium, and that we
ought to look for relatives among the Andine and Mexican forms. Among our
island species /. rupicola differs much in habit from the rest, but all belong together
and most likely represent a special branch of the Andine /yégeron flora which, in
its turn, comes from a boreal stock.

To what I said above about Guaphalium spiciforme nothing can be added
at present. Chile is well provided with poorly limited species related to G. pur-
pureum \.; most of them are badly known. Their boreal parentage can hardly
be doubted.

Abrotanella resembles Lagenophora in its distribution, but is absent from Oce-
ania; the majority inhabits New Zealand with its subantarctic islands (9 sp.), I is
found in Victoria, 2 in Tasmania, 1 in New Guinea and 5 in S. America (W.
Patagonia to Fuegia and Falkland); in addition, one is found on Rodriguez I.
The genus is, as it were, tricentric and its Antarcto-tertiary character indisputable.

The concentration of isolated arborescent Compositae in the Pacific was em-
phasized by BENTHAM; the main groups of the family are represented among them,
and the accumulation of endemic genera in Hawaii and Juan Fernandez has led
to much discussion. The enormous development and differentiation in the family
on American soil is an undeniable fact, but simply to derive the endemic Pacific
genera from America as did Guppy (z2z) does not seem possible. Speculations (comp.
f. inst. SETCHELL 279) led back to Antarctica, but not until Brachionostylum was
discovered in New Guinea and found to be nearly related to Rodzsonza of Juan
Fernandez were we able to stand on tolerably firm ground.

With regard to Centaurodendron, to which Yunqguea seems to be related, the
situation is different. Cextaurea and all the genera of Centaureinae belong to the
Old World with the exception of a single species in N. America and a few in
the Andes, south to Centr.’Chile. The group ranges over Europe with a strong-
hold in the Mediterranean, the Orient, Ethiopia and through Centr. Asia to Japan.
The Chilean species of Centaurea belong to the Plectocephali, but Centauroden-
dron differs not only from this section but from all in sex distribution (243). It

DERIVATION OF THE FLORA AND FAUNA 269

serves no purpose to say that Cezfaurodendron descends from some continental
Centaurea which became arboreous under insular conditions, because the character
of the ray floret is entirely opposed to such a theory and Centaurodendron seems
to be a more ancient type than Cezzfaurea, a relic from an epoch previous to the
final uplift of the Andes.

The four Cichoriaceous genera, to which 7/amnoserzs of the Desventuradas
Is. shows some slight affinity, are even more isolated. The only other genus to
which they bear some resemblance is the Polynesian /7¢chza, but the differences
are too profound to allow us to visualize an Antarctic-Pacific ancestry of the Dev-
droservis group and it remains us to link it with the neotropical element.

Referring to the synopsis given above the Phanerogams are rearranged in
the following way according to their supposed source of origin.

I. Antarcto-tertiary element.—62 sp. (42.2 %).

1. Distribution pattern austral-circumpolar, bicentric or tricentric.—33 sp.

a. Endemic species (17): Cladium scirpoideum, Uncinia Douglasii and costata,
Carex berteroniana, Luzula masafuerana, Drimys confertifolia, Phrygilanthus Ber-
teroi, Acaena masafuerana, Sophora fernandeziana and masafuerana, Gunnera pel-
tata, Masafuerae and bracteata, Apium fernandezianum (?), Pernettya rigida, Plan-
tago fernandezia, Abrotanella crassipes.

6. Species also found on the continent (16): Danthonia collina, Koeleria micra-
thera (?), Oreobolus obtusangulus, Uncinia brevicaulis, phleoides and tenuis, Juncus
imbricatus, capillaceus, acutus, dombeyanus and planifolius, Libertia formosa,
Acaena ovalifolia, Centella triflora, Nertera granadensis, Lagenophora Harioti.

2. Genera only found in S. America, but supposed to be of Antarctic origin.
The species are endemic: Escallonia Callcottiae, Margyricarpus digynus.—2 sp.

3. Endemic genera or species as far as known without continental American
affinities, either suggesting an ancient Antarcto-Pacific track east from Australasia
without reaching America, or having arrived along the road over the Scotia Arc
without leaving any traces in the present S. American flora.—21 sp.

a. Belonging to endemic genera (7): Robinsonia gayana, thurifera, evenia, Masa-
fuerae and gracilis, Symphyochaeta macrocephala, Rhetinodendron Berteril.

6. Endemic species of genera of wider distribution (14): Peperomia berteroana,
margaritifera and Skottsbergii, Santalum fernandezianum, Boehmeria excelsa, Ra-
nunculus caprarum, Fagara mayu and externa, Halorrhagis asperrima, masatier-
rana and masafuerana, Euphrasia formosissima, Coprosma Hookeri and pyrifolia.

4. Endemic species with relatives in the African sector only: Wahlenbergia
Larrainii, fernandeziana, Grahamae, Masafuerae and Berteroi.—5 sp.

5. Endemic family, possibly of old Antarctic, perhaps pre-Tertiary ancestry:
Lactoris fernandeziana.—1I sp.

II. Neotropical-Andean element.—5 4 sp. (36.7 %).
1. Endemic genera or species of neotropical parentage; non-endemic species
Seuth: Amencan-——31 sp:
a. Belonging to endemic genera (4): Juania australis, Ochagavia elegans, Notho-
myrcia fernandeziana, Selkirkia Berteroi.

270 C. SKOTTSBERG

6. Endemic species of non-endemic genera, some of wider distribution (14):
Chaetotropis imberbis, Chusquea fernandeziana, Hesperogreigia Berteroi, Urtica
Masafuerae, Dysopsis hirsuta. Colletia spartioides, Azara fernandeziana, Ugni Sel-
kirkii, Myrceugenia Schulzei, Rhaphithamnus venustus, Solanum fernandezianum,
robinsonianum and masafueranum, Nicotiana cordifolia.

c. Species also found on the continent (13): Stipa neesiana and laevissima,
Piptochaetium bicolor, Chaetotropis chilensis, Cyperus eragrostis and reflexus, Eleo-
charis fuscopurpurea, Peperomia fernandeziana, Parietaria humifusa, Myrteola num-
mularia, Mimulus glabratus, Plantago truncata, Hedyotis thesiifolia.

2. Endemic genera or endemic sections of wide-ranging genera without relatives
in the present continental flora but supposed to descend from extinct neotropical
ancestors.—23 sp.

a. Belonging to endemic genera (16): Podophorus bromoides, Megalachne ber-
teroniana and masafuerana, Centaurodendron dracaenoides, Yunquea Tenzii, Den-
droseris macrophylla, macrantha, marginata and litoralis, Rea neriifolia, micrantha
and pruinata, Phoenicoseris pinnata, berteriana and regia, Hesperoseris gigantea.

6. Belonging to endemic sections (7): Urtica fernandeziana, Chenopodium
Sanctae Clarae, crusoeanum and nesodendron, Eryngium bupleuroides, inaccessum
and sarcophyllum.

Ill. Arcto-tertiary element.—23 sp. (15.6 %).

Genera essentially boreal but extending south along the Andes or reappearing in S.
America.

a. Endemic species of wide-ranging genera (13): Agrostis masafuerana, Sper-
gularia confertiflora (also Desventuradas) and masafuerana, Berberis corymbosa
and masafuerana, Cardamine Kruesselii, Galium masafueranum, Erigeron fruticosus,
luteoviridis, Ingae, Innocentium, turricola and rupicola.

6. Species also found on the continent (10): Trisetum chromostachyum, Carex
Banksii, Paronychia chilensis, Cardamine chenopodiifolia and flaccida, Rubus geo-
ides, Callitriche Lechleri, Empetrum rubrum, Calystegia tuguriorum (?), Gnaphalium
spiciforme.

IV. Palaeotropic element.—2 sp. (1.4 %).

Endemic genus: Cuminia fernandezia and eriantha.

V. Austral element of wide-ranging seaside species.—6 sp. (4.1 %).

Scirpus nodosus and cernuus, Salicornia fruticosa (peruviana), Tetragonia ex-
= a «
pansa, Dichondra repens, Lobelia alata.

Even if Antarctica is recognized as an important source of evolution of both
plants and animals and as a centre from where large-scale migration took place,
a percentage of 42 may seem surprisingly high, and I admit that some genera
or species have been referred to group I with considerable hesitation. This does
not, with very few exceptions, apply to I.1, 22.4% of the total or 53.2% of I.
Luzula and Funcus were placed here because the family was regarded as Antarctic,
but even if this be true it is possible that the sections including our species are

DERIVATION OF THE FLORA AND FAUNA 271

of boreal origin and has spread south. Pernet/ya is a bicentric genus, but it
is much better developed in America and may have originated with Gaultheria
in the Antarctic. The same should apply to \7zcotiana according to MERRILL
(306. 310) who thinks that its actual distribution was attained in the Tertiary pe-
riod by way of Antarctica; however, JV. cord?folia cannot be removed from II. 1.b
and find a better place with I. 1.a. On the other hand, a truly Antarctic genus
may have produced numerous species in one sector and few in another, or the
few may be a remnant of a larger population. In group I. 3 Peperomia berteroana
calls for attention. As I have shown (245) it is so close to P. ¢réstanens7s that
their common origin cannot be doubted and that little prevents us from regarding
them as forms of ove species, very distinct but clearly related to the other two
species endemic in Juan Fernandez, and they point west. This is the reason why
all three were referred to I. Wahlenbergia (I. 4) is another mysterious case, but as
it is neither boreal nor palaeotropical or neotropical but South African (and on St.
Helena), only a far southern ancestry remains to explain the disjunction.

Of the endemic genera belonging to group II. 1 only Yuanza is taxonomically
isolated, Rhodostachys of Chile is brought to Ochagavia by some authors, and
Selkirkia comes very near Hackelia. With regard to the endemic Cichoriaceous
genera authors’ opinions differ. BENTHAM looked upon the arborescent Pacific
Compositae as relics of an old Polynesian flora but did not refer directly to Ant-
arctica as source; GUPPY (727) quoting BENTHAM believed that, with the Hawatian
Lobelioideae, they belonged to an ancient flora of the Pacific which had origi-
nated in America and gained dominance during what he termed “Age of Lobe-
liaceae’’ and ““Age of Compositae’’, respectively. Antarctica seems not to have
meant anything at all to him, but as I have discussed his ideas in some detail
in an earlier paper (248), I shall not enter upon this subject here.

Calystegia tuguriorum is doubtful case, but few people will be inclined to
think that it originated independently in Chile and New Zealand.

In a paper of 1928 (278) SETCHELL discussed what he called the two prin-
cipal elements in the Pacific flora, the Indomalayan and the ‘‘Subantarctic’. With
this he did not understand what, from a geographical viewpoint, I call subant-
arctic, nor the species of old Antarctic genera found in the bogs of Hawaii, the
Pacific flora with which he was particularly concerned, but the group he later
(279) called “the Old pacific and antarctic element’, for it included also the ar-
boreous Lobeliaceae and Compositae, which he believed had a common origin
in high southern latitudes. They had migrated north along different lines, the
Compositae taking an easterly course along the route Juan Fernandez—Hawaii, the
Lobeliaceae, which are absent from Juan Fernandez but have left traces in Poly-
nesia, a more westerly; a third line is called the Dammara (Agathis)—-Podocarpus
line, an Australasian line running over Fiji. What interests us here is that the
isolated genera of Compositae were regarded as Antarctic. As we have seen, |
did not venture to include the Cichoriaceous genera, but I cannot assure that
SETCHELL was wrong.

272 C. SKOTTSBERG

Before quoting some other authors who have paid special attention to the
Antarctic problems I want to add a few remarks on the relations between the far
South and South America—Juan Fernandez. The Antarctic flora invaded the Amer-
ican continent, advancing especially along the rising Andes and in some cases
extending north of the Equator. It must have been a climatically diversified flora,
for the great southern continent must have had coast and inland, lowland and
mountain climates, and consequently the Antarctic plants in S. America have
varying claims on moisture and heat. The less exacting plants are concentrated
in West Patagonia, Fuegia and the Falkland Is., rising higher and higher as they
advance north along the Cordillera, whereas such as require more favourable tem-
perature conditions are found farther north in S. Chile and in the montane region
of the tropical Andes. Father RAMBO, a wellknown expert on the flora of S. Brazil,
called my attention (letter of Nov. 30, 1953) to an Antarctic element in the high-
lands of Rio Grande do Sul, where such genera as Acaena, Margyricarpus, Es-
callonia, Gunnera, Drimys and Phrygilanthus are represented; in 202. 30 he mentions
Araucaria angustifolia, Podocarpus Lamberti, Drimys Waintert, Acaena fuscescens,
Fuchsia regia, Gununera manicata, and Griselinia ruscifolia, elements which, in his
opinion, constitute the last remnants of the old flora that inhabited the southern
lands united until early Tertiary times. |;

HILL (734) was cautious in his judgment of the importance of Antarctica in
the history of the plant world. He quotes SEWARD who had pointed out that a
number of families now largely in the South Hemisphere were present in the North
Hemisphere already in Mesozoic times and that this would lead us to derive them
from there, which most likely is true in certain cases. He also refers to SMUTS,
who looked at the “‘ancient lands of the Southern Hemisphere’ as the cradle of
the peculiar S. African flora, and HILL formulates the following question (p. 1497):

Did the angiosperms originate in the north and migrate southwards and then, having
reached the south, evolve along special lines in lands comparatively isolated from the
land masses in the north, or did certain groups first appear in the South Hemisphere
in an ancient Antarctic Continent and become dispersed northward into our present-
day New Zealand, Australia, South America and South Africa?

With regard to Juan Fernandez he finds that

the present flora suggests that at one time these islands formed part of the Antarctic
continent or were united to the extreme south of Chile, a view which is shared “by
Jhering and Joly, who believe that such a connection may have existed in early Ter-
tiary times and that Kerguelen Island was probably part of a large land mass at the
same period...

and he continues:

It seems also likely, on geological evidence, that Antarctica extended in the Ter-
tiary epoch towards Tasmania and Australia, and so to Asia, and towards New Zea-
land, and the distribution of certain present-day plants in the Australasian region lends
considerable support to such an extension (p. 1479).

Nevertheless, the problem is not as simple as it might seem to be:

DERIVATION OF THE FLORA AND FAUNA 273

We must be prepared to modify our views extensively in those cases which appear
to afford fairly certain evidence of distribution from a Southern centre, but from the
evidence available it seems probable that such genera as Calceolaria, Fovellana, Fuchsia,
Ourisia, Pelargonium, Caltha, Lilaeopsis, Gunnera, Hebe, Pernettya, Azorella, Drapetes
and other Thymelaeaceae, WVothofagus, Eucryphia, Laurelia, probably the Proteaceae,
some of the Ericaceae ... may have originated in some continental area in the South
Hemisphere, whence their descendants spread northwards...

It is of course possible that in early times many of the plants from which these
Southern Genera evolved belonged to the Northern region and gradually migrated to
the south and having reached the Southern regions they there, in a state of some isola-
tion, evolved peculiar types which in their turn spread again northwards ... without
however crossing the Equator and reaching the North Hemisphere. On this view we
may perhaps best regard many of the present day genera which are now only found
in the South Hemisphere (p. 1480).

He adds,

It may be almost impossible in these latter days to make any certain pronounce-
ment as to which natural families or genera, if any, actually originated in the Southern
Hemisphere, when it is remembered that there have been a succession of alternations
of warm temperate and cold glacial periods at various geological epochs not only in
the Arctic but also in the Antarctic regions.

He thinks that the cold spells gave rise to an extensive migration south and
north, respectively, but he did not recognize the glaciations as contemporary in
the two hemispheres.

Of other modern authors, W. H. CAmp (43) interests us here. My Antarcto-tertiary
element lists Drzmys as a classical example of an Antarctic genus; in fact, the
entire family Winteraceae is supposed to be of Antarctic origin. On CAMP’s map,
however, the original seat of the family is placed in the region of the old Bering
Land bridge, from where tracks lead south through Asia to Australia-Tasmania
and through western N. America to S. America. The related Magnoliaceae are
circumboreal and have left numerous fossil remains in the north. Another map
shows Laurelia, Eucryphia, Luzuriaga and Fovellana, all as a rule considered to
be Antarctic; here CAMP has two alternatives, one track leading from the north
as in Dyzmys, another from the south. Other southern families shown on his maps
are Tetrachondraceae, Eucryphiaceae, Centrolepidaceae, Epacridaceae (Hawaii in-
advertently left out), Stylidiaceae, Restionaceae, Liliaceae-Milliganieae, Philesiaceae,
Halorrhagidaceae and Gunneraceae. He emphasizes the fact that of the natural
families, some 300 in number, 103 have a fairly restricted range, and that 80 of
these are on the South Hemisphere. I shall quote him in full.

In the majority of instances an analysis of a group indicates that the primitive
members are on the southern part of the group’s range. If we were to follow the well-
known dicta of Matthews we could conclude that these primitive forms were “‘driven’’
unto the southern land mass extremities by the more highly specialized, better adapted,
and more recently derived groups of the north. The natural corollary to this, therefore,
is that we should find the majority of the peculiar specialized and more recently derived
families farthest away from the primitive forms. However, as can quite easily be seen
in the angiosperms, in most instances this is not the case. I therefore incline to the
conclusion that these southern land masses are not only the original homes of the great

18 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

274 C. SKOTTSBERG

majority of our basic angiosperm groups, but that also it has been on these same southern
land masses where the greater part of their primary evolutionary divergences took place
(pp. 180-18 1)

and he continues,

It seems likely that the angiosperms, as a group, arose on this southern land mass
contemporaneously with the Paleozoic of the northern (Holarctic) land mass and that
the divergences of the basic, generalized familial groups had been accomplished in this
southern land mass certainly by the mid-Mesozoic.

This would make the Jurassic the great period of ascendency of the angiosperms.

Camp's theory does not, at least at first sight, agree too well with his opinion
on the origin of the Magnoliales, always regarded as a primitive group though
not of restricted range; the Winteraceae have a stronghold in New Guinea—Australia—
New Zealand, and the actual distribution of this family testifies to a southern origin.

Recently STEBBINS (257) took up the question of the history of the Ranales.
He points out (p. 8) that this order

includes a high proportion of species which on the basis of all characteristics must be
placed not only in monotypic genera but even in monogeneric or digeneric families.
They are obviously relict types of which the close relatives have long been extinct... .
Finally, distributional studies show that the genera and species are at present strongly
concentrated in eastern Asia and Australasia, and at least one family, the Winteraceae,
may have radiated from the latter center (A. C. Smith 1945). This family was dispersed
through the Antarctic regions. .

We cannot deny the possibility that New Guinea was the birthplace of types
that now generally pass as Antarctic, a theory first advanced by Miss GIBBS (106)
and perhaps strengthened by the sensational discovery there of a great number of
Nothofagus species, but it is quite clear that, if they are bicentric, they must have
migrated across what is now the Antarctic continent in order to arrive in S. America
which again compels us to assume that connections were established on both sides,
and the existence of fossil plants belonging to taxa now living in the two sectors
furnishes additional evidence that Antarctica took an important part in their history.
Anyhow, Wothofagus survives in greater variation in New Guinea than either in
New Zealand or in Chile, though these are situated nearer to the Antarctic continent.

Before Antarctica was recognized as a possible centre of evolution, the Holarctic
region was claimed as the great and only cradle of temperate plant families which
spread south during the Tertiary just as, in pre-Tertiary times, tropical families
had extended to the present Arctic. GORDON (773), although he finds undeniable
proofs of an old Antarctic radiating centre, carefully scrutinizes the possibility of a
northern origin of temperate types now found in the south, referring to OLIVER’s
theory that a genus like Nothofagus originated in N. America and that two tracks
lead south, one to subantarctic America, one across the Bering land to Australia
and New Zealand, but he doesn’t think it probable that we can explain the dis-
junctions of numerous taxa in the south in this way and that the chances for cool
climate types to cross the broad and well-stocked tropical belt must have been small.

To me it would seem more acceptable to fall back on an ancient pantropical

DERIVATION OF THE FLORA AND FAUNA 275

flora, spread from pole to pole before any temperate zones had become sharply
delimited and when the distribution of land and sea was quite different from what
it is now. With the appearance of distinct climatic belts a sorting out of meso-
and microthermic groups followed, and these belts, both north and south, started
to produce their own particular new groups which dispersed toward the equatorial
zone and eventually met and passed it where mountain ranges offered a passage.
This is in agreement with FLORIN’s opinion (95) that the temperate floras, angio-
sperms as well as gymnosperms, developed independently throughout the Tertiary
period in the north and south hemispheres. The march of Antarcto-tertiary types
north corresponds to the march of Arcto-tertiary types south.

Recently Antarctica and the far south in general as the birthplace of the world’s
floras has found an eloquent advocate in LEON CROIZAT (7z), who throws all other
speculative authors in the shade. His ideas were criticised by SPARRE (256), but,
as Juan Fernandez is regarded by CROIZAT as “one of the most interesting domains
of general phytogeography”’, I cannot pass him in silence.

The original southern plant world, CROIZAT says, spread over the globe in
a way not to be deducted from the present map; we have to go back to the map as
it looked during the Cretaceous, but he does not tell which of the constructions
he prefers, only that he dismisses WEGENER’s theories; that everybody who devotes
himself to the study of plant migrations should pay attention not only to the present
configuration of the continents goes without saying. His own ideas of Cretaceous
geography in the south are expressed on p. 252:

It is most probable, sure we would venture to say, that an ancient shore connected
every land in the deep south of our maps, not only, but that the lands, now vanished,
but once extant between the approach of Kerguelen Islands and Magellania are the ultimate
hub of angiospermous dispersal.

Here extends the Patagonian-Mascarene baseline with its two triangles, Natal-Ker-
guelen—Tristan da Cunha (the Afro-antarctic triangle) and Ceylon—Madagascar-—
Mascarenes, the ‘“‘hub’” spoken of above. CROIZAT points at South Georgia as a
proof of the existence of an old Antarctic flora:

The theory that a nebulous ‘Glacial Epoch’ killed off very nearly all the ancient
vegetation of the antarctic islands is shown to be false by the comparatively large plant-
world still endemic in South Georgia, indeed a sizeable flora if we consider that it thrives
in what is now hardly better than a glaciated mountain straight out of the ocean (p.255)

Now, CROIZAT otherwise deals with the phanerogams only; there is not a single
endemic species in South Georgia, but there are many endemic bryophytes and
lichens. And he forgets to mention that the ‘‘nebulous”’ glacial epoch has left its
very distinct marks there, because we know that the lowland was ice-covered where
the phanerogams now form closed communities and that it is very improbable that
the higher flora survived. It is surprising that, in this connection, he does not men-
tion Kerguelen where endemic genera such as Prizglea and Lyadia surely survived.

A second baseline indicates the extension of an Antarcto-Gondwana in the
S. Indian Ocean, a third lies along 60° S. and runs from south and east of Chatham
Is. to contact S. America. From these centres the angiosperms started to disperse

276 C. SKOTTSBERG

during late Jurassic and very early Cretaceous times through three “gates’’ in three
main streams, the West Polynesian, the Magellanian and the African gate. Many
of the genera occurring in Juan Fernandez, endemic or not, are mentioned by CROIZAT;
several of them are, also in my opinion, of Antarctic origin.

Oreobolus. Instead of allowing it to enter S. America through the ‘“Magellanian
gate’’ CROIZAT makes a circuit along his Pacific baseline from New Zealand to Fuegia.

¥uania. | have already discussed the taxonomic position of this genus which
CROIZAT derives from a Mascarene centre in company with related Andine genera.
HUTCHINSON. linked it with Cevoxy/on to form a tribe Ceroxyleae; CROIZAT remarks
that

this may be better taxonomy, we do not know, but does not change substances in the
least. The ingredients that make up the Morenieae, Iriarteae and Ceroxyleae are all from
the same kitchen. In one case /yophorbe speaks, in another so does the face of dispersal.
Everything in the end rhymes to the very same. With or without /yophorbe, seasoned
with this or that sauce, the stew does not change. . . classification of such a genus as
this is bound forever to remain the plaything of opinion (pp. 456-457),

a statement quite characteristic of this author’s method of approaching a phyto-
geographical problem.

Peperomia has its stronghold in tropical America where a number of subgenera
are found. To CROIZAT the genus is Pacific and Juan Fernandez one of the starting-
points of its dispersal toward the heart of S. America, the four island species leading
straight to north Chile. He based this opinion on YUNCKER who wants to bring
them to subgen. Sphaerocarpidium, where, however, only one of them belongs, while
the other three, as was brought forth above (p. 259), point west, one of them inti-
mately related to P. tristanensis, which, unexpectedly it is true, brings the Antarctic
within sight. In a footnote CROIZAT admits that I may be right in my opinion of
the taxonomic position of P. derteroana; none the less he finds the contact Juan
Fernandez—Tristan not at all peculiar. Aypetrum is another example, both “a
mere aspect of the standard dispersal between Africa and South America” (p. 102).
But this “standard dispersal’, as discussed by ENGLER, refers to a number of
tropical genera and families bearing witness of a transatlantic bridge. In CROIZAT’s
opinion the track Mascarenes—Africa-America was a very important route.

Palms and Peperomias are pantropical and range south to New Zealand and
central Chile, and there are many endemic palm genera on the Indian and Pacific
islands south of the Equator, whence follows that the field is open for speculation
and that the possibility of a primary centre in the south may be discussed without
breaking the rules of the game, but to derive exclusively neotropical families from
a Mascarene centre cannot fail to cause astonishment. CROIZAT, dealing with Cac-
taceae, Bromeliaceae, Rapateaceae etc.—and in an earlier chapter he asserts that
Malesherbiaceae also had their starting point in Africa—says that

to credit groups of the kind with “American’”’ origin steps must be taken as a prelim-
inary which no one trained in precise thinking will encourage. . . . AAzpsalis is doubtless
a Gondwanic genus because it is rooted within the triangle Ceylon—Madagascar-Masca-
renes. It well may have reached Brazil and the West Indies from West Africa... . the
baseline of the Cactaceae squarely rests between Patagonia and the Mascarenes, and if

DERIVATION OF THE FLORA AND FAUNA 277

Rhipsalis is Gondwanic it is likely, to say the least, that the archetypes of the family
itself stem from the A/roantarctic triangle (pp. 364, 365).

Much could be said to this, but I shall confine myself to remarking that the
claims of RAzpsalis outside America to true citizenship are still under suspicion and
that the existence of this specialized type of angiosperms in the Gondwana flora
needs proofs of some sort. But CROIZAT does not stop here. On p. 523 we read:

Considering that the New World is wzzformly at the receiving end of angiospermous
tracks throughout the first epoch of migration we are drawn to conclude that none of
the primary angiosperms ever originated in the Americas.

But who will tell us where the first angiosperms were evolved? We have to
go back to times when “the Americas’, as we know them, did not exist and where,
just as everywhere else, all centres of origin are secondary. I cannot see that his
opinion that the primary centre must be looked for in an “Afroantarctic triangle”
rests on a stable foundation, but everybody will agree when he formulates the
following recommendation to the “phytogeographers of the Academic school”:
“Look, and keep silent awhile ere you speak.”

Phrygilanthus and Santalum; compare what will be said below about Coprosma.

Dysopsts. Antarctic according to CROIZAT p. 51; I regarded it as neotropical.

“A large part of the Ericaceae originated with the Empetraceae in the Afro-
antarctic Triangle’, CROIZAT writes p. 381. That the Antarctic has been used as
a migration route of the Gaultherioideae seems certain, let alone where the family
or complex of families had its beginning. CRoIzAT finds that the occurrence of a
species of Pernettya in Juan Fernandez and of another in Galapagos is

one of the most interesting aspects of the Gaultherioideae. . .. a track of this nature is
nothing unusual, of course, because dispersals of this extent and nature are commonplace
between the vicinity of Juan Fernandez and Hawaii, not to speak of the Galapagos and
the Revilla Gigedos (pp. 167, 168).

We have examples of a floristic “‘contact’’ between Juan Fernandez and Hawaii,
but they are very few and to call this type of dispersal commonplace is an inadmis-
sible exaggeration of facts. The American area of Pervet/ya extends from Fuegia-
Falkland to Mexico, and its presence on Juan Fernandez and Galapagos, outlying
stations west of the continent, is not surprising; we need not construct a special east
Pacific track to explain them, nor assume that “the South Pacific is a fundamental
center of the Ericaceae’’.

I have mentioned Ampetrum and the boreal centre of Empetraceae, also that
CROIZAT wants them to have followed the same stream as Ericaceae: ‘‘they begin
their visible dispersal only westward from the Cape region” (p. 349), ‘with the
present Atlantic as the axis of their distribution” (p. 353). Empetrum used to pass
as a classical example of bipolarism, but to those who still believe that bipolarity
exists CROIZOT says: “bipolarism, as we know, is a bugaboo of academic phyto-
geography’’—I am.afraid that some of us didn’t know that.

The case of Cumznia (for which CROIZAT uses the fancy name Fohow1a), whether
or not to be linked with the Old World and Hawaiian Prasioideae, is embarrassing:

278 C. SKOTTSBERG

my discussion with F. B. H. BRowN (2377) made CROIZAT remark that “the debate
proved futile, as could be anticipated’. To attempt a solution and to discuss possible
routes of migration is called “obvious nonsense”.

Of the presumable history of Zxphrasza enough has been said above. In CROI-
ZAT’s opinion there cannot be the question of a north-south track from Asia across
Malaysia to Australia and New Zealand; his map fig. 6 shows the centre far south
in the Pacific, from where tracks lead east to S. America and west to New Zea-
land and Australia and thence from there to Eurasia and along the Aleutian arc to
N. America.

From the hypothetical ‘“‘main angiospermous center’ south of Madagascar,
where Plantago-Palaeopsyllium was born, one branch leads west around S. America
to Juan Fernandez, another east to Polynesia and from there to Hawaii, alterna-
tively reached directly from Juan Fernandez. I preferred the first alternative, but
even if the circumpolar distribution suggests an Antarctic origin I can see no good
reason to place the starting point south of Madagascar.

Coprosma is one of the standard Antarctic dispersal types but lacking in Africa
and in continental America. From its distribution CROIZAT draws the following
conclusions.

This dispersal shows a) An Antarctic range in the southern Pacific which fed Co-
prosma to all lands between Juan Fernandez and E. Java, b) This range could possibly
reach Hawaii from West along the line East Malaysia—Hawaii, c) The trend of this type
of dispersal is all westward from the Americas (Juan Fernandez to New Zealand, Aus-
tralia, New Guinea, Malaysia) ultimately veering again eastward to reach Hawaii (p. 101).

To make Juan Fernandez the source of Coprosma in New Zealand is utterly im-
possible and the statement is not in agreement with CROIZAT’s map, fig. 29, which
shows a baseline along lat. 60° between the longitudes of New Zealand and Fuegia
and arrows pointing west to New Zealand and northeast to Juan Fernandez.

I am not going to enter upon a discussion of the origin of a world-wide family
like Compositae; I shall only quote the following sentence: ‘‘The perfect circum-
polar distribution of Zaraxacum magellanicum ... in itself tells us where is to be
sought the ultimate origin of the Compositae’”’ (p. 63). Even if it is true that this
interesting dandelion also inhabits New Zealand, the conclusions would seem too
far-reaching.

I shall conclude this discussion with a summary of a recent paper by AXELROD
(75) who gives us a palaeontologist’s view on angiosperm evolution.

The cradle of angiosperms stood in the tropical belt, the width of which has
undergone considerable change during the geological ages. The angiosperms origi-
nated and evolved in uplands during Permian to Triassic times and gradually
descended to the lowlands to replace the vanishing floras now extinct or repre-
sented by such remnants as the Cycadophyta. Fossil records, especially of pollen,
make it probable that many families existed in early Jurassic, and from middle
Cretaceous, when angiosperms had gained dominance over gymnosperms and ferns,
the presence of numerous families belonging to very different orders bear witness
of a long history. By that time three different lowland floras had developed, the

DERIVATION OF THE FLORA AND FAUNA 279

Tropic-, Arcto-, and Antarcto-Cretaceous, and it is the latter that interests us here.
AXELROD emphasizes that numerous data allow us to conclude that many families
and genera generally regarded as having evolved in the temperate zones originated
in the tropics.

The data seem to support the view that angiosperms have not had an exclusively
holarctic source, or a wholly austral center of origin. They were being assembled in
both regions by the gradual adaptation of basic tropical groups to extratropical conditions
during the long period of Permo-Triassic down to the Cretaceous. On this basis the
Arcto-Cretaceous and Antarcto-Cretaceous floras of higher temperate latitude represent
vegetation types whose genera were derived largely by the long and continued differen-
tiation of successively derivative members of original tropical and border-tropical angio-
sperms. From this standpoint, the temperate regions to the north (holarctic) and south
(antarctic) are subordinate or secondary centers in early angiosperm evolution.

The conclusions we can draw from this are (1) that the origin and primary seat
of the angiosperms was in the tropics before anything like the present map existed;
(2) that this tropical flora gradually gave birth to a temperate flora also in the
south; (3) that Antarctica became the centre of a varied Antarcto-tertiary flora
which spread north and over S. America reached the region where now stands
Juan Fernandez.

II. Pteridophyta.

Owing to their great age and perhaps also to their greater faculty of dispersal
many of the genera and some species have attained a very extensive range making
it difficult or impossible to assign them to one of the elements proposed in the sub-
division of the angiosperms. Of 23 genera found in Juan Fernandez, 16 are very
wide-spread; some of them belong to the largest fern genera, and even if we avail
ourselves of the generic concept used by COPELAND (69), the situation remains
practically the same. To COPELAND 50-75 % of the living ferns are of austral or
Antarctic ancestry; we shall see to what extent this is true of the Juan Fernandez flora.

With regard to the Hymenophyllaceae COPELAND remarks (67. 174) that “no
other plant family of its size and diversity is quite so conspicuously Antarctic in
origin as this one’.

Trichomanes-Vandenboschia is pantropical; as 2 of the 3 insular species are
decidedly neotropical in their affinities, also 7. phz/7ppzanum is referred to the same
element.

Serpyllopsis is subantarctic-American and best attached to an Antarcto-tertiary
element.

Hymenoglossum should perhaps be referred to the same element, but its re-
lations are just as unknown as those of Serpyllopsis.

Hymenophyllum-Mecodium is a pantropical group, also represented in New Zea-
land; cuneatum and caudiculatum are most likely of neotropical origin and this may
be true also of fuctforme. H. pectinatum stands, COPELAND says, apart from the other
species, but I can find no good reason to call in Antarctic, while the distribution
of Sphaerocionium ferrugineum and its near relatives testifies to their Antarctic origin
in spite of their affinity to ci/zatum (comp. above p. 219). Hymenophyllum s. str.

280 C. SKOTTSBERG

as limited by COPELAND 1938 (66) does not include fa/klandicum (Mecodium accord-
ing to COPELAND p. 94), though it is closely akin to peltatum, and H. rugosum
“Gs to H. tunbridgense what H. falklandicum is to H, peltatum’’ (62. 13). According
to CHRIST (59.146), peltatum and tunbridgense are essentially Antarctic-andine but
found north as far as Mexico, also reported from S. Africa and New Zealand, ex-
tending in insular climates to western Europe. The conclusion would be that also
falklandicum and rugosum should be brought to the Antarctic group.

Hymenophyllum-Meringtum secundum, plicatum and tortuosum are subantarctic-
American in their present distribution; J/er7zg7um is austral and extends north to
tropical Asia, and its origin most likely Antarctic.

Thyrsopteris. Mesozoic fossils supposed to come near this genus have been re-
ported from the boreal zone; besides, from the Tertiary in Chile (see below p. 399).
HILL (734.1477) calls it “a remnant of a once widespread group which migrated
southwards, possibly in Cretaceous times, from the Northern Hemisphere’. With
regard to its present unique station COPELAND (67.175) writes:

As Juan Fernandez can hardly have evolved so peculiar a fern, its most reasonable
origin, as a Juan Fernandez fern, is Antarctica, whether directly or through southern
Chile. European fossils have been referred to this genus, but Juan Fernandez was not
colonized from Europe.

Lophosoria is exclusively neotropical at present, but if we are to believe, as
COPELAND thinks, that the Cyatheaceae derive from Antarctica, it is unlikely that
Lophosoria would be an exception.

The Antarctic origin of Dzcksonza seems very clear to judge from its actual
distribution pattern. To what was said above p. 222 I shall add that along the
track New Zealand—Australia-New Caledonia—Fiji(-Samoa)—Malaysia 14 species are
found, along the Andean path 9, and that the species of Juan Fernandez are not
linked with the neotropical but with southwest Pacific species.

The present distribution of the genus Cystopferzs suggests a boreal origin.

In its wide as well as in its more restricted sense Dryofterzs is world-wide,
and COPELAND’s remark (67.181) that he has little doubt that the group as a
whole is of southern origin (see also 69.122) is perhaps little more than a guess.
I think we can take it for granted that hundreds of species have evolved in the
neotropical region, where the single Juan Fernandez species D. 7xaegualifolia has
its relatives.

Polystichum. Eastern Asia is the centre of greatest concentration; the section
occurring there is, in COPELAND’s opinion, the least primitive, and we have a number
of southern bi- or tricentric forms, among them P. vestédum coll. The endemic P.
berterianum is very close to the circumpolar adzantiforme. To quote COPELAND: “The
case for an Antarctic origin of Polystichum is so clear that the evidence has long
been familiar even to those hesitant to draw the obvious conclusion” (67. 181);
and, “The distribution .. . testifies clearly to its Antarctic origin’ (69. 109).

Arthropieris is the only fern genus found in Juan Fernandez which is absent
from continental America, but otherwise typically austral. Its Antarctic origin is

DERIVATION OF THE FLORA AND FAUNA 281

beyond doubt, and 4. altescandens has its nearest allies in Oceania; comp. above
preee.

Asplenium. COPELAND points out that some groups at least are southern and
that perhaps the same applies to the whole genus: “it migrated in its present
form from Antarctica” (69. 167). Of the species found in Juan Fernandez, ob/iquum
is austral-bicentric, ste//atum close to neotropical species, macrosorum most likely
of neotropical affinity; daveoides comes very near alvarezsense from Tristan da
Cunha and is not, as has been suggested, related to the boreal rufa muraria.

Blechnum. According to COPELAND (69.157) no fern genus is more con-
spicuously austral in its present distribution and thus more evidently Antarctic
in origin. Of the 6 species found in Juan Fernandez auriculatum, chilense and cy-
cadifolium belong to an Afro-american group, Schoft7z and perhaps also valdiviense
of austral-circumpolar type, and /ovgzcauda intimately related to the neotropical
Sprucez; even so it cannot blur the picture of an old Antarctic genus.

Pellaea is an austral-tricentric genus extending north to Canada, but P. chz-
/ensis, endemic in Juan Fernandez, but dangerously close to a widespread American
species, belongs to a group that, according to COPELAND (69. 70), is best allowed
to form a separate genus; I have brought it to the neotropical element.

Hypolepis is pantropical, especially neotropical, but numerous species are
scattered in the equatorial and austral zones: Africa with neighbouring islands,
Malaysia and north to Japan, Australia-New Zealand—Lord Howe—Melanesia—Hawaii
—when COPELAND said “without surviving Chilean representatives” he forgot 7ugo-
sula. This species is bicentric or, including Polypodium villosoviscrdum of Tristan,
tricentric; there is another variety on St. Helena. Thus when COPELAND 67. 177
calls Hypolepis ‘‘an old antarctic genus now at home chiefly in the Tropics but
with two of its paths still occupied’, we can add that also the third path is indicated.

Adiantum is most numerous in S. America, and A. chi/exse and related species
tell us nothing of the earlier history of the genus; COPELAND 69.78: “I suppose
that at least the most of the extant species are of Antarctic ancestry, but the
genus is old and may have lived elsewhere without interruption”; in 67.178 he
, but that the genus

’

says that “Antarctica has played a major part in its history’
“may include elements surviving from other floras and from an age prior to the
great dispersal from the south’. I refer A. chz/ense to the neotropical group.

Pteris. The great wealth of species is in the tropics and only few extend as
far south as to south Chile, S. Africa and New Zealand. P. chilens7s is of neo-
tropical character, perhaps also P. semzadnata, while derteroana belongs to a group
regarded by COPELAND as Antarctic to judge from its distribution pattern (see
above p. 223).

Histiopteris incisa is a pantropical and circum-austral, polymorphous fern with
a number of “local derived species” (COPELAND) in Indonesia and Polynesia. The
genus is, COPELAND says (69.60), “evidently old enough to be a migrant from
Antarctica”.

Polypodium is an aggregation of unities of different origin and history. Gram-
mitis is, in COPELAND’s opinion (67.184), “a plain Antarctic case’, and G. z/-
lardierit “surrounds Antarctica more closely and completely than does any other

282 Cc. SKOTTSBERG

fern’ (69. 211; see above p. 223); it was at that time supposed to be circum-
polar, but is replaced in S. America by P. magellanicum. As mentioned above,
I believe that if a genus Syxzammza with Polypodium Feuillez as the typical species
is accepted, the endemic P. zx¢ermedium finds its place with this, but I fail to see
why COPELAND (68) regards S. Feuillei as ‘clearly Antarctic’, whereas the nearly
related Gontophlebium is called ‘‘a northern genus’. Of the remaining species
P. Masafuerae and P. (Xiphopteris) “#7chomanoides are neotropical and P. (Pleo-
peltis) /axceolatum pantropical but not reported from Australia and not indicating
an Antarctic origin.

We have no good reason to look for an Antarctic ancestry of Elaphoglossum,
though COPELAND thinks that it “may have come from the south” (67. 185), and
i. Lindenii is a neotropical species.

The species of Gletchenia occurring in Juan Fernandez belong to Sécherus,
a genus segregated by some modern authors; II species are scattered over the
austral zone, and in COPELAND’s opinion Gleicheniaceae are ‘‘obviously and en-
tirely of Antarctic ancestry” (69.26), or “entirely Antarctic at some stage of its
history” (67.173); how this should be explained I cannot tell unless he means
that the family characters first evolved in the Antarctic, secondary centres of
evolution having become established in the tropics. The genetic relations between
the species inhabiting the three sectors (see above p. 224) should be studied.
Meanwhile I shall refer the 5 species occurring in southern S. America to the
Antarcto-tertiary element.

The Ophioglossaceae are an ancient family, “scattered with remarkable uni-
formity over the habitable globe’ (COPELAND 69.12). In 67.167 he paid special
attention to Botrychium australe R. Br. which has the peculiar distribution Argentina,
Australia, Tasmania and New Zealand and “may be regarded as an obvious immi-
grant from the south’. It is surprising that the species found in Patagonia and
Fuegia is not this, but the northern 2. duzarza L. We have no reason to regard
Ophioglossum fernandezianum as coming from the south.

Lycopodium was not discussed by COPELAND. L. magellanicum is subantarctic-
circumpolar, and part of the history of the genus may have been enacted in Ant-
arctica, the more so as L. scarzosum, which belongs to another section, is bicentric.

With reference to the discussion above the Pteridophytes are arranged as
follows.
I. Antarcto-tertiary element.—32 sp. (60.4 %).

1. Distribution pattern austral-circumpolar, bicentric or tricentric.—20 sp.
a. Endemic species (2): Hymenophyllum rugosum, Polystichum berterianum.
6. Species also found in S. America (Chile), many of them with a wider
distribution (w) in the S. hemisphere (18): Hymenophyllum secundum, plicatum,
tortuosum, ferrugineum and falklandicum (w), Polystichum vestitum (w), Asplenium
obliquum (w) and dareoides, Blechnum valdiviense and chilense, Hypolepis rugosula
(w), Histiopteris incisa (w), Polypodium magellanicum (w), Gleichenia pedalis,
quadripartita and cf. litoralis, Lycopodium magellanicum (w) and scariosum (w).
2. Species belonging to genera endemic in S. America but supposed to be

DERIVATION OF THE FLORA AND FAUNA 283

of Antarctic origin: Serpyllopsis caespitosa, Hymenoglossum cruentum(?), Lopho-
soria quadripinnata.—3 sp.

3. Endemic species without near relatives in America, suggesting either an
ancient southern Pacific west-east path or an Antarctic-Magellanian track now
not occupied: Dicksonia berteroana and externa, Arthropteris altescandens, Blech-
num Schottii, Pteris berteroana.—5 sp.

A. Species closely related -to S. African ones:—2’ sp.

a. Endemic: Blechnum cycadifolium.

6. Also on the mainland: Blechnum auriculatum.

5. Endemic species very nearly related to a neotropical species: Blechnum
longicauda.—1I sp.

6. Endemic genus without affinities to living genera: Thyrsopteris elegans.
Sas:

II. Neotropical-Andean element.—20 sp. (37.7 %).

a. Endemic species (8): Trichomanes Ingae and philippianum, Dryopteris
inaequalifolia, Asplenium stellatum and macrosorum, Pellaea chilensis, Polypodium
intermedium, Ophioglossum fernandezianum.

6. Also on the mainland, restricted to Chile or more wide-ranging (10): Tricho-
manes exsectum, Hymenophyllum cuneatum, caudiculatum, fuciforme and _ pecti-
natum, Adiantum chilense, Pteris chilensis and semiadnata, Polypodium Masafuerae
and lanceolatum.

c. Non-endemic neotropical species not found in Chile (2): Polypodium tricho-
manoides, Elaphoglossum Lindenii.

Ill. Arcto-tertiary element.—r sp. (1.9 %).

Also in Chile: Cystopteris fragilis var.

The Antarctic element is considerably larger than in the angiosperms, 60.4 %
against 42.2, the neotropical being of the same size, 37.7 and 36.7 %. On the other
hand, the boreal element is very insignificant (even open to doubt). No living
pteridologist has a wider general knowledge of the ferns than E. B. COPELAND, and
even if his new system does not appeal to everybody, we are bound to pay attention
to his theories. In several of his writing he points, as we have seen, to Antarctica
as the main source. On the other hand he admits that

ferns existed well over the world longer ago than I have would try to explain their
presence as immigrants from Antarctica, and must be supposed to have maintained
continuous existence elsewhere. This being so, the preponderance of ferns of apparent
Antarctic origin in the world to-day is surprising indeed (67. 188).

This origin is not readily revealed in the large and world-wide families and
genera; the best proofs are furnished by small families restricted to high southern
latitudes and bi- or tricentric in distribution—most of these families have been
created as a result of the modern splitting process. With a single southern genus
in a large tropical family we cannot feel on safe ground, and if in a large tropi-
cal genus only a solitary species is southern, we are not inclined to classify

284 C. SKOTTSBERG

the genus as Antarctic in origin. But, COPELAND remarks, if the ferns as a group
were of tropical origin they would be expected to be far more abundant north-
ward, where the land areas increase, than southward where the land decreases
—and very much so—in size, and this would be still more so if they were of
northern origin. The conclusion is that if a family or genus is mainly southern
to-day, this fact is a strong indication of its Antarctic origin and that, if the pre-
sent range is wholly southern, the evidence becomes almost conclusive. If the
more primitive families or genera are found to be characteristically southern, a
southern seat of old fern evolution is almost demonstrated (68. 626). But when it
comes to the large families, their history may be much more difficult to read:

A large family, even if of southern origin, cannot possibly be predominantly southern
in present distribution, because tropical and northern species must outnumber the whole
flora of Antarctic America, or even of New Zealand (67. 158).

With this he wants so say, I suppose, that a vigorous family has invaded the
tropics and also extended north under rapid evolution of genera and species con-
cealing the primary origin; he believes this evolution to have been so rapid that
the period since the Miocene has been sufficient to create most of the existing
species and a large part of the genera and for their spread over any expanse ot
suitable land area (l.c.).

We have no evidence that the Tropics as a whole were at any past time unfit for ferns;
and the assumption that this has been the chief place of evolution obviates the neces-
sity of assuming and explaining migration in latitude. It is only when we open our

eyes to anomalies in present distribution that appeal to other places of evolution be-
comes necessary (67. 163).

Great geographical disjunctions, of which the Juan Fernandez fern flora offers
many examples, are such anomalies, and they seem to show that Antarctica has
taken a very important part in fern history.

III. Musci.

In his analysis of the Magellanian moss flora CARDOT (48) distinguished an
Antarctic element and he believed in a common origin of the floras of Magella-
nia and New Zealand (p. 44). HERZOG (z2g) found that Juan Fernandez has “eine
fast vollstandig austral-antarktische Moosvegetation’’, but also possesses ‘‘ein paar
tropisch anmutende Arten: 7hysanomitrium Richardt, Porothamnium fasciculatum,
Pinnatella macrosticta und ein Rhacopilum.’ The monotypical Chilean genus Lam-
prophyllum was also mentioned as a genus of tropical ancestry. A circumpolar
austral-antarctic element is said to dominate in the moss flora of Patagonia and
Fuegia; characteristic genera, also represented in Juan Fernandez, are Dicranoloma,
Ulota, Dendrocryphaea, Lepyrodon, Ptychomnium, Weymouthia, Distichophyllum,
Pterygophyllum, Eriopus, Hypopterygium, Sciaromium, perhaps also Catagontopsis
and Pszlopilum. IRMSCHER (r43) gives many examples of austral-bicentric taxa:
Lepyrodontaceae, Polytrichadelphus, Weymouthia, Pterygophyllum, Sciaromium sect.
Aloma, Hypopterygium sect. Stenobasis, Hypnodendron, and species of Déstecho-
phyllum, Tortula, Macromitrium and Mielichhoferia.

DERIVATION OF THE FLORA AND FAUNA 285

The synopsis pp. 226—233 led to a classification according to the actual geo-
graphic distribution. We have to look for austral-antarctic species and genera
in the Andine-Chilean, Subantarctic-Magellanian and Pacific groups; thereby I
have tried to follow the same principles as applied before. Many of the endemic
species are difficult to place because their author did not compare them with other
species or discuss their position; 11, belonging to large and widespread genera were
left out (see list p. 235). The 8 species of 7hamunium, all endemic, were tentatively
included in group IV. In Natirl. Pflanzenfam. 2d ed. BROTHERUS enumerates 34
species, but none at all are quoted for S. America. Sect. I is entirely austral,
with a single species in tropical Asia (Sumatra, Java, Borneo, Philippines); 2 are
found in Australia and New Zealand, 1 in New Caledonia and 8 in Juan Fernandez.
Of sect. II, 3 species are southern (New Hebrides, E. Australia-New Zealand, New
Zealand-Java-Sumatra), the remainder scattered over the boreal zone, 2 iP
America, 3 mediterrmacaronesian, I England, 1 W. Europe to Japan, 2 Cauca-
sus, 9 E. Asia.

The total number of species included in the following synopsis is 120.

I. Antarcto-tertiary element.—67 sp. (55.8 %).

1. Bicentric or tricentric (t) distribution: Ditrichum affine, Amphidium cyathi-
carpum, Dicranoloma Billardieri (t) and Menziesii, Campylopus introflexus (t),
Thysanomitrium leptodus, Fissidens rigidulus and asplenioides (t), Rkacomitrium
symphyodontum (t), Zygodon intermedius and Menziesii, Rhizogonium Novae Hollan-
diae and mnioides, Bartramia patens (t), Philonotis scabrifolia (t), Rhacocarpus Hum-
boldtii (t), Weymouthia mollis, Leptodon Smithii (t), Pterygophyllum obscurum
and denticulatum, Lopidium concinnum, Rhynchostegium tenuifolium, Polytricha-
delphus magellanicus, Dendroligotrichum dendroides.—24 sp.

2. Endemic or also found in Chile and restricted to the S. American sector,
in either case with Australian-Neozelandic relations, with the exception of Eusti-
chia.—30 sp.

a. Endemic species (9): Dicranoloma fernandezianum, capillifolioides and nigri-
caule, Ptychomnium falcatulum, Distichophyllum fernandezianum, Pterygophyllum
tenuinerve, Eriopus leptoloma and grandiretis, Thuidium Masafuerae.

6. Also in Chile (21): Ditrichum longisetum, Dicranoloma capillifolium, Eu-
camptodon perichaetialis, Rhacomitrium subnigritum and striatipilum, Macromitrium
hymenostomum, Leptostomum Menziesii, Eustichia Poeppigii (other species in
America and Africa), Dendrocryphaea cuspidata, Lepyrodon parvulus, tomentosus
and implexus, Ptychomnium subaciculare and ptychocarpum, Pterygophyllum ano-
malum, Hypopterygium Thouini, Thuidium Valdiviae, Sciaromium pachyloma,
Catagoniopsis berteroana, Hypnodendron microstictum, Psilopilum antarcticum.

3. Endemic species of presumably Australasian and Polynesian affinity, not
or more distantly related to American species: Macromitrium fernandezianum and
Masafuerae, Cyptodon crassinervis, Thamnium rigidum, latinerve, Caroli, Ingae,
crassinervium, proboscideum, assimile and confertum, Distichophyllum subelimba-
tum and assimile.—13 sp.

286 Cc. SKOTTSBERG

II. Neotropical element.—26 sp. (21.7 %).

a. Endemic species (11): Campylopus subareodictyon, Leptodontium fernan-
dezianum, Didymodon calymperidictyon and linearis, Bryum fernandezianum, Pin-
natella macrosticta, Isopterygium fernandezianum, Rhaphidostegium Masafuerae
and caespitosoides, Rigodium robustum and Looserl.

6. Also in S. America (15); Gymnostomum calcareum (widely dispersed), Cam-
pylopus areodictyon, Thysanomitrium Richardi, Mielichhoferia longiseta, Anacolia
subsessilis, Porothamnium fasciculatum and arbusculans, Lamprophyllum splendi-
dissimum, Rhacopilum fernandezianum, Stereodon Lechleri, Rhaphidostegium
caespitosum, Rigodium toxarium, arborescens, hylocomioides and tamarix.

Ill. Chilean element.— 24 sp. (20.0 %).

a. Endemic species (4): Fissidens pycnotylus, Ulota fernandeziana, Philonotis
elabrata, Rhaphidostegium aberrans.

6. Also in Chile, mostly southern (17): Pleuridium Robinsonii, Hymenosto-
mum kunzeanum, Oncophorus fuegianus, Campylopus truncatus, Fissidens lepto-
chaete and maschalanthus, Tortula scabrinervis and flagellaris, Rhacomitrium lori-
forme and convolutum, Stenomitrium pentastichum, Bryum Lechleri, Bartramia
aristata, Philonotis krauseana and vagans, Rhynchostegium complanum, Oligotrichum
canaliculatum.

c. Cosmopolitan (3): Ceratodon purpureus, Rhacomitrium lanuginosum, Funa-
ria hygrometrica.

Note-—On p. 234 Campylopus introflexus was placed here; it was transferred to
I: 1 on account of its circumpolar distribution in the far south, from where it may have
invaded the tropics and migrated north, but it is perhaps just as probable that it is
an old pantropical species which has spread both north and south.

IV. Atlantic-Mediterranean element.— 3 sp. (2.5 %).

Comp. above, p. 235.

a. Endemic: Fissidens crassicuspes.
6. Not endemic: Campylopus polytrichoides, Trichostomum brachydontium.

We have seen above that HERZOG regarded the moss flora of Juan Fernan-
dez as almost entirely austral-antarctic and that the tropical element was small.
Most likely he added the species forming my Chilean element, because they are
concentrated in the moist southern part of the country and in several cases range
south into the subantarctic zone which is, however, no proof of their Antarctic
ancestry. A species like Porothamnium arbusculans is, as far as I know, restricted
to Chile and Patagonia, but the genus is essentially tropical, Szereodon Lechlert
extends from S. Chile to W. Patagonia, but is the only species reported south
of the Equator; Rigodium is American except 2 species found in Africa and
essentially tropical. On the other hand it is difficult to draw a limit between
groups II and III, but the species with a southern area are so numerous that I
felt obliged to make this distinction. The Atlantic element will perhaps disappear
when the distribution becomes better known.

DERIVATION OF THE FLORA AND FAUNA 287

The bi- or tricentric species found in Juan Fernandez represent a minor frac-
tion only of the mosses common to S. America and New Zealand. SAINSBURY’s
new Flora (270) contains nearly 60 species with this type of distribution, and to
these some occurring in Australia and Tasmania, but not in New Zealand, may
be added. Detrichum affine, Trichostomum brachydontinm, Rhacomitrium symphyo-
dontum, Bartramia patens, and Pterygophyllum denticulatum, \isted for New Zealand
by BROTHERUS (34), are not recorded by SAINSBURY.

Consequently, what Miss FULFORD (see below p. 289) says about the Hepati-
cae is true also of the mosses.

IV. Hepaticae.

In an interesting paper on the distribution of the Hepaticae DOMIN (76) lays
stress upon that they are just as specialized geographically as the flowering
plants, and
einen uralten, heutzutage in den gemissigten und kialteren Gebieten sozusagen erstarr-
ten, sich von ihren ganz speziellen Standorten nicht weiter ausbreitenden, wenig an-
passungsfaihigen Typus darstellen, welcher auf der nérdlichen Hemisphire seinen Ent-

wickelungsgang in weit zuriickliegenden Epochen durchgemacht und bereits zur Tertiar-
zeit in den Hauptziigen beendet hat (p. 3).

He thinks that the evolution still continues in the tropics. Numerous genera are
widespread, but the majority of genera and species inhabit the tropical and south
temperate rain forests in America, Africa (perhaps not so rich?), Malaysia, Austral-
asia and Oceania. Austral-antarctic genera mentioned by DOMIN are Schzstochi/a,
Balantiopsis, Adelanthus (hardly austral), Lophocolea, Chiloscyphus, Trichocolea, and
many genera with their greatest wealth of species in the tropics are well repre-
sented in the temperate and cool south, such as Azccardia, Symphyogyna, Pla-
giochila, Madotheca, Lepidozsia, Radula, Frullania, while few, in cases very few,
species are found in the boreal zone. From HERZOG's handbook the following
genera may also be quoted as tropical-austral, or essentially or exclusively south-
ern: Hymenophytum, Acrobolbus, Tylunanthus, Famesoniella, Saccogyna, Lepicolea
and Marsupidium, to mention only genera also found in Juan Fernandez. Lefz-
dolaena is restricted to the south temperate and cold zones. In the following ar-
rangement I have been guided by these authors.

I. Antarcto-tertiary element.—84 sp. (67.7 %).

1. Bicentric or tricentric (t) species: Marchantia berteroana (t) and foliosa,
Riccardia insularis, Metzgeria decipiens and violacea (t), Hymenophytum flabellatum,
Symphyogyna hymenophyllum, Monoclea Forsteri, Pallavicinia xiphoides, Jame-
soniella colorata (t) and grandiflora (t), Acrobolbus excisus, Mylia repens, Lophocolea
pallidevirens and muricata (t), Marsupidium piliferum, Bazzania cerina, Lepidozia
sejuncta (t) and plumulosa, Lepicolea ochroleuca (t), Lepidolaena magellanica,
Schistochila splachnophylla, Frullania magellanica (t).—23 sp.

2. Restricted to the American sector, endemic in Juan Fernandez or also
found on the mainland, connected with species in the Australia-New Zealand sec-
tor S61 Sp:

288 Cc. SKOTTSBERG

a. Endemic (17): Riccardia adglutinata and leptostachya, Metzgeria multifor-
mis, Tylunanthus silvaticus, bilobatus and densiretis, Lophocolea papulosa, angulata
and submuricata, Trichocolea opposita, Schistochila Skottsbergii, Balantiopsis hians
and lancifolia, Lopholejeunea spinosa, Strepsilejeunea squarrosula and macroloba,
Cololejeunea Skottsbergii.

6. Also found in Chile (44): Riccardia fuegiensis, breviramosa, variabilis and
nudimitra, Metzgeria decrescens, Symphyogyna circinata and Hochstetteri, Jame-
soniella maluina and oenops, Tylunanthus limbatus, Mylia fuscovirens and ligulata,
Lophocolea rotundifolia, fernandeziensis, chilensis, attenuata, textilis and divergenti-
ciliata, Chiloscyphus integrifolius and lobatus, Saccogyna squarristipula, Adelanthus
sphalerus, Lepidozia bicuspidata, pseudozoopsis, fernandeziensis and Jacquemontil,
Trichocolea verticillata, Schistochila berteroana and pachyla, Balantiopsis cancellata,
chilensis and purpurata, Radula hastata, microloba, Mittenii and Dusenii, Madotheca
chilensis and subsquarrosa, Frullania Eckloni, chilensis, lobulata and stipatiloba,
Brachiolejeunea spruceana, Strepsilejeunea acuminata.

II. Neotropical element.—1 3 sp. (10.5 %).

a. Endemic (3): Fossombronia fernandeziensis, Anthoceros Skottsbergii, Le-
jeunea reticulata. :

6. Also in tropical South America, extending to Chile, or southern, but pre-
sumably of tropical origin (10): Megaceros fuegiensis, Androcryphaea confluens,
Anastrophyllum leucocephalum, Bazzania peruviana, Harpalejeunea oxyota and
setifera, Siphonolejeunea nudicalycina, Aphanolejeunea asperrima and diaphana,
Colura bulbosa. —

Ill. Chilean element.— 27 sp. (21.8 %).

Chilean species without tropical connection, nor suggesting Antarctic rela-
tions.

a. Endemic (5): Solenostoma obtusiflorum and rostratum, Plagiochila fusco-
brunnea, Lepidozia fragillima and disticha.

6. Also in Chile, especially in the south (18): Solenostoma crassulum, Ana-
strepta bifida, Plagiochila gayana, fasciata, hyadesiana, deformifolia, chiloensis,
rectangulata, remotidens, pudetensis, homomalla, neesiana, riparia, squarrosa,
robusta, elata and Notarisii, Herberta runcinata.

c. Cosmopolitan (4): Plagiochasma rupestre, Reboulia hemisphaerica, Lunularia
cruciata, Marchantia polymorpha.

Even if not the Juan Fernandez Hepatics are almost entirely Antarctic as
HERZOG thinks, the Antarcto-tertiary element is proportionally larger than in any
other group, and it is not impossible that genera like Wegaceros or Harpalejeunea,
the island species of which were referred to the neotropical element, are Antarctic,
and that this is true also of Bazzania, represented in I and I, but on the other
hand we must admit that pantropical genera like Rzccardia, Anthoceros, Mega-
ceros, Bazzania, Lepidozia, Radula, Madotheca, Frullania etc. may have extended
far south during a warmer period and given rise to important groups all around
the circumpolar belt, and that we have to show that such disjunct southern groups

DERIVATION OF THE FLORA AND FAUNA 289

are interrelated in such a manner that we can derive the genera from a common
Antarctic centre. This is a problem that only an expert and all-round hepatologist
can solve. In our island case the result may be that the Antarctic element will
shrink and the Neotropical swell; we may have to merge the Chilean element in
the Neotropical. Disregarding the cosmopolitan species, anthropophilous and anthro-
pochorous and very likely of recent introduction, we find that the bulk of the
Chilean species is formed by the 14 species of Plagiochila; we need a critical
revision of this gigantic assemblage before we shall know anything of its origin
and history. Also small but widely scattered genera offer difficulties: Axastrepia,
Adelanthus and Herberta, for instance.

The discontinuous distribution of many southern genera and several species
was commented upon by Miss FULFORD (703. 846):

The distribution patterns indicate that they were contemporaries at a time when free
migration was possible between South America, Africa, Australia, New Zealand and the
Sikkim area in N.E. India. They also indicate that the Antarctic Continent has been of
great importance in the distribution of genera and identical species in South America, the
Antarctic Islands [vead Subantarctic], Australia-New Zealand and probably Africa.

Finally, let us repeat our comparison of the four phyla analyzed. No obviously
Arcto-tertiary (Boreal) species were distinguished among the Hepaticae. The Chilean
element accepted in the mosses and hepatics has been combined with the Neotropical
to form a Neotropical-Chilean group (NC); A, Antarctic; B, Boreal.

Table V.
A NC B
ANOSIOSOETONS Sy GG i 60a 8 Bela 4252 30.7 15.0
PASTA OINNAWES 6 S46 0 a 6 6 oo WOz BO 1.9
IMOSSESENeechas cts ss Th 3s | SEO 41.7 2.5
LENGIORNENES 5" Ei, Veil co. LOM ecient OT) 32.3 —-

V. Lichenes.

Unlike the Archegoniates the Lichens offer insuperable difficulties when we
try to trace eventual centres of origin and evolution. Of 67 genera represented in
Juan Fernandez no less than some 55 have a wide, in cases world-wide distribu-
tion; they are called cosmopolitan, subcosmopolitan, tropical, temperate and so
forth, and with regard to about 70 species the situation is the same. Only ina
very limited number of instances a genus, or a group of related species is, to
judge from its present distribution, concentrated to a limited area, tempting us
to consider it a centre of evolution. On the other hand we must not forget that
lichens are lichenized fungi; modern lichenologists do not regard them as a spe-
cial phylum but range them with the different fungus orders and families accord-
ing to the taxonomic position of the fungus component. Nothing prevents us
from assuming that the same lichen species originated in different places widely
apart where both fungus and microscopic alga happened to be present at the time
when lichen-forming fungi existed as independent organisms. Nor should it be

19 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

290 Cc. SKOTTSBERG

forgotten that numerous green and bluegreen algae are cosmopolitan and that the
fungus spores as well as the thallogenous diaspore (in fact more important for
dispersal) of lichens are quite resistant and adapted to long-distance transporta-
tion, also that they have a greater facility to get established where they happen
to land than the spores of ferns and bryophytes.

During early Tertiary times—and certainly much earlier—South America had
a rich and varied lichen flora of tropical and subtropical character. With the grad-
ual climatic differentiation and the evolution of cold-resistant forms the rising
Andes provided stations for both northern and southern lichens; migrations in
both directions helped to equalize the Boreal-arctic and Austral-antarctic floras and
furthered the origin of a bipolar element.

With our present insufficient knowledge of the distribution of the lichens all
we can say with regard to a majority of species or genera is that they are tropi-
cal-subtropical or temperate and either northern or southern; it is rarely pos-
sible to decide in favour of the north or the south as a primary source of the
Andean temperate flora. The exceptions from this rule are few; for instance, gen-
era like Pachyphiale, Lemmopsis and Massalongia may be classified as northern,
while Byssocaulon, Pseudocyphellaria, Nephroma and Stereocaulon are southern or,
if bipolar, have a southern centre making it permissible to conclude that the
Antarctic continent has played an important part in their history and that the
bicentric distribution of so many species, also in the larger genera, testifies to an
Antarctic origin or at least indicates that a transantarctic migration route once
existed. Thus I have ventured to distinguish an Antarcto-tertiary element also in
the Chilean-Juan Fernandez lichen flora. Here as in other groups the possibility
that a bicentric species may have reached its stations from the north must be
taken into account; that a circumglobal species has reached southern Chile and
New Zealand is no proof of a transantarctic connection.

Below an attempt is made to distribute the Juan Fernandez lichens among
geographical-genetic groups. Only 164 species could be included. No place at all
could be assigned to many of the endemic species—see above p. 254—and the fol-
lowing non-endemic ones with their disjunct areas were also excluded: Baczdia
delapsans, Usnea dasypogoides and subtorulosa, Caloplaca rubina, Buellia halophiloides
and fernandesiana.

The arrangement proposed is far from satisfactory and will, I am sure, be
subjected to criticism. With our very imperfect knowledge of the real distribution
of lichens called “wide-spread” or even “‘cosmopolitan’’, mistakes in assigning a
species to a certain element are unavoidable. Thus, many were referred to group
I with great hesitation because it is impossible to decide if they have reached their
austral-bicentric stations (south Chile, New Zealand) independently from the north,
or if Antarctic routes are involved. The species lumped under IV surely represent
several different distribution patterns.

DERIVATION OF THE FLORA AND FAUNA 291

I. Antarcto-tertiary element.—58 sp. (35.4 %).

b, Austral-bicentric; t, Austral-tricentric species.

a. Endemic species (5): Psoroma vulcanicum, cephalodinum, dasycladum and
angustisectum, Pseudocyphellaria berteroana.

6. Non-endemic (53): Sphaerophorus melanocarpus, Phaeographina scalpturata
(b), Byssocaulon niveum (b), Leptogium phyllocarpum (b), Parmeliella nigrocincta (t)
and pycnophora, Pannaria fuegiensis and rubiginosa(b), Massalongia carnosa, Psoroma
pholidotum and sphinctrinum (b), Lobaria crenulata (b), Pseudocyphellaria argyracea
(t), intricata (b), fragillima (b), subvariabilis (b), chloroleuca (b), cinnamomea (b), hir-
suta, Guillemini, gilva (t), mougeotiana, aurata (t), nitida, endochrysea (b), Durvillei
(b), flavicans (b), Freycinetii (b) and Richardi (b), Sticta Weigelii (b), lineariloba,
latifrons (b) and laciniata, Nephroma plumbeum, cellulosum (b), antarcticum (b) and
australe (b), Catillaria melastegia (b), Megalospora versicolor (b), Phyllopsora parvifolia
(b), Cladonia pycnoclada, didyma (b) and aggregata (t), Stereocaulon patagonicum,
ramulosum (b) and implexum (b), Coccotrema granulatum (b), Lecanora albellina,
Placopsis chilena, fuscidula (b) and parellina (b), Myxodictyon chrysostictum (b),
Buellia halophila (b).

II. Andean tropical to temperate element.— 54 sp. (32.9 %).

a. Endemic species (9g): Arthonia subnebulosa and berberina, Enterostigma
Skottsbergii, Coenogonium velutinum, Lecidea avium, Pertusaria hadrocarpa and
Skottsbergil, Lecanora masafuerensis, Caloplaca orthoclada.

6. Non-endemic (45): Arthopyrenia cinchonae, adnexa and planorbis, Pyrenula
aspistea, mammillana and Kunthii, Pyrenastrum chilense, Arthonia complanata,
Graphis intricata and Dumastii, Dirina limitata, Schismatomma accedens, Thelotrema
lepadinum, Dimerella lutea, Physma chilense, Leptogium moluccanum, tremelloides,
cyanescens, Menziesii and callithamnion, Lecidea leucoplaca and icterica, Catillaria
endochroma and leucochlora, Bacidia endoleuca and subluteola, Toninia bullata,
Lopadium leucoxanthum, Baeomyces chilensis, Acarospora xanthophana, Per-
tusaria polycarpa, Melanaria melanospora, Parmelia laevigatula, abstrusa, nilgher-
rensis, soredica and microsticta, Menegazzia sanguinascens, Usnea angulata, Bom-
byliospora dolichospora, Caloplaca subcerina, Theloschistes flavicans, Physcia picta,
Anaptychia pectinata, Cora pavonia.

III. Boreal element.—14 sp. (8.5 %).

a. Endemic: Lemmopsis polychidioides.

6. Non-endemic (13): Verrucaria microspora, Arthonia cytisi, Gyalecta jenensis,
Pachyphiale cornea, Racodium rupestre, Lecidea enteroleuca and viridans, Bacidia
arceutina, Acarospora smaragdula, Lecanora dispersa and saxicola, Parmelia pilosella,
Buellia concinna.

IV. Pantemperate-Bipolar to Cosmopolitan element.— 38 sp. (23.2 %).

Normandina pulchella, Diploschistes actinostomus and scruposus, Peltigera
rufescens and polydactyla, Lecidea latypea and mutabilis, Catillaria intermixta,

292 C. SKOTTSBERG

Rhizocarpon geographicum and obscuratum, Cladonia bacillaris, coccifera, furcata,
eracilis, pyxidata, fimbriata and pityrea, Pertusaria leioplaca, Lecanora coarctata,
atra, polytropa and chrysoleuca, Placopsis gelida, Candelariella vitellina, Parmelia
laevigata, revoluta, cetrata, saxatilis, conspersa, perlata, cetrarioides and caperata,
Ramalina linearis and usnea, Usnea florida, Caloplaca elegans, Buellia stellulata,

Anaptychia hypoleuca.

Chapter III.

Composition, distribution and relationships of the Fauna.

There are no indigenous reptiles, amphibians, freshwater fishes or mammals
on the islands; of the introduced mammals, goats, rats and mice were naturalized
centuries ago and during the last 20 years also rabbits and Nasua rufa.

Aves.

LONNBERG (774); names in brackets used by GOODALL-JOHNSON-PHILIPPI (770).

Indigenous land-birds.

Turdus magellanicus King (T. falklandii magellanicus). Both islands. North
Chile to Patagonia and Fuegia; according to 57. V. 224 not known to be migratory.
“More richly coloured with buff below than specimens from the mainland avail-
able to me’, LONNBERG wrote l.c. 3, a colouring characteristic of 7. falkland-
cus (Quoy et Gain), comp. 5z. V. pl. XIII. In zzo this is called 7. f. falklandi and
the island form is referred to magellanicus.

Anaeretes fernandestanus (Phil.) (Spizitornis f.). Endemic on Masatierra. Azae-
vetes is a neotropical genus of 7 species (Ecuador to N. Argentina and Chile).
A. fernandezianus is related to a Chilean species.

Aphrastura masafuerae Phil. et Landb. Endemic on Masafuera. A second
species ranges from Centr. Chile to Patagonia and Fuegia.

Cinclodes oustaleti Scott ssp. baeckstroemit Lonnb. Both islands, endemic but
very near the typical species (Antofagasta—Chiloé). A genus of 13 species (Ecuad.
to Argent., Patag., Falkl. Is.).

Eustephanus fernandensis (King as 7rochtlus) Gould (Thaumaste f.). Endemic,
forming an endemic genus according to zro. 300: “‘Coloracion totalmente diferente
de la de cualquier Picaflor que habita el continente.’’ These authors leave open
the question whether or not £. deyboldzi Gould of Masafuera is distinct but list it
as Th. f. leyboldi and the form from Masatierra as 7h. f. fernandensis. LONNBERG
Pp. 7 gave good reasons for considering them as identical. The humming-bird
seems to be extinct on Masafuera.

Lustephanus galeritus Mol. (as Trochilus; Orthorhyncha sephanoides Lesson
et Garnot 1827—but MOLINA’s name must be about 50 years older; Sephanoides
s., 770). Masatierra and on the mainland from Centr. Chile to Fuegia. Said to
migrate to the coast and spend the winter there (774), a statement not quoted
by GOODALL.

DERIVATION OF THE FLORA AND FAUNA 293

Asio flammeus Pontoppidan (A. f. suinda Vieillot). Masatierra. The typical spe-
cies almost cosmopolitan; swzvda ranges over South America from Venezuela to
Fuegia.

Cercnets sparverius (L.) ssp. fernandensis Chapm. (Falco s. f.). Masatierra,
endemic, the species distributed from North America and the West Indies to northern
South America, another ssp. in Chile. A genus of 28 species and very wide
distribution, but not recorded for Oceania.

Buteo erythronotus King ssp. exsul Salvin (B. polyosoma e.). Endemic on Masa-
fuera, an accidental visitor to Masatierra; typical evythronoftus ranges from Peru
to the Magellan Straits and Falkland. On Masafuera the principal food of the
buzzard are rats, mice and young goats’ kids, all introduced by man, but it has
been observed attacking petrels and thrushes. A widespread genus of 33 species
(Amer., Euras., Austral., Ocean.).

Breeding sea-birds.

Fregetta grallaria Vieillot. Masatierra and Santa Clara and also Desventu-
radas (San Ambrosio); coasts and islands of the Indian and Pacific Oceans.
4 species, subtropical-tropical seas.

Puffinus creatopus Coues. Masatierra and Santa Clara; California—S. Chile,
where it breeds on Mocha I. and islands in the vicinity of Chiloé. Migrates during
the winter to Peru and along the coast north to Alaska, returning south in No-
vember. The genus (28 sp.) is world-wide.

Pterodroma neglecta Schleg. Masatierra and Santa Clara; also on San Am-
brosio and further reported from Lord Howe and Kermadec Is. Strolls north during
the winter. A genus of about 30 species spread over the south hemisphere and
extending north to the north Atlantic and to Japan.

Pterodroma externa Salvin (P. e. externa). Endemic on Masafuera, migrates
north as far as Costa Rica. Another race breeds on Tristan da Cunha, a third
on Kermadec Is.

Pterodroma cooki Gray ssp. defilippiana Gig). et Salvin. Endemic to Santa
Clara, Masatierra and Desventuradas. Typical cookz on New Zealand, P. cook? orten-
talis Murphy on the coast of Pert and Chile, but breeding places unknown;
defilippiana is said to extend its flights to Peru.

Pterodroma cooki ssp. masafuerae Lonnb. (P. leucoptera Masafuerae). Masafuera,
endemic. At first LONNBERG felt inclined to identify the bird with P. c. deucoptera
“in spite of the zoogeographical difficulties for such a theory” (p. 15); LONNBERG’s
opinion is strengthened by the fact that, whereas defilippiana and J/eucoptera are
surface-breeding like neglecta, externa and masafuerae are burrowing.

Two aliens are naturalized in the islands, a melanistic form of Columba livia
Briss. in the 18th century and now common, and Lophortyx californicus Shaw et
Nodd, introduced 1912 or 1913. Several landbirds have been observed as acci-
dental visitors, Crymophilus fulicarius and Butco obsoletus migrants from the north
and Belonopterus chilensts, Cathartes sp., Cygnus melanocoryphus, Haematopus ater
and Czrcus maculosus from the opposite coast. Petrels, albatrosses, Cape pigeons

294 C. SKOTTSBERG

and penguins are occasionally seen around the islands but do not breed
there.

Of the 15 species breeding on the islands 3 are endemic; of the remaining
12, 6 are represented by endemic subspecies of which, however, 2 also breed on
the Desventuradas Islands. Including these, 60% of the birds are endemic, a high
figure in animals as mobile as birds are. Otherwise, the poverty of the island ornis
is noteworthy, as also the fact that the affinities of the landbirds are all with
5S. "America:

With regard to the actual distribution we can distinguish the following two
groups.

I. South American (especially Chilean) group.——1o sp.

a. Endemic (6): Anaeretes fernandezianus, Aphrastura masafuerae, Euste-
phanus fernandensis, Cinclodes oustaleti baeckstroemi, Cercneis sparverius fer-
nandensis, Buteo erythronotus exsul.

4. Not endemic (4): Turdus magellanicus, Eustephanus galeritus, Asio flam-
meus, Puffinus creatopus.

II. South Pacific group.—5 sp.

a. Endemic (3): Pterodroma externa externa, cooki defilippiana and cooki
masafuerae.
6. Not endemic (2): Fregetta grallaria, Pterodroma neglecta.

The first group includes of more tropical birds Anaeretes, Cinclodes and Eu-
stephanus, and of more temperate 7urdus, Asio, Cerchners, Buteo and Puffinus.
Of the endemic species Lustephanus fernandensis is the most notable, in certain
characters a unique type in the family Trochilidae. The second group is of par-
ticular interest as including, beside the widespread frigate-bird, four species of
Pterodroma not breeding on the mainland, where, perhaps, a special race of
P. cooki breeds. The genus is essentially austral-circumpolar, as it were tricentric,
with Tristan da Cunha representing the African sector. Cases like those of P. zeglecta
and eaterna call for a common source and suggest that Prerodroma belongs to
an Antarcto-tertiary element which inhabited the coasts and islands of Antarctica
n preglacial times.

Oligochaeta.

MICHAELSEN (787) regards all the earth-worms of Juan Fernandez as adven-
titious. The single strictly South American Kerrza saltenszs was, he believes, in-
troduced from Chile with the human traffic, and this is also true of the three species
of Allolobophora, introduced to Chile from Europe, and by /redericia galba. The
occurrence of Pachydrilus verrucosus offers more interest. It was known from Great
Britain, the Hebrides, S.W. Africa and Fuegia, everywhere living on the seashore;
on Masatierra it was not found on the beach but inland in a freshwater stream.

Hirudinea.

The leech discovered in 1917 in the highland of Masafuera was described
as a new species of the Australian genus Philaemon, Ph. skottsbergi L. Joh. (£47).

DERIVATION OF THE FLORA AND FAUNA 295
Whether or not the two species described from Samoa and Madagascar, respec-
tively, belong to PAz/aemon remains to be settled. JOIANSSON expressed some
doubts with regard to the position of the Juan Fernandez leech and after his death
the question was taken up by NYBELIN who showed that it should form a separate
genus, Vesophilaemon (188). With its allies it forms a small austral group, pos-
sibly of old Antarctic ancestry. It must, of course, have a host; it was found in
the extremely wet Dicksonza forest, where Pterodroma cooki masafuerae makes
its burrows, the only possible host existing here (see also 747. 442). Truly no
leech has been collected on the bird, but extremely few specimens of this have
been taken care of and examined.

Crustacea.
Amphipoda (56).
Orchestia chilensis (chiliensis) Milne-Edw. Both islands, terrestrial and found

from near the shore to almost 600 m altitude. It is a bicentric species, known
also from Chile and New Zealand.

Isopoda (277).

Beside 3 cosmopolitan species 2 endemic ones have been found, both be-
longing to widespread genera, Ligza “itiginosa and Philoscia mirifica, the latter
referred to a new subgenus.

Arachnoidea.
Araneae (22).

I am indebted to Professor ALB. TULLGREN, who supplied much information
on the distribution of the genera. BERLAND lists 24 species, of which 4 are cos-
mopolitan and also occur in Chile; of the remaining 20 one, belonging to the very
large and widely spread genus Avaneus and perhaps new, was left unnamed. The
other 19 are enumerated below. Here as in the following + signifies an endemic
species, + + an endemic genus. Mt = Masatierra, SC = Santa Clara, Mf = Masafuera.

Ariadna maxima Nicolet. Chile-—Mt, Mf. The genus widely distributed
(N. and S. Amer., Afr., E. Ind., Australia).

+ Theridion Baeckstroemi Berl.—Mt. The genus is cosmopolitan.

Th. gracile Keyser]. Chile.—Mf.

+Lephthyphantes Fernandest Berl.—Mf. A cosmopolitan genus, but only a
single species recorded for S. America (Patagonia).

+ Macrargus pacificus Berl.—Mt. The genus is known from northern N. Amer-
ica and Europe, but TULLGREN (in litt.) doubts that it has a wide distribution
in America; thus, its appearance in Juan Fernandez is rather unexpected.

+M. australis Berl.—Mf.

+Leptorhoptrum (2?) Platet F. Cambr.—Mt. Doubtfully referred to this Eu-
ropean genus, from which it differs in certain characters: ‘‘l’épigyne est d'un type
tout a fait différent’’ (22. 430).

+ Tmeticus Defoet ¥. Cambr.—Mt. The genus used to be quoted from N.
America and Europe, but TULLGREN informed me that a great many species are
now referred to other genera.

296 C. SKOTTSBERG

Meta nigrohumeralis ¥F. Cambr.—Mt, endemic? (see l.c. 430). The genus is
known from all continents.

+Selhirkiella alboguttata Berl.—Mt. The genus is related to the neotropical
Guolus, known from Peru, Brazil and Chile, and appears not to be restricted to
Juan Fernandez; BERLAND has seen a very closely related species from Valdivia
(22. 432).

Mecysmauchenius segmentatus Simon. Patagonia and Fuegia.—Mt. A genus
of 2 species, the second one from the Magellanian region.

+ Misumenops Sjoestedti Berl—Mt. An American, especially N. American
genus.

+Gayenna Skottsbergi Berl.—Mt. A S. American, especially Chilean genus
of numerous species.

G. maculatipes Keyserl. Chile.—Mt.

+ Oxysoma Delfint Simon.—Mt. A S. American genus.

+ Philisca ornata Ber|.—Mt. A subantarctic-magellanian genus extending north
into Chile.

+Ph. mgens Berl.—Mt.

+Lycosa Fernandezi F. Cambr.—Mt. The genus is cosmopolitan.

Evophrys quilpuensis Simon. Centr. Chile-—Mt. The genus is known from
Centr. and S. America, Europe, S. Africa and Japan.

Of the 19 species enumerated 13 or perhaps 14 are endemic in the islands;
15 (11 or 12 endemic) are restricted to Masatierra, 3 (2 endemic) to Masafuera
and a single Chilean species found on both islands. The only conclusion we can
draw from these figures is that most likely only a minor part of the spiders occur-
ring on the islands is known. It is quite possible that there is a marked differ-
ence between the two islands, but it is not probable that Masafuera is so poor
and that Santa Clara is devoid of spiders. Only a short visit was paid to this islet.

The fauna makes the impression of being almost entirely neotropical or, at
least, S. American, with the exception of the two species of Macrargus, which
are of boreal parentage; of the doubtful Lepftorhopirum nothing can be said. The
presence of a southern, eventually Antarctic element is indicated by JZecysmauche-
mius, possibly also by Phzl7sca, but so far there is no sign of a bicentric group.
Whether it can be distinguished in subantarctic America I cannot tell.

Acarina.

TRAGARDH (268) enumerates 28 species, of which 2 are cosmopolitan, the re-
mainder endemic. He points out that the collection, the first ever made in Juan Fer-
nandez, undoubtedly represents only a small part of the acarofauna; this is evident
already from the fact that not a single species came from Masafuera or Santa Clara.
Whether the cosmopolitan species are late arrivals or not is impossible to tell, but
very likely they are. This would mean that the entire acarofauna is endemic, and
new investigations will not change its independent character unless some species
are discovered on the mainland. Of the 23 indigenous genera only one—probably

DERIVATION OF THE FLORA AND FAUNA 297

a second will have to be described—is endemic and several have a wide distri-
bution.

The scant knowledge of this neglected group in these regions, particularly
in Oceania, is to be regretted; it certainly does not yet lend itself to zoogeo-
graphical speculations. Nevertheless it deserves to be mentioned that Luv/ergus
similis Trag. belongs to a genus hitherto recorded only from New Zealand and
that Phyllhermannia dentata Trag. is related to a neozelandic species; the genus
is also found elsewhere.

Pseudoscorpionidea (79),

+ + Asterochernes vittatus Beier.—Mt. The genus has its greatest resemblance
to Thalassochernes Beier from New Zealand.

+ Chelanops insularis Beier.—Mt.

+Ch. kuschelt Beier.—Mt. Related to a Chilean species.

+ Geogarypus bucculentus Beier.—Mt.

+ Parachernes kuschelt Beier.—Mt, Mf.

++ Protowirthius fernandestanus Beier.—Mf.

+P. robustus Beier.—Mt.

Neotropical elements are present, but species with their relatives in the
Australian-Polynesian region are in dominance and part of the fauna shows not-
able archaic characters (l.c. 205).

Myriapoda.

The very small and incomplete collection—no specimens were brought from
Masafuera or Santa Clara—was studied by VERHOEFF (274). In order to get some
information on the distribution of the genera I asked Dr. OTTO SCHUBART of
Pirassununga, Brazil, for assistance, and he most liberally put his wide knowledge
of this group at my disposal (letter, Aug. 27, 1954). Several changes had to be
made in the nomenclature; the names used by VERHOEFF, if different, have been
put in brackets.

Diplopoda.

Brachyiulus pusillus Leach (Microbrachyiulus litoralis Verh.). Indigenous in
western Europe, adventitious in N. America and Argentina.

Brachydesmus superus Latzel. A European species, adventitious in N. America
and Argentina.

+ Aulacodesmus insulanus (Verh.) Schubart (Semnosoma, Verh.). Endemic. A
genus of 16 species, distributed over Chile and Argentina and belonging to the
austral family Sphaerotrichopidae (S. Amer., S. Afr., Madag., Nossi Bé, Austral.,
Tasm., N. Zeal., N. Caled., Hawaii).

Nopoiulus venustus Meinert (pulchellus Leach). Widely distributed in Europe,
introduced to N. America and Chile.

Cylindroiulus frisius oceanicus Verh. Typical friséus (C. Owent Bollman)
introduced to N. America, Argentina (also in forma oceanicus), S. Africa and St.
Paul's I.

298 C. SKOTTSBERG

Chilopoda.

+ Nesogeophilus laticollis (Attems) Schubart (Geophilus, Verh.). Endemic. The
genus, which has not been reported from S. America, includes after the latest revi-
sion by ATTEMS (as subgenus of Geophilus) 11 species (1 S.W. Austral., 1 N. Zeal.,
1 N. Caled., 1 Annam, 3 Jap. and 2 Eur,).

+ Nesogeophilus baeckstroem? (Verh.) Schubart (Geophilus, Verh.). Endemic.

Schizotenia alacer (Pocock) Silvestri. Chile, south to Fuegia, Argentina. A
genus of 6 species (3 Chile and Argent. to Patag., Fueg., 1 E. Austral., 1 N. Zeal.,
T@hatham’ 1s.)

Lithobiomorpha africana Porat (Lamyctes insignis Pocock, insignis baeckstroemi
Verh.). Widely distributed over Africa; also Tristan da Cunha, St. Paul’s I,
S.W. Australia and Hawaii. The genus very wide-ranging [N. Amer., W. Ind.,
S. Amer. (also Chile), Afr., E. Ind., Austral., Tasm., N. Zeal., Chatham Is., N.
Caled., Kermadec Is., Guam, Hawaiil.

If we exclude the 4 species regarded, rightly I presume, as introduced with
the human traffic, 5 species remain, 3 of these endemic in Juan Fernandez. This
is indeed a very small number, but in spite of being so few, they tell a story of
an austral-circumpolar, presumably Antarcto-tertiary element.

Collembola.

Of the 8 species distinguished by SCHOTT (276), the first ever collected in
Juan Fernandez, 2 inhabit Chile, 3 are known from various parts of the world
and 3 endemic. As long as so little is known about the distribution of this group
it does not lend itself to zoogeographical speculations. The occurrence of widely
dispersed boreal species in S. America and other parts of the south hemisphere
(Australia, New Zealand etc.) is noteworthy, but whether their wide range is due
to the great age of Collembola or a result of later dispersal is unknown.

Thysanura (222, 293).

+TIsolepisma annectens Silvestri.—Mt, Mf. The specific epithet refers to the
intermediate position between /solepzsma and Heterolepisma; the species is com-
pared with forms known from Africa and Australia.

+ + Kuschelochilis Ochagaviae WWygodz.—Mt. A monotypical endemic genus
related to Allomachilis and Nesomachil7s from Australia, but not, as far as known,
to an American genus.

Among the Invertebrates treated above the endemic leech offers great in-
terest. Of Arachnoidea the Pseudoscorpionidea include a remarkable Antarcto-
tertiary element, whereas the true spiders, strangely enough, are quite disap-
pointing in this respect, even more so than the centipedes.

Insecta.

In order to get an idea of the zoogeography of the island insects I asked a
number of specialists for information on the general distribution of genera and

DERIVATION OF THE FLORA AND FAUNA 299

species. For their readiness to supply me with the necessary data I am much
obliged to Dr. OLOF AHLBERG, Stockholm (Thysanoptera), Dr. KJELL ANDER,
Linképing (Orthoptera), Dr. PER BRINCK, Lund (Coleoptera), Mr. NILS BRUCE,
Gardby (Coleoptera), Dr. LARS BRUNDIN, Stockholm (Coleoptera), Mr. FELIX BRYK,
Stockholm (Lepidoptera), Dr. W. E. Cuina, London (Hemiptera), Dr. K.-H. Forss-
LUND, Stockholm (Trichoptera), Dr. G. J. KErRicH, London (Hymenoptera), Dr.
K. Princis, Lund (Orthoptera), Mr. Bo TyepER, Falun (Neuroptera) and Dr.
B. P. Uvarov, London (Orthoptera).

JOHOw (z50) enumerates 26 species of insects from Juan Fernandez; some finds
may, I presume, have escaped his notice, but probably not many, and it is evident
that the entomofauna was very little known at that time. During our survey 1916-17
a fair number of insects were collected and many novelties were described in vol.
3 of this work, but the collection gave the impression of being very fragmentary.
The intense collecting undertaken in 1951 and 1952 by the Rev. Dr. GUILLERMO
KUSCHEL revealed, however, the existence in the islands of a surprisingly rich
and varied insect world. As Dr. ALEXANDER, the wellknown specialist on Tipu-
lidae, expresses himself (4.35): “Father Kuschel’s collecting has completely
revolutionized our knowledge of the insect fauna of the islands in many groups, in-
cluding the crane-flies’’—only 3 species were known, the number now amounts to 37.

Until now only a part of Dr. KUSCHEL’s large material has been worked up
by specialists, and I can only refer to what has been published (142, 208, 292,
293, 309, 314), but for some groups we now have sufficient data to form an
opinion of the zoogeographical position of the islands as far as the insects go. At
the end of 1954 Dr. KUSCHEL joined my new expedition to the islands and brought
back a third very large collection. When all his material has been studied, the insect
fauna of Juan Fernandez will be better known than that of most isolated islands. At
present about 340 indigenous species have been recorded, of which about 230(70 %)
are regarded as endemic. Dr. KUSCHEL (letter, Oct. 16, 1955) calculates that of
a total of about 600 species collected by him, about 360 still await publication:
Among them are 25-30 flies, probably over 50 butterflies, many endemic, at
least 180 beetles (more than 120 weevils, of which 4 have been introduced
accidentally, the remainder being endemic), and some 40 hymenopters.

Orthoptera.
Dermaptera (225).

+ Euborellia annulipes (Lucas).—Mt, SC, Mf. The genus S. Amer., E. Afr.,
Orient, Ind., Ceylon, Tasm.

Anisolabis Bormansit Scudd. Galapagos Is., Easter I.—Mt. A large genus of
world-wide distribution.

Saltatoria (535, 225).

+ Hoplospyrium Skottsbergi Chopard.—Mt. An American genus, the species
related to species from N. America and Chile.
Trimerotropis ochracetpennis Blanch. Chile.—Mt. The genus is American.

300 C. SKOTTSBERG

Corrodentia.
Isoptera.

+ Kalotermes gracilignatus Emerson.—Mt. The only Termite known from Juan
Fernandez. “The wing venation is close to that of Kalotermes brouni Froggatt
from New Zealand” (83. 393).

Mallophaga (266).

Puffinus creatopus and Pterodroma neglecta and externa are infested with the
same mallophagous parasites found on these and related birds in other regions. A
new Halipeurus is mentioned but not described.

Thysanoptera ( 3).

Aecolothrips fasciatus L. Boreal.—Mt.

+Physothrips Skottsbergi Ahlb.—Mt. The genus is distributed over N. Amer-
ica, Europe, W. Asia and Australia.

+Sericothrips ineptus Ahlb.—Mt. Perhaps nearest to a Californian species.
The genus is otherwise confined to Europe, where it is widely spread.

Thrips tabact Lindem. N. America.—Mt. The genus known from N. America,
Eurasia, N. Africa and Australia.

The two non-endemic species may have been introduced accidentally. Both
were found in the spathe of Zantedeschia aethiopica, cultivated and naturalized.

Neuroptera (785, 92, 125).

++Conchopterella kuscheli Handsch.—Mt.

+C. maculata Handsch.—Mt.

Gayomyta falcata (Blanch.). Chile, Argentina.—Mt, Mf. A small S. American
genus.

+ Hemerobius Sioestedti Navas (H. fumosus Esb.-Peters., H. nigrinus Esb.-
Peters.).—Mt. An almost world-wide genus, absent from the S. hemisphere except
for the Andean region (Colomb., Pert, Boliv., Braz., Argent., Chile).

+H. Skottsbergi Navas.—Mt, Mf.

Trichoptera (274).

+ Australomyia masatierra Schmid.—Mt. The genus is known from Chile,
Patagonia and Falkland Is.

+A. masafuera Schmid.—Mf.

Verger Porter? Nav. Centr. Chile-—Mt. A Chilean genus.

Lepidoptera (73).

A great number of genera and species will have to be added when Dr.
KUSCHEL’s material has been determined.

DERIVATION OF THE FLORA AND FAUNA 301

Tineomorpha.
Gelechidae.

+ + Apothetoeca synaphrista Meyr.—Mt. The genus is closely allied to the large
and widespread Gelechia (N. and S. Amer., Galdp., Palearct., Macaron., S. Afr.,
Australia).

Oecophoridae.

+ Depressaria relegata Meyr.—Mt. Near a species from the Andes of Ecuador.
The genus is Holarctic and also found in S. Africa.

Endrosis lactella Schiff—Mf. Widely spread, domestic.

Tineidae.
Monopis croctcapitella Clem.—Mt, Mf. In most parts of the globe, domestic.

Pyralimorpha.

Pyralidae.

+Crambus fernandesellus Hamps.—Mt. A world-wide genus.

Elasmopalpus lignosellus Zell. Centr. and S. Amer.—Mt.

Ephestia kuehniella Zell. Widespread, domestic.

+ +lernandocrambus Baeckstroemt Auriv.—Mt, Mf. The genus nearly related
to Crambus.

+f. brunneus Auriv.—Mt.

+P, fuscus Auriv.—Mt.

++Fuania annulata Auriv.—Mt. Similar in some ways to Prochostola Meyr.
(S. Afr., Australia).

Nomophila noctuella Schiff. Cosmopol.—Mt, Mf; adventitious.

+Pionea fumtpennis (Warren) Hamps.—Mt. A world-wide genus.

Scoparia Ragonott Butl.—Mt, Mf. Chile. A very widespread genus.

Geometrina.
Geometridae.

+Eupithecia halosydne Prout.—Mt. A widespread genus, but not found in
Australasia.

+E. (?) zuepta Prout.—Mt.

+k. physocleora Prout.—Mt.

+ Lobophora msularis Auriv.—Mt. An essentially Palaearctic genus.
Lortricidae.

+ Crocidosema insulana Auriv.—Mt. A S. American genus.

+kulia griseiceps Auriy.—Mt. The genus Holarctic, also in Hawaii; few else-
where. :

+E. Robinsont Auriv.—Mt.

+E. striolana Auriv.—Mt.

Noctuina.
Noctuidae.

Copitarsia turbata Herr.-Sch. Venezuela, Colombia.—Mt. The genus in Mex.,
Centr. and S. Amer. (Argent., Chile).
Feltia malefida Guén. Amer. (south to Chile); Macaronesia.—Mt.

302 C. SKOTTSBERG

+ +Hoplotarsia magna Auriv.—Mt. Related to Copitarsza.

Leucania impuncta Guén. Chile-—Mt. A bipolar genus (Palaearct., N. Zeal.).

+Lycophotia Baeckstroemt Auriv.—Mt. The genus widespread (Arct., Amer.,
Eur., S. Afr., Madag., N. Zeal.). )

L. messium Guén. Chile, Patag.—Mt.

Rachiplusia nu Guén. Patag., Urug., Argent., Chile.—Mt.

Syngrapha gammoides Blanch. Chile-—Mt, Mf. A Palaearctic genus; Mex.,
SH Amer:

Rhopalocera.
Pyrameis carye Huebn. Widespread in S. America and probably introduced

in Juan Fernandez.—Mt.
Diptera (84, 742).

Acroceridae (208).

+ Ogcodes kuscheli Sabr.—Mt. A temperate genus, recorded from all con-
tinents, but only 2 species known from the mainland of S. America.
Anthomysidae (128).

Anthomyza cursor (Kieffer). Cosmopol., also S. Chile.—Mt.

Calliphoridae (255).

Callyntropyga humeralis (Walker) Souza L. et Alb. (C. Selkirki Enderl.). Chile
Concepcion).—Mt, Mf.

Paralucilia fulvicrura (Desvoidy) Aub. et Baxt.—Mt, SC, Mf.

Sarconesia chlorogaster (Wiedem.) Arrib. Chile, Easter I—SC, Mf.

Sarconesiomima bicolor Souza L.. et Alb. Chile (Santiago).—Mt, SC. A mono-
typical genus.

Cecidomyidae (84, 100).

++Psadaria pallida "nderl.—Mt, Mf. Related to Campylomyza Meigen.
Chloropidae (209).

Hlippelates australis Sabrosky |/f. (Cadrema) metadlicus Enderl. non Beck.
(=H. flavipes (Loew) Sabr.)|. Ecuad., Pert, Argent., Chile-—Mt, probably ad-
ventitious.

Culicidae.

Culex interfor Dyar.—Mt, Mf.
Dolichopodidae (126).

+fydrophorus kuscheli Harmston.—Mt.

FH. poliogaster (Phil.) Harmston. Chile-—Mt, SC.

+Sympycnus fernandezensts Harmston. Near a Chilean sp.—Mt, Mf.
Ephydridae (289).

Dimecoenia caesia (v.d. Wulp) Wirth. Argent., Uruguay.—Mt, SC.

+ Discocerina fumipennis Wirth.—Mt. Near a Chilean sp.

Flyadina certa Cresson. Chile.—Mt, Mf.
Hydrellia vulgaris Cresson. Guatem., Boliv., Chile.—Mt.

DERIVATION OF THE FLORA AND FAUNA 303

+Scatella angustipennis Wirth.—Mf. An almost Cosmopolitan genus.
+S. argentifacies Wirth.—Mt.

+S. drachyptera Wirth.—Mt.

+S. decemguitata Wirth.—Mt, SC, Mf.

+§. discalis Wirth.—Mt.

+S. fernandesensis Wirth.—Mt.

+ S$. kuscheli Wirth.—Mt, Mf.

+S. lutea Wirth.—Mt.

+S. marginalis Wirth.—Mt.

+S. masatierrensis Wirth.—Mt.

+S. minima Wirth.—Mt.

+S. nanoplera Wirth.—Mt.

+S. pallida Wirth.—Mt.

+S. pilimana Wirth.—Mf.

+.$. stenoptera Wirth.—Mt.

+S. vittata Wirth.—Mt.

+ Scatophila fernandeztana WWitth.—SC.

S. medifemur Wirth. Chile (Coquimbo).—Mt, SC.

Heleidae (288).
+Dasyhelea australis Wirth.—Mt, Mf. Near a Chilean sp., genus cosmop.
+Forcipomyia tenuisquamtpes Wirth.—Mt. A widespread genus (N. and S.
Amer., Eur., Afr., Austral.); one species common to Paraguay and Australia.
+F. sanctaeclarae Wirth.—Mt, SC.

Helomyzidae (128).
Blaesochaetophora picticornis (Bigot) Henn. S. Chile.—Mt.

Prosopantrum flavifrons TYonn, et Mall. (Cnemospathis Baeckstroemi et
Schoenemanni Enderl.). Chile, S. Africa, New Zealand.—Mt, Mf.

Lonchaetdae (128).

Lonchaea patagonica Malloch. Chile.—Mt.
Muscidae (128).
Austrocoenosia ignobilis (Stein) Hennig. Chile.—Mt.
Craspedochaeta limbinervis (Macq.) Hennig. S. Chile.—Mt, Mf.
Delia platura v. sancti-jacobi (Bigot) Hennig. Chile—Mt, Mf.
Euryomma peregrinum (Meigen) Hennig. Peru, Chile.—Mf.
Fannia anthracina (Walker) Hennig. Chile.-—Mt.
F-. canalicularis (L.) Cosmopol., also in Chile-—Mt, Mf.
F. punctiventris Malloch. S. Chile.—Mt.
Fucellia intermedia Lundbeck (Egeria masatierrana et masafuerana Enderl.).
Bur, Oceania.—Mt, Mf.
Hydrotaea cyanetventris Macq. Chile.—Mt, Mf.
Limnophora patagonica Malloch. S. Chile, Patag.—Mt.
+ Notoschoenomyza kuscheli Hennig.—Mt, Mf.
Ophyra caerulea Macq. Centr. Chile to Fueg.—Mt.

304 C. SKOTTSBERG

+ Schoenomysina emdent Hennig.—Mf.
+ Syllimnophora lispomima Hennig.—Mt, Mf.

Mycetophilidae (r00).
+Lxechia furcilla Freem—Mf. Near a Chilean species; world-wide genus.
+Leta malleolus Freem.—Mt. Allied to a species reported from Bolivia,
Pert and Brazil; the genus world-wide.
Macrocera funerea Freem. Chile-—Mt. A world-wide genus.
+Mycetophila angustifurca Enderl—Mt, Mf. The genus world-wide.
M. conifera Freem. Chile.—Mf.
M. cornuta Freem. Chile.—Mt.
M. flavolunata Freem. Chile.—Mt, Mf.
M. (2) tusecta Freem. Chile.—Mt.
Mf.
+M. subfumosa Freem.—Mt.
+Paraleia nephrodops (Enderl. s.n. Selkirkius) Freem.—Mt.
P. nubilipennis Walker. Chile-—Mf. The genus neotrop., Austral., Tasm.
Sciophila ochreata Phil. Chile-—Mt. A world-wide genus.

M. spinosa Freem. Chile.

Piophilidae (128).

Piophila caset (L.). Cosmopol., domestic.—Mt, Mf.

P. foveolata Meigen. Cosmopol.—Mt.
Phoridae (84).

+Lioyella guanfernandesica Enderl.—Mt, Mf. A European genus.
Phryneidae (128).

Phryne fuscipennis Macq. S. Chile.—Mt, Mf.
Platyp esidae.

Microsania pallipes Meigen. Cosmopol.—Mf.
Psychodidae (84, 2172).

Psychoda cinerea Banks. Cosmopol., also Chile—Mt. A widespread genus.

+P. masatierrensis Satchell—Mt, Mf. Possibly = the following.

+P. minutissoma Enderl.—Mt.

P. severint Tonnoir. Widespread in temperate regions.—Mt, Mf.
Sarcophagidae (255).

Hypopygta varia (Walker) Townsend. Chile-——Mt, SC, Mf.
Scatopsidae (84, T00).

++Masatierra ferruginea Enderl.—Mt. Related to the European Rhaeboza
Enderl. FREEMAN does not mention JJasatierra.

Scatopse fuscipes Meigen.—Mt, Mf. A world-wide genus.

S. notata (L.). Cosmopol., introduced with the traffic—Mt.

Sciavidae (160).

+ Bradysia fusca Freem.—Mt, Mf. A world-wide genus.

DERIVATION OF THE FLORA AND FAUNA 3905

+B. media Freem.—Mt, Mf.

+Mertanina kuscheli Freem.—Mt, Mf. Another species in Brazil.

+Psilosciara nitens Freem.—Mt.
Simultidae (294).

+ Gigantodax kuschelt \Wygodz.—Mt, Mf. An essentially Chilean-Patagonian
genus with single species as far north as Mexico.
Sphaeroceridae (84, 204).

Archiborborus submaculatus Duda. S. Chile, Patag—Mt.

++Gyretria binodatipes Enderl—Mt. The genus is perhaps identical with
Skottsbergia Enderl. and this is merged into Leftocera by RICHARDS who,
however, does not list the two Gyvetria species described by ENDERLEIN, but
not found in Dr. KUSCHEL’s material.

+G. crassicosta Enderl.—Mf.

Leptocera brachystoma (Stenhammar) Richards. Cosmopol., also in Chile.—
Mt, Mf. The genus is widespread and well represented in S. America.

+L. cultellipennis (Enderl. ut Skottsbergia) Richards —Mt.

L. darwint Richards. Chile, Argent.—Mt, Mf.

L. divergens Duda. Pert, Boliv., Chile, Argent—Mt, SC.

+L. duplicata Richards.—Mt.

+L. ellipsipennis Richards.—Mt.

L. flavipes (Meigen) Richards. Eur., N. Afr.—Mt.

L. mediospimosa Duda. Cosmopol.—Mt.

L. pectinifera (Villen.) Richards. Eur., Falkl. Is.—Mt.

L. pulchripes Duda. Argent., Parag., Urug.—Mt.

+ +Phthitia alexandrt Richards.—Mt. The genus must be very near Lepéocera.

+Ph. selkirki (Enderl. ut Pterodrepana) Richards.—Mt.

+Ph. venosa Enderl.—Mt.

Syrphidae (96).
Allograpta exotica (Wiedem.) (A. Skottsbergi Enderl.). Neotropical. An
American genus of at least 16 species, the majority in S. America.—Mt, SC, Mf.
+A. robinsoniana Enderl.—Mt. Mf.
Melanostoma fenestratum (Macq.) Fluke. Chile—Mt. An American genus.
+M. Lundbladi (Enderl. ut Carposcalis) Fluke.—Mt.
Eristalis tenax (1..). Cosmopol., also in Chile-—Mt, Mf.
Sterphus aurifrons Shannon. Chile—Mt.

Tachinidae (70).
Incamyia chilensis Aldrich. Chile-—Mt, SC, Mf. AS. American genus.
+Phantastosiphona kuscheli Cortés—Mt. A Centr. American genus.
Tendipedidae (288).
Anatopynia vittigera Edw. S. Chile, Patag—Mf. A world-wide genus.
+ Clunio fuscipennis Wirth.—Mf. A large, wide-ranging genus.
+ Hydrobaenus fernandezensis Wirth.—Mt, Mf. The genus world-wide.

20 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

306 C. SKOTTSBERG

H. pratorum (Goetgeb.) Coe. England, Chile, Patag.—Mt.

H. pusillus (Eaton) Coe. England, Kerguelen.—Mt.

+ Podonomus acutus \WWirth.—Mf. A genus of numerous species in southern
S. America, few elsewhere (Eur., N. Amer.).

+P. discistylus Wirth.—Mt, Mf.

P. kiefferi (Garrett) Edwards. Brit. Columb., Chile, Eur.—Mf.

+P. kuscheli Wirth.—Mf.

P. nigrinus Edwards.—S. Chile.—Mf.

+P. selkirkt Wirth—Mt, Mf.

Tanytarsus flavipes (Meigen) Townes. N. Amer., S. Amer., also Chile,
Eur.—Mf.

Tipulidae (4).

Evioptera pilipes (Fabricius). Cosmopol.—Mt, SC, Mf.

+Limonia (Dicranomyia) affadilis Alex.
ranging genus.

+L. amphionts Alex.—Mt.

+L. axterasta Alex.—Mt.

+L. harpax Alex.—Mt.

+L. kuscheliana Alex.—Mt.

+L. masafuerae Alex.—Mf.

+L. pedestris Alex.—Mf.

+L. selkirki Alex.—Mt, Mf.

+L. stuardot Alex.—Mt.

L. trituberculata Alex. S. Chile, Patag—Mt, SC (an endemic variety).

+L. venatrix Alex.—Mt.

+L. veneris Alex.—Mf.

+L. yunqueana Alex.—Mt.

+Molophilus amphacanthus Alex.—Mt. A world-wide genus well developed
on the mainland.

+M. antimenus Alex.—Mt, Mf.

+M. appressospinus Alex.—Mt.

+M. arciferus Alex.—Mf.

+M. canopus Alex.—Mt.

+M. defoeanus Alex.—Mf.

+M. distifurcus Alex.—Mt.

+M. filiolus Alex.—Mt.

+M. filius Alex.—Mt.

+M. masafuerae Alex.—Mf.

+M. multifidus Alex.—Mt.

+M. neptunus Alex —Mt.

+M. pectiniferus Alex.—Mt.

+M. rectispinus Alex.—Mt.

+M. selkirkianus (Enderl. ut Archimolophilus) Alex.—Mt.

+1. styx Alex.—Mf.

Mt, Mf. A very large and wide-

DERIVATION OF THE FLORA AND FAUNA 307

+M. tridens Alex.—Mt.

+M. variatus Alex.—Mf.

+M. yunquensts Alex.—Mt.

+ Shannonomyia kuschelt Alex.—Mf. An American genus, well represented
also in Chile and extending north to Canada.

+.Sh. masatierrae Alex.—Mt.

+ Sh. selkirkiana Alex.—Mt.

+ Tipula baeckstroemt Alex.—Mt. The genus world-wide.

Coleoptera.

As yet little has been published about the beetles collected by Dr. KUSCHEL;
to judge from what is known the number of island species no doubt will be
multiplied.

Anobtidae (195).

Anobium punctatum De Geer. Cosmopol., introduced.—Mt. Beside the typical

species an endemic ssp. described by Pic (Mt).
Mt.
+ Calymmaderus atronotatus Pic—Mt, Mf. Near a Chilean species. Numerous

A. striatum Ol. Cosmopol., introduced.

species in N. and S. America.

++Masatierrum impressipenne Pic—Mt, Mf. A genus near Megorama Fall.,
a small N. American genus.

Stegobium (Sitodrepa) paniceum (L.). Cosmopol., domestic.—Mf.

Xyletomerus pubescens ssp. kuscheli Pic—Mt, the ssp. endemic. A north
American genus.

X. pubescens var. picettarsis Pic (fumosus var., Pic)—Mt. The variety endemic.

Anthribtidae (153).
++ Opisolia lenis Jordan—Mt. Related to Lucyclotropis Jordan (Centr. and
S. Amer.).

Bostrychidae (166).
Neotertus pulvinatus Blanch. Chile-——Mt. A small genus reported from Pert

and Chile.
Prostephanus sulcicollis Fairm. et Germ. Chile.—Mt.

Carabidae (5, 260).

Bembidium inconstans Solier. Chile-—Mt. A world-wide genus.

L. punctigerum Solier. Chile.-—Mt.

Laemosthenes complanatus Dejean (Pristonychus, Gory, Andrewes). Cosmopol.,
also Chile; introduced.—Mt, Mf.

+Metius eurypterus Putzeys—Mf. A S. American genus, mostly in the far
south.

M. flavipes Dejean. Chile—Mt, SC.

+M. kuschelt Straneo.—Mt.

+M. ovalipennits Straneo.—Mf.

308 C. SKOTTSBERG

+ Plerostichus kuscheli Straneo—Mt. A world-wide genus.

+ Pt. selkirki Andrewes.—Mt, SC.

+Pt. skottsbergt Andrewes.—Mt.

+ Pt. walkeri Andrewes.—Mt.

++ Trachysarus basalis Straneo—Mt. The genus presumably endemic (260.
138).

Mt.
Mt.

bicolor Straneo.

T. emdent Straneo.
T. kuscheli Straneo.—Mt.

T. ovalipennis Straneo.—Mt.
T. pallipes Germ.—Mt, Mf.
T. punctiger Andrewes.—Mf.
ik

+ 4+ + + + 4+

+7. sericeus Andrewes.—Mf.

+ Trechisibus baeckstroemi (Andr.) Straneo.—Mf. An American genus.
T. femoralis Germ. Chile.—Mt (end. ssp.), SC.

+7. kuscheli Jeannel.—Mt.

+ Variopalpus crusoet Reed.—Mt, SC.

Chrysomelidae (282, 18).

++Minotula fernandezsiana Bechyné.—Mt. The genus is related to /Hypno-
phila (W. Eur.—Japan).

+M. kuschelt Bechyné.—Mt.

+I. nitens Weise.—Mt.

Ciotdae (166).

+ Cis bimaculatus Germ.—Mt. A world-wide genus.

+C. fernandeztanus Lesne.—Mt.

+C. rufus Germ.—Mt.
Cleridae (213).

Necrobia rufipes De Geer. Cosmopol.—Mt, introduced.
Coccinellidae (282).

Eviopis opposita Guér. Chile-—Mt. The genus ranges from Vancouver I. to
Patag. and Fuegia.
Colydiidae (200).

+ Pycnomerodes masafuerensis Pope—Mf. Beside the two Juan Fernandez
species there is a third in New Zealand.

+P. masatierrensis Pope.—Mt.

+ Pycnomerus insularis Grouvelle-—Mt. The genus N. and S. Amer., E. Ind.,
Japan, Australia, N. Zeal. (numerous), Samoa.

+P. germaint Pope.—Mf.

Cryptophagidae (36).
Cryptophagus atomarioides Grouv. (Selkirki Bruce’). Chile—Mt. A world-
‘Wetter’ 19:9. 1954.

DERIVATION OF THE FLORA AND FAUNA 309

wide genus, the subg. J/zzonomus, where the island species belong, in Eur., N. and
Centr. Afr., Macaron., Centr. Asia, probably also N. Amer.

+C. Skottsbergi Bruce.—Mt.

+C. splendens Bruce.—Mt.

++Cryptosomatula longicornis Bruce.—Mf.

+Loberoschema convexum Bruce——Mt. Beside the island species 3 in Bo-
livia and 2 in Chile.

+L. discoideum Bruce.—Mt.

Curculionidae (z2). Dr. KUSCHEL, who specializes in this family, estimates the
number of species collected by him to exceed one hundred.

++Anolethrus gracilis Auriv.—Mt.

++ Apteronanus dendroseridis Auriv.—Mt.

+A. (?) gunnerae Auriv.—Mt.

Aramigus Fullert Horn. A widespread noxious beetle.—Mt.

+Caulophilus (?) nigrirostris Auriv—Mt. The genus in southern U.S.A.,
Centr. and S. America. It does not exist on Juan Fernandez (KUSCHEL in litt.).

+ Cyphometopus masafuerae Auriv.—Mf. 3 species in Chile, where the island
species most likely also occurs (KUSCHEL in litt.).

++ Fuanobta ruficeps Auriv.—Mt.

++ Fuanorhinus Robinsont Auriv.—SC.

Ottorrhynchus rugosostriatus Goeze. W. and S. Europe.
introduced via Chile.

Mt, accidentally
+ + Pachystylus dimidiatus Wollaston.—Mt.

+P. nitidus Auriv.—Mt.

+ Pachytrogus crasstrostris \Wollaston.—Mt.

+ Pentarthrum affine Wollaston.—Mt. A widespread genus, found on many
oceanic islands.

+P. nigropiceum (Phil.) Auriv.—Mt.

+P. nitidum Wollaston.—Mt.

+P. rufoclavatum Auriv.—Mt. Close to P. afzcale Broun from New Zealand.

+ +Platynanus arenarius Auriv.—Mt.

+P. Baeckstroemt Auriv.—Mt.

+P. hirsutissimus Auriv.—Mt.

+P. quadratifer Auriv.—Mt.

+P. sericatus Auriv.—Mt.

+P. Skottsbergi Auriv.—Mt.

+ Strongylopterus nitidirostris Auriv.—Mt. The genus is also found in Chile
and New Caledonia.

S. ovatus Boh. Chile.

Mt, Mf.

Dermestidae (104).
Dermestes vulpinus L. Cosmopol., introduced.—Mt.
Dytiscidae (299, 117).

Antsomeria bistriata Brullé. Chile. —Mt. The genus known from Tristan da Cunha.

310 C. SKOTTSBERG

+Lancetes Baeckstroemt Zimmerm.—Mf. The genus austral bicentric.
Rhantus signatus ssp. kuscheli Guignot.—Mt, Mf. An endemic variety of

a Chilean species.

Lathridiidae (196).
+ Coninomus curtipennis Pic.—Mt. Near C. dimidiatus Belon (Boliv., Chile).
An essentially European genus.

C. subfasciatus Reitt. Chile-—Mt.

Melasidae (308).

+ Pseudodiaeretus Selkirki Flet—Mt. An Argentine genus.
Mycetophagidae (196).

Mycetophagus chilensis Phil. Chile-—Mt. The genus in Eur., Asia, Afr. and Amer.
Nitidulidae (196, 107).

+ Cnips acuta Gillogly—Mt. A Chilean genus.

+C. atrata Gillogly.—Mt.

+C. diversa Pic—Mt.

+C. fernandezia Gillogly.—Mt.

+C. mucronis Gillogly.—Mt.
Scarabaeidae (215).

Aphodius granarius 1. Cosmopol., domestic.—Mt.

Pleuropharus caesus Creutz. Reported from N. Amer., Chile, Eur., Orient,
Madagascar.—Mt, introduced.

Scolytidae (309).
Guathotrichus corthyloides Schedl. Chile-—Mt. The genus in N. and S. America.
Phloeotribus willet Schedl. Peru and Chile.—Mt.
Staphylinidae (25).
+Atheta Robinsont Bernhauer.—Mt. Near a Brazilian species. A cosmopoli-
tan genus of about 2000 species.

+Eleusis semtrufa Fairm. et Germ.—Mt. A genus of about 150 species, S.
Amer. (also Chile), Austral., N. Zealand.

+Medon crusoéanus Bernhauer—Mt. A cosmopolitan genus of about 500
species.

+ Ocyusa Baeckstroemi Bernhauer.—Mt. About 40, mainly Palaearctic.

Phitonthus nitidipennis Solier. Chile—Mt. A cosmopolitan genus of about
800 species.

+ Trogophloeus Skottsbergi Bernhauer.—Mt. A cosmopolitan genus of about
350 Sp.
Temnochilidae (196).

+ Phanodesta cribraria (Blanch.). Includes Ph. angulata Reitt—Mt, SC. A
Chilean genus.

+Ph. robusta Pic.—Mt.

+Ph. variegata Germ.—Mt.

DERIVATION OF THE FLORA AND FAUNA 311

Tenebrionidae (104, 196).

Blapstinus punctulatus Solier. S. Amer., also Chile-—Mt, SC. Numerous spe-
cies in N. and Centr. Amer., some in S. Amer.

+Enneboeus Baeckstroemi Pic.—Mt. Near a species from Panama. The genus
Mex.—Colomb., Tasmania.

Nycterinus gractlipes Phil. Chile-—Mt. Numerous species in Chile.

Hymenoptera.
Aphelinidae (211).
Aphelinus jucundus Gahan. N. America.—Mt, Mf.

Bethylidae (191).

+Cephalonomia skottsberg? Brues.—Mt.

++Lepidosternopsis kuscheliana Oglobin.—Mt.

+ Peristerola maculicornis Oglobin.—SC. A widely distributed genus of about
25 species.

+P. sanctae clarae Oglobin.—SC.
Braconidae (187).

+ Apanteles evadne Nixon.—Mt, Mf. A wide-spread genus.
+A. morroensts Nixon.—SC.

Aphaereta minuta (Ns.).—Mt.

+Opius kuscheli Nixon.—Mt, Mf.

+O. scabriventris Nixon.—Mt.

Dryinidae (191).

+ Haplogonatopus insularis Oglobin—Mt. A genus of 6 species (N. Amer.,
J. Fern., Australia, Pacif. Is.).

++Idologonatopus nigrithorax Oglobin.—Mf. A genus related to the former.
Elachertidae (211).

++Kuschelachertus acrasta De Santis —Mt, Mf.
+Pseudelachertus semiflavus De Santis—Mt. The genus otherwise Aus-
tralian.

Encyrtidae (211).

+Hemencyrtus kuscheli De Santis—Mt, Mf. A neotropical genus.

Entedontidae (211).

+Achrysochris bicarinata De Santis—Mt, Mf.

Euparacrias phytomyzae (Bréthes) De Santis. Chile, Argentina —Mt, introduced.

+Omphalomorphella elachertiformis De Santis. Said to come near an Aus-
tralian sp.—Mt, Mf.

Eulophidae (211).
+Diaulomyia calvaria De Santis ——Mt, Mf. Allied sp. in Australia.

312 C. SKOTTSBERG

Formicidae (283).
Ponera trigona Mayr var. opacior Forel. N. Amer., W. Ind., Chile, Argent.
—Mt. The typical species in Brazil. Probably spread with the traffic.
Prenolepis obscura Mayr ssp. vaga Forel. Melanesia—Mt. The typical spe-
cies Java, Australia, Hawaii, another variety N. Guinea and Melanesia.
Tetramorium guineense (Fabricius). An African ant, now widely spread with

the human traffic—Mt.

Ichneumonidae (206).

Enicospilus purgatus Say. Temperate N. and S. America.—Mt, Mf. An al-
most world-wide genus (Amer., Eur., Afr., Austral., N. Zeal., Hawaii).

+ Hemiteles Baeckstroemi Roman.—Mt. An almost world-wide genus, less
rich in the tropics.

+H. masafuerae Roman.—Mf.

+ Holocremna (?) juaniana Roman.—Mt. The genus is known from Europe.

Metelia (Paniscus) gerling? Schrottky. Chile—Mt. Range of genus very
wide, including S. Amer., Falkl. Is., Rodriguez, Austral., N. Zeal.

Stilpnus gagates Grav. var. Robinsont Roman.—Mf. The. typical species in
Europe, the genus also in N. America and Greenland.

Triptognathus aequicinctus Spin. Chile-—Mt.

Mymaridae (190).
Anagrus incarnatus Hal. Palaearctic—Mf, undoubtedly introduced.
++Cremnomymar fernandezi Oglobin.—Mt.
+C. wnperfectus Oglobin.—Mt.
++ Nesopolynema caudatum Oglobin.—Mt.
Polynema fuscipes Hal. Palaearctic—Mf, supposed to have been accidentally
introduced.

+ +Scolopsopteron kuscheli Oglobin.—Mt.

Oo
co)

Rhynchota.

Heteroptera.

Anthocoridae (21).

+ Buchanantella devia Bergroth—Mt. Related to B. continua B. White from
Madeira; other species reported from Tasmania and Hawaii.

Lyctocoris campestris Fabr. Cosmopolitan, probably adventitious.—Mt.
Ly gaeidae (21, 161).

++Micrymenus kuscheli Kormilev—Mt. Most nearly related to Metagerra
B. White from New Zealand (KORMILEY).

+iM. seclusus Bergroth.—Mt.

+Nystus Baeckstroemi Bergroth—Mt, Mf. An almost cosmopolitan genus
with numerous species in New Zealand and Melanesia, east to Samoa; greatest
concentration in Hawaii. V. Baeckstroemz is closer to N. Huttont B. White from
New Zealand than to any American species (KORMILEV).

+ + Robinsonochoris tingitoides Kormilev.—Mt. Forms a separate tribe.

DERIVATION OF THE FLORA AND FAUNA 5 SS

Miridae (50).

+Derophthalma fernandestana Carv.—Mt. A neotropical genus (Braz., Argent.
Urug.).

+ +Kuscheliana masatierrensts Carv.—Mt.
Nabididae (21).

Nabis (Reduviolus) punctipennis Blanch. Chile—Mt, Mf. A world-wide genus.

’

Reduviidae (292, 310).

Empicoris (Ploeariodes) rubromaculatus (Blackb.). Almost cosmopolitan.—Mt,
probably adventitious.

+Metapterus additius Wygodz.—Mt. A wide-ranging genus (Amer., also
Ghile, 5. Eur., N. Afr, W- Asia).

+M. kuschelt Wygodz.—Mt.

+M. masatierrensts Wy godz.—Mt.

Plotaria chilensis (Phil.) Kuschel.—Mt, Mf. Almost cosmopolitan; also in
Chile and probably adventitious in Juan Fernandez.

Homoptera.
Aphididae (communicated by Dr. KUSCHEL): 4 introduced species on garden plants.
Cicadellidae (57).

++Kvansiella kuscheli China.—Mt.

Delphacidae (314).

+ Nesosydne sappho Fennah.—Mt. A genus known before from south and
central Pacific islands including Hawaii, but never reported from America.

+ lV. minos Fennah.—Mt, Mf.

+ WV. oreas Fennah.—Mt.

+V. calypso Fennah.—Mt, Mf.

+N. philoctetes Fennah.—Mt, Mf.

+NV. vulcan Fennah.—Mt.

+ Delphacodes kuscheli Fennah.—Mt. A widely distributed genus.

+D. (Sogata) selkirkz (Muir) Fennah.—Mt.

Fasstdae (21).

++ Alloproctus amandatus Bergroth.—Mf.

Some soogeographical statistics.

Orthoptera.—Of the four species known 2 are endemic but of American
affinity, one a Chilean species and one reported from Galapagos Is. and Easter I.

Neuroptera.—Endemism strong, 4 species of 5, 2 forming an endemic genus,
the fifth an American species. Relations presumably Andean.

Lepidoptera —Of the 26 indigenous species 18 (69 %) are endemic, and there
are four endemic genera. Of these, Afothetoeca and Fernandocrambus are related
to world-wide genera also represented in S. America, //op/ofarsza to an American

314 C. SKOTTSBERG

genus; the systematic position of Fuanta has not been stated. Together they
include 6 species. Of the remaining 12, 7 belong to genera with a large to world-
wide area including at least some part of America, and one belongs to an American
genus. Lulia (3 sp.) and Lobophora are essentially boreal. The 8 non-endemics
are found in Chile or in some other part of S. America.

The total absence of all indigenous Rhopalocera is remarkable.

Diptera.—At present 157 named species belonging to 27 families have been
reported. Nine or ten species at least have been introduced with the human traffic»
147 are thought to be indigenous and of these 94, 64%, are endemic. Con-
sidering our insufficient knowledge of the dipterofauna of Chile, too much weight
should not be laid on these figures, but even if quite a few of the insular en-
demics will, in the future, be discovered on the mainland, I trust that a fair
number will remain, sufficient to show the peculiar character of the fauna. Be-
sides it can be foreseen that Dr. KUSCHEL’s new material will bring to light
some remarkable additions.

The fauna is not a haphazard crowd of wind-drifted flies. It gets its stamp
less from the few endemic genera—of 10 new genera proposed by Enderlein
only 4 remain—than from the presence of six non-endemic, 5. American or
more wide-ranging genera with six or more species each, Jolophilus with 19
(all end.), Scatella with 16 (all end.), Lzmonia with 13 (12 end.), Leptocera
with 10 (3 end.), Mycetophila with 7 (2 end.) and Podonomus with 6 (4 end.),
together 71 species of which 56 (79%) are endemic.

The Neotropical-Chilean character of the fauna is obvious. This is what we
expect quite apart from what we may think about the history of the fauna, but
the almost total absence of even a small austral-circumpolar or Pacific element
is noteworthy; the only examples would be Prosopantrum flavipes (austral-
tricentric) and /ucellia intermedia, said to be distributed over “‘Oceania’’. There
are some striking cases of disjunction, suggesting bipolarity (Lioyella, Flydrobaenus,
Podonomus Kiefferz), but the distances will perhaps be lessened when the dis-
tribution becomes better known.

Coleoptera.—| want to emphasize that of Dr. KUSCHEL’s collections ony 5
families have been worked out; it is to be regretted that no list of the Cur-
culionids is available. On the other hand I believe that the beetles inhabiting
the Chilean mainland are better known than the flies and some other insect
groups so that the proportion between endemics and non-endemics will not
undergo very great change in the future.

The number of named species hitherto reported from Juan Fernandez is
103, belonging to 19 families, perhaps little more than 1/3 of the species found
there. Eleven species are anthropochorous. Of the remaining 92 no less than 74 are
endemic—80 %, only 20% having been found elsewhere. Future research will
alter these figures, I suppose, a number of island endemics will be stated to
extend to Chile and vice versa, but on the other hand we have good reason
to expect that practically all Curculionids collected but not yet described will

prove to be endemic; of 22 indigenous species enumerated by AURIVILLIUS 21
were described as new.

DERIVATION OF THE FLORA AND FAUNA ZI5

So far 49 indigenous genera are cited, of which 10 are endemic; five of
these are Curculionids.
The non-endemic species are, with one exception—Aylelomerus pubescens, a

N. American species represented by 2 endemic varieties—also found in Chile.
As yet no austral-bicentric or -tricentric species have been found. However,
Pycnomerodes with 2 species in Juan Fernandez, 1 in New Zealand and none
elsewhere, as well as Pycnomerus, Strongylopterus, Eleusis and Enneboeus suggest
the existence of a small austral, possibly Antarctic element, even if the area
in cases extends north of the Equator.

Hymenoptera.—TVhe 35 registered species, 5 of them adventitious, cannot
represent but a minor portion of the fauna. Just as in all other insect groups
endemism is strong, 23 species are endemic (76.6 %) and of the 26 genera 6. The
affinities were, as a rule, not indicated by the authors; very likely they are,
with some exceptions, with 5. American forms. //af/ogonatus is essentially south-
ern, Prenolepis obscura is a southern, mainly Pacific ant. Aphelzus jucundus and
Stilpnus gagates are said to be boreal, but in these as in other similar cases
the possibility of accidental introduction must be considered.

Flemiptera—The 21 indigenous species—there are 7 adventitious ones—are
by no means a fair representation of the RKhynchota inhabiting the islands. Dr.
KUSCHEL’s collection contains twice as many species, more than half of them
endemic; of the 21 named species 20 are restricted to Juan Fernandez. Where
3 of the 4 endemic genera have their relatives I cannot tell, but the fourth, J/zcry-
menus, is most nearly allied to a genus in New Zealand. Buchananiella is quoted for
Madeira, Tasmania and Hawaii, but not from America, Wyszus, a world-wide
genus, has a stronghold in New Zealand and in the Pacific, and the single is-
land species stands nearer to a species endemic in New Zealand than to its
American congeners. It seems likely that we have to do with an austral-
antarctic element.

Mollusca.

The following synopsis is based on information supplied by Professor NILS
ODHNER who put his unique knowledge of this group at my disposal. The
additions and changes in his earlier list (759) are entirely due to him, and I
thank him for invaluable assistance. Possibly the new material brought back by
our 1954-55 survey will give additional taxonomic results, but they will not alter
the zoogeographical position of the fauna.

Lindodontidae.

++Amphidoxa helicophantoides Pfeiff—Mt. The genus (only 2 species known)
is related to Stephanoda. |

+A. marmorella Pfeiff—Mt.

+Charopa (Endodonta, s. lat.) zzvoluta Odhner.—Mt. Numerous species,
Polyn.2 NY Guint, ‘Australy: N)<Zeal:

+C. occulta Odhner.—Mt.

+C. skottsbergt Odhner.—Mf.

316 C. SKOTTSBERG

+ Punctum conicum Odhner.—Mt. A genus of a rather small number of spe-
cies, in various parts of the world.

+P. depressum Odhner.—Mt.

+ Radiodiscus masafuerae (Odhner) Pilsbry—Mf. An American genus of
few species, found in N. America (Arizona) and in S. America, south to Patagonia.

+ Stephanoda arctispiva Pfeiff—Mt. About 30 species in western S. America
(Chile, south to Fuegia).

+S. ceratoides Pfeiff.—Mt.

+S. guadrata Férussac.—Mt.

+S. selkirki E. A. Smith—Mt.

+S. tessellata Muehlf.—Mt, Mf.

Helicidae.
Helix aspersa Muell. Cosmopol., introduced to Chile.—Mt.
Limacidae. °

Agriolinax agrestis L. Very widespread, introduced to Chile.—Mt, Mf.
Limax arborum Buch.-Chant. As the former.—Mt, Mf.
Milax gagates Draparn. As the former.—Mt, Mf.

Succinetdae (subfam. Succineinae).

+Succinea cumingt Reeve—Mt. Succinea is taken in its old sense; it has
been split up, and the Juan Fernandez species belong to a group which must
bear a different name. It is reported from N. America, Panama, Galap. Is., St.
Helena, S. Africa, Hawaii and Tahiti.

+S. fernandi Reeve —Mt.

S. fragilis King. (syn. S. texta Odhner). Hawaii.

+S. gayana (D’Orbigny) Odhner.—Mt.

+S. masafuerae Odhner.—Mf.

+S. pingutis (Pfeiff.) Reeve —Mf.

+S. semiglobosa Pfeiff—Mt, SC.

Mt.

Tornatellinidae.

++Fernandeszia bulimoides Pfeiff. (incl. consimilis Reeve).—Mt.

+I. conifera Reeve —Mt.

2+. cylindrella Odbner.—Mt. Possibly identical with Zornatellina (Elasma-
tina) éurrita Anton, credited to Oparu I.

+F. diaphana King. —Mt.

+F. expansa Pilsbry.—Mt.

+F. tnornata Pilsbry.—Mt.

+F. longa Pilsbry.—Mt.

+f. philippiana Pilsbry.—Mt.

+f. splendida Anton.—Mt.

+f. tryoni Pilsbry.—Mt.

+f. wilsont Pilsbry.—Mt.

1
Rapa seems to be the name commonly used.

DERIVATION OF THE FLORA AND FAUNA 317

+ Tornatellina aperta Odhner.—Mt. A Pacific genus of over 50 species, ranging
from E. Ind. and Japan over Micron., Polyn. (incl. Hawaii), Melan. (Kermadec
Is., N. Caled.) to Austral. and N. Zeal.

1. bilamellata Anton. Recorded from Oparu I.

+7. callosa Odhner.—Mt.

+7. conica Anton.—Mt.

+7. plicosa Odhner.—Mt.

7. reclustata Petit—Mf. According to ODHNER probably identical with 7. Za7-
vita Anton.

+7. trochiformis (Beck) Pfeiff.—Mt.

7. trochlearis (Beck) Pfeiff. Oparu I.—Mt.

+ Tornatellinops minuta (Anton) Pilsbry et Cooke.—Mt. A Pacific genus of
22 species, reported from Japan, Philipp. Is., Polyn. and N. Zeal.

Zonttidae.

Hyalinia alliaria Miller. A widely distributed, anthropochorous species.—Mt.
FT, cellaria Miller. As the former.—Mt. Mf.

Forty-six species are enumerated; of these 6 have been introduced through
the human traffic. Of the remaining 40, 35 (87.5%) are supposed to be en-
demic; 13 belong to the two endemic genera. The occurrence of Succinea fragilis
in Hawaii and Juan Fernandez and nowhere else in surprising, and 4 species
are credited to Oparu (Rapa) Island, but the distribution is perhaps not too well
known. The poverty of Masafuera, where only 5 species have been collected, 3
of them restricted to this island, is, I daresay, only apparent. Additional forms
have been found later and still await study. On Santa Clara only empty shells
of a Masatierran species were found. This islet seems entirely unfit for land-shells.

Only two well-defined geographical groups are distinguished, to which a
third of wider extension is added.

I. American element.—S sp.
Amphidoxa (2), Radiodiscus (1), Stephanoda (5).
II. Pacific element.—23 sp.
Charopa (3), Fernandezia (11), Tornatellina (8), Tornatellinops (1).

III. Austral (or more widespread) element.—g sp.

Punctum (2), Succinea (7, see above).

Chapter IV.

Continental and Oceanic islands.

For a clear distinction between the two main kinds of islands WALLACE (278)
is as a rule referred to as the leading authority. From a geographic-geological
viewpoint an island, usually neovolcanic or coralline, which does not stand upon
a continental shelf, is called oceanic. If situated on the shelf there is a strong

318 C. SKOTTSBERG

possibility that, at some period of its existence, it has formed part of the con-
tinent. This is e.g. the case with the Falkland Islands. A truly oceanic island
lacks a continental basement of old, granitic or sedimentary rocks; at least, their
presence has not been demonstrated. It is, on all sides, surrounded by deep water
and a rise of perhaps thousands of metres is required to bring it into contact
with a continent. MAyYR (779), however, argues that, from a biological viewpoint,
every island, whether situated on a continental shelf or not, is oceanic which has
received its entire living world across the open ocean—consequently it must be
shown that every kind of organism present on the island has or once had the
faculty of migrating across the sea and establishing itself, either the species ac-
tually found or their ancestors.

It goes without saying that the answer to the question ‘continental or oceanic?”
should in the first place be looked for in the history of the oceans. With regard
to the Pacific our knowledge of its origin and history is incomplete, and even
if modern oceanographical research has supplied a wealth of information on the
hydrography, the nature of the sediments and so on, large parts of the southern
Pacific are little known and soundings so few that we cannot form but a very
general idea of the bathymetrical conditions and the configuration of the bottom.
The northern half is of course far better known. As it is, we must admit that
little or nothing has come to light that is opposed to the theory of the perma-
nence of the Pacific Ocean. It is, with few exceptions, from the biologists’ camp
that the theory has been attacked, particularly by phytogeographers; the majority
of zoologists seem to accept the conclusions arrived at by physiographers and
geologists. It is easy to understand, however, that many biogeographers, struck
by the perplexing disjunctions in the distribution of plants and animals, started
to build bridges across wide expanses of sea, in cases with a generosity that led
to absurdities. I have no reason to enter into details, our problem concerns Ant-
arctica, southern South America and Juan Fernandez, but even so it seems worth
while to quote a number of modern scientists, mainly geographers and geologists,
who have expressed their opinion on the nature and history of the Pacific Ocean.

Geotectonics of the Pacific Basin.

BAILEY WILLIS (28%) thinks that a suboceanic pressure works against the con-
tinents surrounding the Pacific, resulting in an expansion of the suboceanic mass
and a deepening of the basin which, in its turn, has a displacing effect on the
continental margin. He summarizes p. 367-368:

The consideration of the general facts of the geotectonics of the Pacific basin thus
leads us to regard the great ocean as a dynamic realm, within which the peculiar char-
acteristics of its rocks have facilitated the internal forces of the earth. The effects have
been as a whole to deepen the basin in consequence of the expansion of the under-
lying rocks. The expansion has in turn crushed the continental margins and raised the
great cordilleras. Geologic studies of the mountain ranges have demonstrated that the
actual orogenic period began in the Jurassic or possibly somewhat earlier in the Mesozoic.
Of the earlier periods we know but little, but the fragmentary records indicate that
periods of orogenic activity alternated with those of quiescence.

DERIVATION OF THE FLORA AND FAUNA 319

WILLIS did not question the permanerice of the basin; what interests us here
more particularly is his belief in the instability of the marginal regions.

In H. EK. GREGORY’s view the Pacific basin inside the deep troughs is an old
sink and it follows that all the islands within this sink are truly oceanic. Con-
sidering geological evidence alone there has been no significant change in the
position of Polynesian land masses since Pleistocene, most likely since early Ter-
tiary time: “There is no geologic evidence that any Polynesian island stood in
Jurassic or Cretaceous seas’ (775.1673). Still he thought that due regard should
be taken to objections raised by other branches of science, and he did not extend
the unaltered permanent basin outside the deep troughs. When, a little later,
another prominent geologist, J. W. GREGORY (776), expressed a different opinion,
this attracted a good deal of attention. The Pacific had, he says, been claimed
to have existed in its present shape and size throughout geological time, a hypo-
thesis almost universally adopted by geophysicists and geologists, but from
a biological viewpoint this theory did not satisfy. GREGORY was no believer in
large-scale transmarine migration of either plants or animals and consequently
inclined to consider the arguments put forth by the opponents to the permanence
theory. He counted with a number of Pacific seas separated by stretches of land,
and he looked upon the region where atolls serve as proofs of subsidence, a sub-
sidence which gradually enlarged the basin until it reached its present size, as
originally continental.

ANDREWS (6) who was a firm believer in successful transoceanic migrations
of all kinds of organisms and knew more about geology than most biogeogra-
phers, shared H. E. GREGORY’s opinion: islands situated within the area bounded
by the ocean deeps such as Hawaii, Marquesas, Society and Juan Fernandez, all
differing in their geological structure from the continents, are oceanic. Parts
of ANDREWS’ interesting paper deserve to be quoted here.

An examination of the continental blocks proper and the great western island
arcs suggests that they have had similar histories, whatever great differences may other-
wise exist between them. Japan, Eastern Australia, New Zealand, New Guinea, Fiji...
may be taken as examples. Each has a foundation of ancient folded and metamor-
phosed sediments, such as conglomerates, grits, quartzites, sandstones, slates, shales,
and limestones, and each of these foundations has been subjected to marked plutonic
intrusions of granitoid nature. Upon this foundation have been accumulated sediments
similar to those mentioned above, together with lavas not only of basic but also of
acid types. These, in turn, have been folded, overthrust, and invaded by plutonic rocks.
This generalization is true even though as yet no consensus exists concerning the age,
or ages, of the folded sediments and plutonic intrusives of the foundation rocks. It
would appear, however, that the foundation rocks of the island arcs which occur mar-
ginally to the continents of Asia and Australia are not as old as the earliest members
of the continental nuclei. This leads to the consideration of island arcs situated more
centrally within the Pacific Ocean. For this purpose, these may be considered as in-
cluding all the Pacific islands lying oceanward of the great island arcs mentioned above.
The principal examples include the Hawaiian Islands, the Marquesas, Juan Fernandez,
Easter Island, the Society Islands, the Cook Group, the Line Group, Micronesia, Samoa,
and the Ladrone, Caroline, and Pellew groups; Tonga and the Hebrides occupy a
peculiar position, mentioned below (p. 202—3).

320 C. SKOTTSBERG

Field observations show that these islands have had histories which present marked
differences from those of the continents and their marginal island arcs. Thus, they
appear to be composed almost entirely of volcanic material, mainly basic, together with |
“coral reef’ formations, whereas granitoid intrusives and acid lavas are jacking, to-
gether with the sediments invariably associated with “continental” areas. And not only
is this so, but the volcanic ejectamenta of these inner groups do not appear to con-
tain fragments of granitoids and sediments such as might be expected from volcanos
discharging through a foundation of rock formations such as compose the continents
(p. 203).

Another interesting feature is the peculiar topography of that portion of the Pacific
floor which separates the great island arcs and lands of ‘continental’ character from
the more central groups. Thus, on the American side of the Pacific, the “continental”
lands are separated from the groups—such as Hawaii, the Marquesas, the Society Is-
lands, and Juan Fernandez—by a series of deep discontinuous ocean trenches, prac-
tically collinear (p. 203).

The question is whether these trenches are of quite the same nature and
date from the same period as the deeper trenches arranged oceanward from the
great western island arcs. It is unfortunate for the advocates of a “‘continental’’
origin of the Juan Fernandez flora that these islands are situated on the wrong
side of the trench. However, the Galapagos Islands occupy a similar position,
and still they have been claimed, on good grounds, once to have been united
with Central America. In this connection another quotation from ANDREWS with
regard to the New Hebrides and Tonga is of interest.

Island groups which are difficult to place exactly in this scheme are the New
Hebrides, Tonga, and possibly the Pellews and the Ladrones. A profound deep lies
between New Caledonia and the New Hebrides, and this is suggestive of a noncon-
tinental origin of the group. On the other hand, the occurrence of mineral deposits
such as copper, iron, and nickel, of large kauri, fig, myrtaceous, and other trees, and
of animals such as lizards, turtles, ducks, pigeons, and parrots, suggests that they may
well have formed, at some earlier time, portions of a continental margin which later
became involved in a powerful movement within the marginal Pacific, resulting in the
gradual submergence of these outer portions, the present Hebrides, Tonga, and so on,
being built upon such sinking area. This certainly is suggested for the New Hebrides
and for Tonga, while the Pellew and Ladrone islands also have had complex histories,
which would well repay close attention, in their structural, petrological and biological
aspects (p. 203).

An additional point of interest is the association of great ocean deeps with youthful
volcanic zones, and inasmuch as the trough and crest of an earth undulation are parts
of the same structure, it is a legitimate inference that the great Pacific deeps or trenches
are relatively youthful structures (p. 203).

ANDREWS summarized his idea of the Pacific basin in a number of points,
which, with very slight verbal alterations, form the introduction to his paper on
the origin of the Pacific insular floras (7.613-14):

1. The continents bordering the Pacific have been larger, at various times, than
they are at present.

2. The great bordering island arcs of the Pacific—such as the Aleutians, Japan,
the Philippines, the Netherlands East Indies, New Guinea, Fiji, New Caledonia, and
New Zealand—have been connected directly with the continental lands. Certain of these

DERIVATION OF THE FLORA AND FAUNA 321

island groups—such as Fiji, New Caledonia, and New Zealand—appear to have been
isolated at much earlier periods than others, such as New Guinea, the East Indies,
and Japan.

3. The ancient borderlands of the continents have, in part at least, suffered un-
dulatory submergence. Compensatory forms are the deep ocean trenches, on the one
hand, and the mountain ranges of the continents, on the other.

4. The Pacific is a relatively deep and unstable area, whereas the Atlantic—with
the exception of the broad intersected belt of activity directed toward the equator—
has, on the whole, been relatively stable since the Palaeozoic.

5. The western area of the Pacific appears to possess a more complex structure
than the eastern, owing to the earth’s rotation, the width and weakness of the Pacific
base, and the resistance opposed to this activity by the stable continental masses of
Eurasia and the Australian—Sahul area.

6. The islands of the Pacific lying within the area bounded by the ocean trenches
have not had continental histories, nor do they appear ever to have had actual and
direct land connections with the continents.

7. The New Hebrides and Tongan Islands, and possibly also the Pellew and Mari-
anne groups, appear to partake in some measure of the nature both of “‘continental”’
and “‘oceanic’’ islands.

We shall have occasion to return to ANDREWS’ opinion on the origin and his-
tory of Pacific floras. Here it seems convenient to draw attention to the numerous
submarine cones called “‘guyots’ recently discovered in the sea between Hawaii
and the Marianas and discussed by HEss (732). About 160 flat-topped peaks,
presumably truncated volcanic islands, rise from gooo to 150cO feet above the
deap-sea bottom. In most cases their flattened summit was sounded in about 800
fathoms. HEss’ working theory is that they were formed on land, sunk to their
present level and levelled by sea action—this would mean that they stood with
their summit at sea level long enough to be exposed to wave action. Thus,
drowned reefs could be expected, but no such are reported: the guyots are,
HEss thinks, very ancient structures dating from a “proterozoic episode of vul-
canism”’, they are of pre-Cambrian age and consequently do not lend themselves
to biological speculations. There may, however, have existed later islands, both
Paleozoic and Mesozoic, but all have disappeared beneath the surface of the ocean,
either built up again by reef-building organisms or sunk to a depth where these
cannot live. The high volcanic islands are very young, perhaps not even Tertiary
but Pleistocene or recent, because the rocks could rarely be proved to be of Ter-
tiary age. We shall see by and by that the endemic insular floras and faunas cannot
be anything like recent. ‘““Oceanic islands’, HESS continues, “are and have always
been slowly sinking relative to sea level’’ as a consequence of the accumulation
of bottom sediments causing the water level to rise. The red clay increases I cm
in 10000 years, the globigerina ooze the same amount in 5000 years. Thus millions
of years are needed to account for even a very moderate submergence. Besides,
raised shore-lines are, in many cases, unmistakable proofs of local emergence.
The melting of the great inland ices ought to have had a greater influence.

Finally, let us listen to one of the foremost authorities on geophysics, GUTEN-
BERG (223). He finds (p. 7) that there is

2I — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

322 C. SKOTTSBERG

growing accumulation of evidence that the Pacific basin shows unique features which
are not duplicated in any other oceanic or continental area of the earth. There is no
feature on the surface of the earth which compares in dimensions and importance with
the Marshall line, within which the younger eruptive rocks are basaltic rather than
andesistic. This discontinuity in the material of the crustal layers is called here the
boundary of the Pacific Basin.

This is in conformity with what I have quoted above from other sources.
However,
certain areas of the Pacific Ocean (near its borders, for example), at least part of the
region between South America and the Easter Island rise, or between the Marianas

and the Asiatic continent, show indications of continental layers. For the latter. petro-
graphical and geophysical evidence agree.

As seen on the map, GUTENBERG goes a good way beyond the Juan Fer-
nandez-San Ambrosio rise, but the Easter Island shield which, excepting the vi-
cinity of this island, is covered by very deep water, belongs to the wide basaltic
centre, where continental layers are lacking—in contrast to the Atlantic where
‘‘sranitic layers of the continents continue far out under the bottom . . . probably
at least some continental rocks underlie its bottom throughout its area’.

Turning to the speculations of biologists I shall quote some representatives
from the two opposite camps. ARLDT (&) did not draw his conclusions merely
from facts of distribution but compiled a wealth of geological, palaeontological,
bathymetrical dates and so on, and constructed a series of maps illustrating the
distribution of land and sea through earth’s history. A Cretaceous Oceania united
South America with Australia + New Zealand, it disappeared during Eogene and
left the west coast of South America in the same position as to-day. From what
he says about Juan Fernandez it appears that he regarded these islands as con-
tinental (see below p. 376), while still admitting the possibility of oversea migra-
tion from the coast. CAMPBELL was for a long time a supporter of the land-bridge
theory. He regarded the Hawaiian Islands as formerly much larger and more closely
connected with land masses to the southwest, having become isolated during
early Tertiary time coincident with the uplift of the great cordilleras (zg). Later,
when discussing the Australasian element in the Hawaiian flora he expresses him-
self very positively: “We are justified in assuming the former existence of land-
masses of considerable size, connecting more or less directly both Australia and
New Zealand with Hawaii” (gs5.221); and when, for the third time, he took up
the history of the Hawaiian flora, he expressed himself as follows (46. 181):

We may assume that the Hawaiian Archipelago, as it now exists, is but a remnant of
a much larger land-mass which has been in subsidence for a long period, and that extensive
subsidence has also occurred throughout Polynesia, and to a lesser degree in Australasia.
One argument for this assumption is the great development of coral reefs in the Pacific,
especially in Polynesia and northeastern Australia. The existence of active coral reefs
involves continuous subsidence and the absence of large land-masses in mid-Pacific, with
the innumerable small coral islands and reefs, can be explained most satisfactorily on the

theory that the latter are remnant of submerged land-masses of large size—possible even
of continental dimensions.

DERIVATION OF THE FLORA AND FAUNA 323

It is surprising that, fourteen years later, he was converted to wegenerianism
in the form modified by Du To!r, thinking that “this theory would best explain
most of the problems in the geographical distribution of the floras of the Southern
Hemisphere” (47.70)—he seems to have forgotten how badly Hawaii fits into
the picture.

GUILLAUMIN, in his paper on the floristic regions of the Pacific (778), devotes
a chapter to its geotectonic history.

Un vaste continent paralléle a l’équateur a du relier les régions australiens, canaque,
et néozelandaise a l’Amérique du Sud tandis que la région malayo-polynésienne et le
domain hawaiien, formant peut-étre un continent, en étaient séparés.

The dissolution and disappearance of this hypothetical continent took place
before the end of the Jurassic, so that its direct influence on the distribution of
angiosperms must have been slight; ARLDT’s ““Ozeanis’’ was more useful. The
Melanesian extension of Australia-New Zealand, including Fiji, Kermadec, Tonga
etc., persisted longer and is perhaps better founded, but when GUILLAUMIN thinks
that his hypotheses “‘ne sont pas en contradiction avec les données géologiques
ni avec ce qu'on sait du relief sousmarin du Pacific’, he moves on unsafe ground.
Most authors who deny any considerable reduction of the surface of the Pacific
admit that Melanesia forms an exception. Thus Guppy (z2z) who otherwise is a
firm believer in oversea migration thinks that “we should rope in Fiji with all
the large islands westward and southward as originally continental’ because they
lie inside the gymnosperm line. All non-coniferous ones are excluded. GERMAIN
(705.973) goes much farther. Discussing the Hawaiian region he thinks that it
may have extended far eastward. The distribution of some animals, for instance
the eel, proves that Hawaii was “‘part of an oriental Pacific continent, perhaps
also united with the Juan Fernandez and Easter islands’, and he suggests that
many of the Polynesian elements in Hawaii travelled by way of South America
and Juan Fernandez (p. 1009).

Among botanists SETCHELL, pointing to the evidence furnished by geology,
expresses his opinion on Pacific paleogeography in the followings words.

I see no necessity of postulating any fundamental changes from the point of view
of the permanence of the Pacific Ocean as such, and the purely volcanic origin, prob-
ably in Tertiary times, of the islands existing in it, in much the same position as we
now find them (279. 301).

Another prominent botanist who has strong claims to the title “Defender of
the Oceanic Faith’, is FOSBERG. He refuses to recognize any Pacific islands at
all as continental, even Fiji (98.164 etc.); though the flora is “plutot de nature
continentale’”’ it is not necessary to count with terrestrial connections to explain
its characteristics. Among zoologists, MAYR (779) and ZIMMERMAN (298) belong to the
same camp, and so do most of them, but it happens that certain animal groups,
particularly the land molluscs, offer serious difficulties and have led otherwise
conservative zoogeographers to take refuge in ‘‘a mid-Pacific land’. So for in-
stance MUMFORD (783. 247):

324 C. SKOTTSBERG

Evidence with regard to the nature and distribution of land snails is of the utmost
importance, for it is from the nature of the land-snail fauna in Polynesia that Mr. PrtsBry
has reached the conclusion that the whole of Polynesia, with Hawaii, was once a great
continental land mass. artw/ia—an ancient generalized type of land snail—is wide-
spread in Polynesia and Melanesia and not elsewhere; whereas many groups (e.g. He-
licidae and Arionidae) found widely spread in the world are absent in Polynesia, etc.
Professor Buxton is of the opinion that Pilsbry has been more successful than any other
writer in establishing a case for the early existence of a mid-Pacific continent.

But how early or how late? To explain the absence of modern land molluscs
we may have to go back to late Tertiary times only, and we should need other
proofs of a distribution of land and sea sufficiently different from the present one;
perhaps they will be found. But we cannot simply fill out the Pacific basin with
land and leave the surrounding continents unaltered. Often enough due regard
was not taken to such circumstances. SWEZEY (262), who for his own part believed
that the entire insect fauna of Hawaii owed its presence to accidental migration,
quotes two of the authors of “Fauna Hawaiiensis’ who did, in his opinion with-
out any reason, build bridges where they found that they needed them, MEYRICK
and Lord WALSINGHAM. MEYRICK, an authority on Microlepidoptera, when stating
that among the endemic Hawaiian genera three were of south Pacific affinity, pos-
tulated “the former existence of a considerable land area (now submerged) be-
tween New Zealand and South America’, a land mass which compares to ARLDT’s
South Pacific bridge—such a land still exists and is not submerged: Antarctica!
—and he also believed in a ‘‘Palaeonesia’’ extending from Rapa to Hawaii and
from Pitcairn Island to the Society and Cook groups. Little room is left for the
water of the ocean, but he does not argue that these land masses were contem-
poraneous. Lord WALSINGHAM who based his opinion on the distribution and
relationships of the Microlepidoptera regarded the Hawaiian Islands as representing
the summits of mountain ranges formerly belonging to a continent, ‘‘a lost Paci-
fica’; if not accepted, ‘‘some other theory possibly even less acceptable must be
devised’’—equally beyond the possibility of exact proof.

It is easy to understand that, in all these discussions and speculations, Hawaii
in its isolated position and with its rich flora and fauna should be the object of
the main interest, and I shall end this chapter with a review of E. H. BRYAN’s recent
contribution, ““The Hawaiian Chain’ (zo), equally instructive as popularly written.
BRYAN believes in a continental Melanesia but regards Hawaii as oceanic, but
this does not prevent him from accepting the Leeward islands as remnant of a
long, deeply submerged ridge, ner from admitting that the Hawaiian Islands proper
may have been greater and, in part at least, united. The chain is supposed to
have emerged first at its extreme western end, where we now only find the rem-
nants of once larger islands; it appeared some time during Tertiary, the forma-
tion proceeding toward the east, with the island of Hawaii, where ejection of
lava still occurs, as the youngest link, the chain having been completed by the
end of the Pliocene. BRYAN, who has a wide knowledge of the Hawaiian fauna, is
no friend of land bridges and less so of submerged continental masses, but un-
like many other zoologists he admits that it is very difficult to explain the fauna

DERIVATION OF THE FLORA AND FAUNA 325

without the help of some kind of closer contact with other lands. He looks
around for bridges in the shape of “‘stepping stones’, just as MAyR does, ‘‘tem-
porarily connected or sufficiently close together’. “It is,’’ he continues, “a long
distance between Guam and Fiji and Tahiti and Hawaii, but if there were numer-
ous other islands spread conveniently between ...’’; speaking of the weevils of
Necker Island and of Nihoa, Wake and Laysan where, with the exception of Nihoa
with its endemic palm and Laysan, once the home of an endemic form of Sav-
talum, no suitable host plants exist, he finds strong evidence for their represent-
ing “the last remnants of former forest insects, surviving along a route of migra-
tion, a land bridge of the past’’—this applies, I daresay, to an earlier connection
between the links of the broken Hawaiian chain. However, it seems to me that
rows of ‘conveniently spaced stepping stones’, sufficient to offer routes of migra-
tion from several directions, involve tectonic movements of considerable magnitude.
Few conscientious bridge-builders would argue that, for instance, a solid land
mass extended from Melanesia. and Indonesia to Hawaii as a continuous open
road, it might have risen gradually from west to east—when the land upon
which the present Hawaiian Islands were built, was above the sea, the western
part of the bridge had disappeared; all that was left was a detached, “‘advanced”’
portion of a borderland, the home of a facies of the Australian-Malaysian fauna
and flora, which gradually took possession of the rising volcanic soil of Hawaii.

Chapter V.

The Pacific Ocean and Continental Drift.

It serves no purpose to dwell here at any length on WEGENER’s original
hypothesis, with which every biogeographer is familiar, but it may be useful to
scrutinize its bearing on Pacific problems in general and Juan Fernandez in par-
ticular. Before the breaking up of Pangaea, the Pacific Ocean was twice as wide
aS now, an enormous water desert where no islands enlivened the seascape. The
entire sial crust revolved west, where festoons were successively split off from
the Asiatic-Australian land mass, got stuck in the sima forming one island arc
after the other, bordered on their outside by deep trenches. The Americas trav-
elled at a greater speed away from Europe-Africa, and the Pacific became less wide.
It is true that most of the island chains in the Pacific trend NW-SE, but there
are many islands that do not follow this pattern, among them Juan Fernandez;
nevertheless, even if he did not expressly state this, it seems clear that WEGENER
regarded all of them to be of the same origin. In an earlier paper (237) I re-
ferred briefly to what he said about Juan Fernandez; here I shall quote him in full
(280. 116).

Die pazifischen Inseln (mitsamt ihrem submarinen Unterbau) werden in der Ver-
schiebungstheorie als von den Kontinentalschollen abgeloste Randketten betrachtet, die
bei der allgemeinen, vorwiegend westlich gerichteten Bewegung der Erdkruste tiber den
Kern allmahlich nach Osten zuriickgeblieben sind. Ihre Heimat ware hiernach, ohne auf

326 C. SKOTTSBERG

Einzelheiten einzugehen, auf der asiatischen Seite des Ozeans zu suchen, der sie jedenfalls
in den betrachteten geologischen Zeiten erheblich naher als heute gelegen haben miis-
sen. Die biologischen Verhiiltnisse scheinen dies zu bestatigen. So haben nach Grise-
bach und Drude die Hawaiinseln eine Flora, die am nachsten verwandt nicht mit
Nordamerika ist, das ihnen doch am nichsten liegt, und von dem heute Luft- und
MeeresstrOmung herkommen, sondern mit der alten Welt. Die Insel Juan Fernandez
zeigt nach Skottsberg gar keine Verwandtschaft mit der doch so nahen Kiiste von
Chile, sondern mit Feuerland, Antarktika, Neuseeland und den anderen pazifischen
Inseln. Doch sei hervorgehoben, dass die biologischen Verhaltnisse auf Inseln allge-
mein schwerer zu deuten sind als diejenigen auf grésseren Landrdumen.

In the 3rd edition p. 59 we read after “‘Inseln”’: “Dies passt vorziiglich zu unserer
Vorstellung, dass Siidamerika, nach Westen wandernd, sich ihr erst in letzter
Zeit so weit genahert hat, dass der Florenunterschied auffallend wird.” In the
following editions this sentence was excluded.

Certainly I never said anything like that and I fail to see where WEGENER
got his strange ideas; just as many other writers I have pointed out that the Andean-
Chilean element is stronger than any other. Even to a firm believer in the festoon
theory the Juan Fernandez and Desventuradas Islands ought to offer insuperable
difficulties. WEGENER built his theory on the island arcs accompanying the Asiatic-
Australian continental border; geologically these arcs are continental, but when
he came to island chains like Hawaii, the Marshall Islands and the Society Islands
—and we can add Marquesas, Tuamotu etc.—all of which are situated outside
the deep trenches, neovolcanic and regarded as built up from the depths of
the ocean—he was driven to assume that they have a sialic basement hidden
under the basaltic layers. He thinks that this assumption is supported by pend-
ulum observations, the force of gravitation being greater over the islands than
over the open ocean where, of course, a sial cover is incompatible with his dis-
placement theory.

WEGENER’s theories were taken up by Du ToirT and presented in a modi-
fied form (Sz); I shall quote his attitude toward the festoons.

As WEGENER has observed, they are all comparable in size, regular, linked to-
gether en échelon and convex to the Pacific; each shuts off a large portion of sea and
fronts an oceanic deep, while the concave side bears a row of volcanoes. To Suess we
owe the conception of the development of successive arcuate asymmetrical fold-waves mi-
grating outwards from the more stable ‘‘Amphitheatre of Irkutsk’’, which led to progressive
expansion of Asia towards the Pacific. While the hypothesis has since had to be ap-
preciately modified, its fundamental ideas have been brilliantly confirmed by subsequent
investigations... Significant are the oceanic fossae that immediately front the convex
sides of the arcs—foredeeps subsiding in advance of the outward-moving geoantoclines
and incidentally tracts of marked coastal instability (pp. 186-187).

How far did this outward-movement, these advance-folds proceed? Does Du
To1r allow all the Pacific island chains to be linked up here? When the great
WNW swing of Asia is replaced by an expansion toward the Pacific, the system
of rifts in the ocean floor, over which the island chains as claimed by most
geologists were formed, did not exist, because there was no tension to account
for them; instead, series of ripples were crumpled up on the floor. The transfor-

DERIVATION OF THE FLORA AND FAUNA 327

mation of the geanticline structures into rows of islands is explained by Du Torr
as follows; their crest

could become deepened by crustal tension and broken into segments to form an island
chain before vanishing... limbs were intermittently built up and destroyed during the
Cretaceous-Tertiary through stretching in the direction of their length while they were
still compressed by forces at right angles thereto (p. 293).

Trying to apply these ideas to South America, which according to Du Toir
as well as WEGENER was pressing into the Pacific basin, the Juan Fernandez—
Desventuradas—Merriam ridge could be compared to an advance-fold. But it is not
convex to the Pacific, nor fronted by a fossa—this is situated on the wrong
side and may well stand in causal relation to the upheaval of the Cordillera. To
think that the submarine ridge emerging in the Juan Fernandez and Desventu-
radas Islands is the easternmost advance-wave from a western borderland seems
too phantastic.

Du Tott’s idea of the geological character of the ocean floor differs from
WEGENER’s. Seismographic records, he says, scarcely bear out that the Pacific
floor must be composed of basalt—the records could readily accord with a
granitic layer up to about 10 km thick (p. 212). He was no believer in a more
or less unlimited oversea migration of plants and animals, nor in land-bridges,
and he critizises J. W. GREGORY and the bridge-constructing biogeographers: they
are wrong, and the displacement hypothesis interprets otherwise. But when he
speaks of the extensive ‘‘march into the ocean of crustal waves, thereby leaving
their parent continents far in the rear’ and of the “rhythmic intrusion, culminat-
ing in the three migrations of the Cretaceo-Eocene, mid-Tertiary and late Ter-
tiary” (p. 214), these advance-folds, when crumpled up from the ocean floor, were
absolutely devoid of every sign of terrestrial life and without a trace left of the
sial cover. I fail to see that they can solve any biogeographical problems—we
have to fall back on oversea dispersal. WEGENER’s festoons were at least split
off from the borders of a continent and left behind with their fauna and flora. With
regard to Juan Fernandez we shall perhaps be able to find a less adventurous
explanation of its history.

Two years after the appearance of his book, DU ToiT summarized his theo-
ries in a paper which I think it is worth while to quote (82. 75-76). The base-
ment of the Melanesian islands is, he says, for the most part continental; the
ocean floor consists of a relatively thin structure of sial underlain by sima, but
this does not allow us to regard the sial as continental, because it may be a
product of magmatic differentiation from the sima. He points to the parallelism
between the great Tertiary folding-zones, most evident along the west coast of
the Americas, and the trend of the coast line, and he thinks that the “‘compres-
sive phases’’ were contemporaneous all around from New Zealand across Antarc-
tica to South America. Coming back on the advance-folds he remarks that some
of GREGORY’s hypothetical bridges or land-masses could well have been of this
nature. The procedure is illustrated by a map showing the pressure direction
and the formation of island arcs—except on the American side, where the sea
is a blank.

328 C. SKOTTSBERG

Numerous biologists have found a solution of all or most of their difficulties
to explain the present distribution of the organic world in the theory of conti-
nental drift, combined with large-scale pole-wanderings. In view of this the opi-
nions expressed by modern geologists and physiographers cannot be passed in
silence. A symposium, arranged in 1950 (65) offers an opportunity to get ac-

quainted with their attitude.
J. H. T. UMBGROVE, The case for the crust-substratum theory, pp. 67-71.

SuEss’ terms sial and sima were petrographic. WEGENER attributed different
physical properties to these types of rock; sial should be rigid but elastic, sima
viscous. These statements lack foundation. Sima (basalt) has a higher melting-
point, approximately 1300°C, sial (granites) approximately 700°C. The crystal-
line crust of the sima layer is at least as strong as the continental sial. Still the
sial blocks were supposed to advance through the sima. UMBGROVE concludes
that continental drift is impossible at present. This granted, was it perhaps pos-
sible in bygone times? The answer is fetched from the Atlantic and Indian oceans
with the intervening African continent; if the floor of the oceans originated as
thought WEGENER, the processes must have taken place during early Precambrian.
UMBGROVE asks if not the thick blanket of sediments would have been squeezed
and piled up in front when America ploughed westward. He calculates that,
considering the size of the westward drift, a plateau 200 km wide at sea level
would have been formed in front; instead, “‘the continental slope is one of the
steepest in the world and is fronted by deep-sea troughs’’—here we have, how-
ever, to consider the late upheaval of the Andes. And if, as WEGENER’s theory
requires, the Atlantic originated in comparatively recent times, how are we to
explain the enormous thickness of its bottom sediments, according to HANS PET-
TERSSON a maximum of 10000 feet, “‘representing a time-span of 300 to 400
million years’, which would bring us back to the Palaeozoic. If continental drift
ever occurred, UMBGROVE asserts, it took place some 3000 million years ago and
consequently loses every shade of interest to the biogeographer.

HAROLD JEFFREYS, Mechanical aspects of continental drift and alternative theories.

JEFFREYS definitely rejects WEGENEk’s theory on geophysical grounds;
WEGENER gave to the sima properties which the material has not; basic rocks
are stronger than acidic, sima stronger than sial. The strength of the ocean floor
must be overcome, if drift shall result. “I seriously suggest’, he concludes, “that
no more time be spent on discussion of this theory until a mechanism for it is
produced; what it has done, and continues to do, is to distract attention from
the serious problems of geophysics’ (p. 80).

S. W. WOOLDRIDGE, The bearing of Late-tertiary history on vertical and hort-
zontal movements of the continents.

The nature of the mechanism of the “uplift”? may remain in doubt; its reality can
hardly be gainsaid... in so far as such complementary foundering took place during
Tertiary times, it is not directly relevant with the drift problem ... Vertical movements
offer an alternative solution to some ofthe biological problems. Biological evidences

DERIVATION OF THE FLORA AND FAUNA 329

of former facilities for exchange of biota are in themselves neutral in deciding which
alternative should be preferred without paying attention to the ambiguity of much of
the geological evidence and the grave geophysical difficulties (p. 81).

The progress of knowledge of geophysics certainly seems to point to thinking in
terms of land bridges and their coastal margins.

The later findings of seismology, indicating three distinct rock-shells each prob-
ably capable of both vitreous and crystalline states, carry the possibility, as Jeffreys
has clearly insisted, of vertical movements on a major scale. Such strong vertical move-
ments are indeed not merely to be expected; they are widely and unargubly evidenced
by the facts of geomorphology no less than those of stratigraphical geology (p. 82).

R. F. Joyce, Zhe relation of the Scotia Arc to Pangaea, pp. 82-88.

JOYCE made an attempt to reconstruct WEGENER’s Pangaea in Lower Palaeo-
zoic time. To him continental drift is a possibility only, he says that 7f Pangaea
did in fact exist, his arrangement of the actual continents and islands—as usual,
the Pacific island world is not involved—at the opening of the Palaeozoic era is
more in accord with the known data than in Wegener's reconstruction (p. 87).

Two biogeographers took part in the symposium.

H. E. Hinton, The Wegener-Du Toit theory of continental displacement and the
distribution of animals, pp. 74-79.
HINTON rejects the liberal construction of bridges to suit the demands of
specialists. Beside the usual apprehensions he adds the following.

A further objection to the past existence of transoceanic bridges... is the nearly
complete absence of deep-sea deposits on the existing continents, since we would expect
some of the latter to have been also elevated from the sea floor (p. 75).

This seems reasonable; nobody earnestly proposes to fill the surface of the
globe with land, leaving no or little space for the water. Bridges of any consider-
able size cannot have been contemporaneous, and transgression on part of what
is now land seems inevitable, if tracts of ocean floor were exposed. We have
ample proofs that considerable transgressions occurred, but these ancient seas may
have been too shallow to be of much importance as a support of the bridge
hypothesis. HINTON thinks that we require relatively few inter-continental con-
nections, the most important being Brazil—Africa and Australia—Antarctica, but not
in the form of bridges—of such, he admits isthmian and shelf-bridges, nothing
more. Thus, for those who insist on direct inter-continental contact, the only
hypothesis at their service is the drift theory, and he asserts that most of the
modern biogeographers accept the broad outlines of this theory, unless it is claimed
that sliding occurred long before the end of the Mesozoic. As we have seen,
however, this is exactly what has been claimed. For his own part he puts much
faith in chance dispersal across broad stretches of open water.

R. Goon, The distribution of the Flowering Plants in relation to theortes of con-
tinental adrift.

After some remarks on the general distribution of plant families, grasses and
Compositae taking the lead almost everywhere—which seems quite natural in view

330 C. SKOTTSBERG

of the size and enormous climatic amplitude of these families—the preponder-
ance, in nearly all floras of dicotyledons (2 1/, to 4 times as many as the mono-
cots), and the absence of any strongly marked South Pacific flora south of Hawaii
and east of New Guinea, New Caledonia and New Zealand (which ought to show,
I suppose, that the island swarms are not fragments of a larger land mass), GOOD
continues p. 74:

A common expression of the theory of continental drift postulates that the sunder-
ing of various continental masses began in a relatively remote geological period and
that it has continued without notable cessation ever since. The distribution of Angio-

sperms does not seem to be in special accordance with any such particular course of

events. It suggests more strongly that sundering occurred after these plants as a group

had become well diffused, let us say by the Eocene. Similarly I know of nothing in
the distribution of Angiosperms to show that drift is still going on. Can it be that
continental drift has in fact been, not a persistent process, but an intermittent feature
of geological time, as for instance have the great glaciations? If so, and if it can
further be shown that the most recent of these drift ages took place not earlier than
the beginning of the Tertiary, then at last the plantgeographers will have to hand a
master-key to most of his perplexities.

It will be difficult to shape a key that will fit a lock constructed in this fashion.
As the glacial periods were interrupted by interglacial periods, so the periods
of sundering should have been interrupted by periods not of a standstill but of
the land-masses coming into contact again, last time during the Jurassic-Creta-
ceous era, which saw the origin, evolution and dispersal over the globe of the
Angiosperms, to be followed by the last drift age.

With regard to the relations and disjunctions between South America (with
Juan Fernandez, etc.) and Antarctica, Australia and New Zealand, vertical move-
ments seem to offer a less distant possibility.

In a most cleverly written chapter Goop (z09. 344-360) discussed land-
bridge versus continental drift. If we cannot, he says, accept the former, nor
put our trust in dispersal, a changing position of the continents is the only way
out of the difficulties. This may be true, but we have seen that this theory does
not help us to solve the problem concerning the oceanic islands. GORDON (z73)
points out that the occurrence of a small but important subantarctic element in
the Pacific, reaching north to the Hawaiian Islands where it is better displayed
than in any of the Polynesian or Melanesian groups, makes it impossible to deny
both land-bridges and the efficiency of transoceanic dispersal without providing
for migration with the help of shifting continents. GOOD

has run into an impasse over the Pacific islands like Hawaii. He has rejected the land-
bridge hypothesis in favour of continental drift.... But he excludes continental drift so
far as the islands are concerned, for he accepts them as truly oceanic, not continental frag-
ments. Yet he will not accept overseas migration. Well, I can’t see what explanation
remains, if all these three are excluded, but the plants are there (p. 148).

WULFF (297) found that the biogeographers have good reason to support
WEGENER’s theory; much speaks against it, but he trusts that the difficulties
will be overcome by and by. He recognized that, with regard to the Pacific,
they are very considerable and call for a modification of the theory.

DERIVATION OF THE FLORA AND FAUNA 331

Chapter VI.

Transoceanic migration.

In some of my earlier papers I touched upon the great problem of long-
distance dispersal and the supposed efficiency of the transporting agents; see for
instance 237z.20-30, where, however, only the flora was concerned. This time
also the faunas are, to some extent at least, considered, and I shall quote a num-
ber of authors, old and modern, who have expressed their opinion for or against
overseas transport as the only possible means by which the isolated islands of the
Pacific have received their indigenous flora and fauna.

Advocates of large-scale overseas migration.

Among earlier authors ENGLER (85) and GRISEBACH exercised great influence
on their contemporaries. They divided the world into flora domains, regions and
districts, characterized by a combination of certain important elements and by a
greater or lesser degree of endemism, but still they never doubted the facility with
which plants travelled across the oceans; progressive endemism was the unavoid-
able corollary, but relict endemism was recognized as important. The attitude of
this school is adequately expressed by GRISEBACH (325. 469):

So merkwiirdig es auch sein mag, dass sogar einzelne Holzgewiachse sich hier
iiber das Stille Meer verbreitet haben, so ist ihre Wanderung doch aus der antarktischen
MeeresstrOmung, den herrschenden Westwinden, oder durch Mithilfe der Seevégel,
vielleicht auch durch alte Verkehrswege wohl hinlanglich zu erkléren, ohne dass die
Annahme von Landverbindungen in der Vorwelt gerechtfertigt ware, die durch keine
geologische ‘Tatsache gestiitzt wird.

By some the case of Krakatau was quoted as a proof that plants and animals
are able to travel across water barriers; it is mentioned by HAYEK (304), but
with reservation:

Wie die Besiedelung einer Insel erfolgt, haben die oben angefiihrten Beobachtungen
bei der Wiederbesiedelung des Krakatau gelehrt... Aber die Entfernung des Krakatau
von den niichst gelegenen Inseln ist keine allzugrosse, sie betragt nur etwa 18—40 Km,
also Entfernungen die auch durch die Flugtiere nicht allzuschwer tiberbriickt werden
konnen (p. 251).

When we have to deal with islands, separated by thousands of miles from
all continents, the difficulties are of much greater magnitude, and he continues:

Und doch miissen wir-annehmen, dass auch die weit entfernt gelegenen Inseln ihre
Pflanzendecke von den zunichst gelegenen Festlaéndern (und Inseln) erhalten haben,
wenn auch vor undenklichen Zeiten und ganz allmihlich. Dafiir spricht auch der Um-
stand, dass die Flora dieser Inseln keineswegs von der iibrigen Flora der Erde grund-
verschieden ist,, sondern denselben Pflanzenfamilien angehort wie diese, demnach von
derselben abstammen muss.

The only exception known (at that time) was Juan Fernandez (Lactoridaceae).
But he admits that there are grave difficulties:

332 C. SKOTTSBERG

Selbst bei Inseln, die erst in relativ junger Zeit vom Festlande abgetrennt worden
sind, ist ein weiterer Austausch der Florenelemente zum mindesten wesentlich erschwert
und eine weitere Zuwanderung von Elementen der Festlandsflora wenig wahrscheinlich.

If this be true, how was immigration over thousands of miles ever possible?

Several writers who have paid special attention to Pacific problems occupy,
more or less dogmatically, the same standpoint as HAYEK. SETCHELL, with whom
I had the privilege to discuss this subject on various occasions, was already quoted
p. 271; I shall add here what he says, in the same paper, about migration (279. 300).
He found that I was “too narrow” in my allowances for migration possibilities;
he believed in “migration over very considerable breadth of barrier, whether of
sea or land’, and absence was not a result of failure to migrate successfully, but
could be explained by obstacles to establishment. He regarded the oceanic islands
as Tertiary, but in his summary pp. 307-309 admitted the possibility of their
being considerably older, late Mesozoic or early to middle Tertiary, which would
give time for extensive progressive evolution of endemic taxa and for the dying-
out of their continental ancestors; or they had developed in other directions,
making the relationships difficult or impossible to recognize. Geologists, however,
refuse to give even the Hawaiian Islands a greater age than late Tertiary or even
Pleistocene. It goes without saying that travel facilities are different in different
cases; spore-plants are supposed to spread more freely than seed-plants, but even
these are supposed to be quite capable. Thus STEBBINS (379. 537):

The seeds of plants may occasionally be transported over many hundreds of miles
of ocean and may establish themselves on Oceanic islands like Hawaii, Juan Fernandez,
St. Helena and the Canary Islands.

FLORIN has, he writes, shown that conifers of the south hemisphere have migrated
freely from Australasia to South America and vice versa, whereas mammals are
unable to pass and are absent from oceanic islands—but is it not customary to
place them on a par? STEBBINS’ Antarctic connection does not include land-bridges,
for “it existed for plants, but not for vertebrates’ (but what about birds?). He
looks for assistance in lost islands between Antarctica and New Zealand; on the
opposite side the width of open water is not so great, and seeds can still be carried
from South America to Antarctica without much difficulty.

As mentioned before, no botanist has greater confidence in long-distance dis-
persal than FOSBERG:

. . transoceanic migration across at least 2500 miles without stepping-stones is not
only a possibility but a relatively common occurrence (99. 867).

FOSBERG's subject was the American element in the Hawaiian flora, but in order
to explain the presence of the dominant Australasian element we must count with
still greater distances. AXELROD (rg) quoting FOSBERG takes a modified position.
In case of distances not exceeding some 200 or 300 miles there are no difficulties,
‘‘a complete flora can transgress such a barrier without the loss of any significant
floristic units’, A greater distance results in “waif assemblages’, but many will
find it impossible to regard e.g. the Hawaiian flora as a haphazard accumulation

DERIVATION OF THE FLORA AND FAUNA 333

of waifs and their descendants. What AXELROD says about migration probabilities
during different geological epochs is of greater interest.

Since plants are controlled largely by climate, and since climate has been changing
during geologic time, it follows that plants comprising different communities have had
different possibilities at different times . . . probabilities for long-distance migration were
much higher for tropical plants in the Eocene than they are to-day. Temperate forest
species had a much higher probability from late Cretaceous to middle Tertiary, it is
low now. Steppe plants had a higher probability during Pliocene than now. Desert spe-
cies have a higher probability to-day than at any time before.

His conclusions are drawn from the size and area of populations shifting with the
extension of climatic regions.

For my own part I have expressed my opinion on overseas migration in the
Pacific on various occasions (237, 378, 248) and I am not going to repeat the dis-
cussion here. My general conclusion was that the effect of transoceanic migration
has been largely overestimated.

Guppy (227), who allowed birds, winds and currents to stock all oceanic
islands with plants, arrived at the conclusion that this traffic was a thing of the
past and that migration had practically ceased altogether. 1 expressed my doubts
that it had ever been effective, in any case with regard to seed-plants.

I have already remarked that SETCHELL laid stress upon what he called the
CEB (climatic-edaphic-biotic) factor complex. The main difficulty for the vagabond
plants was not to cover the distance, be it ever so great, but to become a successful
member of a community already established in the place where it happened to
alight, and this difficulty increased as time went by; most surfaces of the earth,
he says, are already stocked with closed vegetation, making it impossible for new
arrivals to gain a foothold (279. 300). His ideas are clearly expressed in 278 (p. 874).

As the islands have become more and more completely stocked each with its quota
of plants and animals and have undergone various vicissitudes, particularly of elevation,
erosion, etc. its hospitality to migrating germules necessarily has become less and less,
the Biotic factor has become more complex and the Edaphic factor has also suffered change.

In my view the result could just as well be the opposite, for these “‘vicis-
situdes’’, emergence, erosion, volcanic activity and so forth create new soil, a more
varied topography, a multitude of different habitats, all of which ought to give
newcomers increased opportunities to get established.

With SETCHELL, ANDREWS underlines the importance of CEB; genera
expected to occur in Hawaii but absent “‘were not amenable to germination
and survival after transport’’.

Long before SETCHELL, J. D. HOOKER, WALLACE and others had paid
attention to the obstacles for the successful establishment of newcomers, WALLACE
believed that St. Helena had become stocked with plants during early Tertiary
time; later there was no room left, and the flora had changed so completely that
no plant was recognized as an insular form of a continental species.

We meet with SETCHELL’s line of thinking in a recent paper by W. B. TAYLOR
(263.572). In recent volcanic islands are many unstocked habitats to begin with,

334 C. SKOTTSBERG

but each new species would mean competition, and the entry of an additional
species would be very difficult and consequently of rare occurrence. This may
be so, but there are many communities of a more open character than the forest,
and even a closed forest is not like a tin packed with cigarettes; young secondary
forest associations, steppes and savannas ought to offer good housing grounds
to an intruder, supposing that he likes climate and soil. Experience shows that
numerous aggressive plants brought by man, not only herbs but also trees and
shrubs, find suitable living conditions even in undisturbed natural communities.
In Juan Fernandez I have had occasion to follow the invasion by Avzsvotelia maquet
(chilensis) and to witness the fabulous ease with which it crowds out the native
vegetation, and to observe Ugni Molinae springing up on the ridges where the
plant cover was open and, from there, to enter the dense native brushwood.
And they are only two of the many successful weeds, a third one, equally dan-
eerous but of quite recent introduction, is Rubus ulmifolius. All three have fleshy
fruits and are eaten by man and birds and propagate themselves rapidly. They
are common on the opposite mainland (where, of course, the brambleberry was
introduced from Europe), but man, no bird, carried them across to the islands.
Similar examples are, I presume, offered by almost all oceanic islands. I just hap-
pened to read a book on Cape Verde Islands, where a naturalist tells us about
Lantana camara spreading like wildfire and menacing the little there is left of
natural savanna and steppe (320).

Most zoologists favour the theory of long-distance dispersal. Mammals are,
as a rule also bats, flying foxes and the like, excluded, but of birds some are able
to cover very large distances, winged insects are carried off to distant places
where they never wanted to go, and so forth. I shall quote some zoogeographers
who, with reference to the Pacific, have expressed their opinion on overseas transport.

PERKINS, in his introduction to Fauna Hawaiiensis (793. XLVI), wrote:

All the islands being volcanic and having been built up from a great depth of ocean
at various periods, their entire fauna naturally originated from immigrants derived from
other lands. These immigrants must have arrived either by flight, like the birds, or in
drift like the flightless insects and probably the land Mollusca.

Drifting logs were often regarded as an important kind of conveyance, but they
come from North America and what they bring of animals, PERKINS says, would
serve no purpose because it is unlikely that the passengers would become acclim-
atized in Hawaii. The fauna must have come from the warmer parts of America,
from Australia, Polynesia etc. “at rare intervals from the Eocene until now’. If
we have to believe the geologists, no Hawaii existed in the Eocene—and how
did those, who arrive “now’’, manage to become endemic genera and species ?—
non-endemic flowering plants not brought by man are few.

In some instances GULICK (779) admits the possibility of land connections,
but Hawaii, Juan Fernandez, Galapagos, St. Helena, etc. etc., have always been
isolated. The question whether their fauna shows that “the ancestors possessed
an almost inconceivable capacity for passing uninjured over vast stretches of ocean”’
is answered in the affirmative. It is significant that the Galdpagos archipelago

DERIVATION OF THE FLORA AND FAUNA 335

“‘was successfully reached by a giant tortoise’; already WALLACE entertained the
same idea

a strictly terrestrial animal crossing the ocean.

Some biogeographers prefer one dispersal agent, some another, most have
confidence in all, but it goes without saying that different types of plants and
animals have availed themselves of different kinds of transport. I shall quote
GULICK (rzg. 414) first.

It is possible to go far toward a first diagnosis of the degree of a land’s insularity by
noting how exclusively itis peopled by types with a known capacity for colonizing across
vast expanses of ocean. Our summary up to this point reveals very nearly which these
forms may be. Quite a majority of them, both plants and animals, show characters that har-
monize with wind-storm transportation. A respectable majority of the larger-seeded palms
and some tough-lived earth-inhabiting invertebrates, suggest transportation by water or on
drift-wood. Such seeds and invertebrate eggs as can withstand the digestive tracts of a bird,
have a very substantial travelling radius by that means, easy 500 miles in the routine
seasonal migrations, and possibly stretching in the extremest cases to almost transoceanic
distances. ... Dioecious plants and separate-sexed animals are statistically at a disadvan-
tage, as compared to the reversed condition, because of their poorer chance of achieving
fertilization. The ability to take a journey in a gravid condition helps the chances greatly,

“Types with a known capacity for colonizing’—GULICK proceeds from
what should be proved, for their occurrence on isolated islands is in itself no
proof of oceanity. Under his angle the great number of dioecious endemic
phanerogams in Hawaii ought to have surprised him. It almost seems as if he
believed that entire specimens with roots and all managed to reach a distant
island and get established; surely, if only a male or an unfertilized female arrived;
all was in vain until a mate of the opposite sex turned up; a pregnant female
would of course do better (bye the bye, WALLACE tells a story of a pregnant
boa constrictor arriving on a West Indian island with drift-wood and in good
condition). I guess we can leave these chances aside, for plants spread by
means of seeds, and whether wind-blown, epizoic or endozoic (provided they
do not, as many assert, discharge their droppings soon after the departure),
there is every chance that more than one seed of the same kind is brought; a
single many-seeded berry is enough, and a bird picking drupes fills his stomach.
A seed portion of a dioecious species gives, under ordinary conditions, 50 % of
each sex. In the Hawaiian flora we find, GULICK says (p. 418), “a preponderance
of plants spread by wind-carried spores and minute seeds’; species with drupes
and berries are, however, numerous. As an example of a presumably definite
case of bird rather than wind carriage he mentions the Hawaiian species of
Vaccinium, which he derives from North America. Their presence is most in-
teresting, ““as the distances involved must be very close to the extreme physio-
logical maximum that land birds can traverse, and still carry fruit seeds in their
droppings’. To me it appears as a bad case of constipation. Besides, the Ha-
waiian Vaccinia are not related to North, American groups but belong to a
special section.

MAYR, an extreme ‘‘oceanist’’ who refuses to admit land connections for
either Fiji or New Caledonia, in his paper on the Pacific bird fauna (779) includes

330 Cc. SKOTTSBERG

a general survey of the dispersal chances for other animals and also for plants.
The special instance cited (p. 197) is not very convincing.

Birds are excellent flyers and thus capable of rapid and active spreading .. . capable
of crossing considerable stretches of open sea to settle in new territories. ‘There is abun-
dant evidence of this, such as the resettlement of Krakatau Island, the recent arrival of
Australian birds in New Zealand, and the colonization of unquestionably oceanic islands.

Not even the arrival of Australian birds in New Zealand brings conviction;
the colonization of “unquestionably oceanic islands’ certainly does, if we can
prove it. MAYR continues p. 198:

The possibility of transport by floats or in logs is not to be underestimated. Many
tropical currents have a speed of at least 2 knots, that 1s, about 50 miles a day, or 1000
miles in three weeks. It is probably not a great task for a wood-boring insect to survive
3 weeks in a drifting log. Air currents are, however, of uncomparably greater importance
than sea currents. Even slight winds are of great influence on the distribution of floating
and flying animals, as recent investigations have shown. It is astonishing how rich the
“aerial plankton”’ is, even up to altitudes of 1000 meters and more. Normal winds would,
of course, not account for the spreading of molluscs, flightless insects, and other small in-
vertebrates. However, most of the islands, with which we are concerned, are situated
within the zone of tropical hurricanes, the lifting force of which is quite extraordinary... .

The fact that there are small molluscs and flightless insects on such typical oceanic
islands as Easter Island, Juan Fernandez and Saint Helena is almost unassailable proof
that such a method of dispersal is a reality. Tropical hurricanes carry for hundreds and
even thousands of miles.... The result of the recent surveys im the Hawaiian Islands,
the Marquesas, and on Samoa indicate that there are indeed very few animals that
cannot be transported across considerable stretches of the sea by winds, waves, other
animals or man.

On p. 201 MAyR adds some general remarks:

The means of dispersal of most plants and animals are much more extensive than
was formerly realized, and even rather irregular distributions can be explained without
the help of land bridges. Dispersal across the sea is, of course, most obvious for birds,
and ornithologists were among the first who accepted the ideas of the permanency of
continents and oceans. Most entomologists are also beginning to realize that they can
solve most of their distribution difficulties without land bridges. The conchologist,
however, postulates even today continental connections between all or nearly all the is-
lands where land shells exist.

As we shall see below (p. 350) MAyr declared himself unable to explain how
land shells are dispersed.

It seems that, also with regard to the birds, MAYR contradicts himself, for in
the same paper (p. 198) he asserts that most birds, particularly on tropical islands,
precisely the islands we are discussing, are extraordinarily sedentary, and as an
example he mentions that of 265 species known from that part of New Guinea
which is opposite New Britain, a distance of 45 miles, only 80 occur on New
Britain, and the situation in Western Papuan islands is even more conspicuous;
he mentions two islands only 2 miles apart, with rather different fauna. “Literally
hundreds of similar instances could be listed ... all of them indicating the sedentary
habits.” One is likely to remember Guppy’s fruit-eating pigeons which were
thought to be responsible for the dissemination across the Pacific of seeds too
large for other birds. The “pigeons” are, now at least, restricted in range and of

DERIVATION OF THE FLORA AND FAUNA 337

very little use on longer distances. I cannot help drawing the conclusion from
this that the sedentary habit was acquired after the great colonization had taken place.
Few phytogeographers have had greater faith in the capacity of wide-ranging
marine birds to carry diaspores than GRISEBACH. In his discussion of bipolar
species found in the far north and the far south but not at all in intermediate
zones he selected Genteana prostrata Haenke as the best example. Its distribu-
tion is due, he says, to the wanderings of Dzomedea exulans which,
abweichend von der Lebensweise der meisten anderen Zugvégel, iiber beide Hemisphiren,
von Kap Horn bis zu den Kurilen und Kamtschatka, wandert und die Standorte jener
Pflanze in der arktischen und antarktischen Flora in Verbindung setzt. Mit der Beute,
die dieser Vogel verschlingt, kann er auch Samen von Pflanzen, welche, mit den Fliissen
ins Meer gespiilt, in den Magen der Fische iibergehen, in einzelnen Fiillen ausstreuen,
so dass sie an fernen Kiisten aus seinem Diinger aufkeimen (325. 460).

I have not come across any comments on this bold theory. It is difficult
to take it seriously, but to GRISEBACH the only gap in his argumentation was
that nobody had happened to witness such an event. If he is wrong, he asks,
why is there no trace of this Gezézana in the Andes, where it would thrive just
as well as in the Alps and in the mountains of Asia? To this should be re-
marked that G. prostrata is a polymorphous species of wide range and that it
does occur in the Andes from Colombia to Chile, suggesting that it has mi-
grated south along the mountains without the assistance of the albatross.

WALLACE (278. 259) tells us, on the authority of MOSELEY, naturalist to the
“Challenger” expedition, of the great albatross breeding on Marion Island in the
midst of dense, low herbage; I can add that this bird also breeds on South
Georgia and on some other southern islands, but as far as I know they do not
shift breeding places, and even if they did, they do not go on shore between
the breeding seasons. TAYLOR, in his important paper on Macquarie Island (263),
tells us about a giant petrel which was captured, tagged and released on this
island and shot on South Georgia, 8000 km away, four months later, but these
birds are often seen on land where they attack the penguin chickens; this was
at least the case on Paulet Island in the Antarctic. Whether they aid in the
dispersal of diaspores is unknown.' TAYLOR quotes an observation, made on
Macquarie, that seeds were found adhering to the feet of an albatross. These
birds, when building their nests, regurgitate an oily fluid which makes seeds stick
to their feet. Macquarie Island was ice-covered during the Glacial epoch and the
plants, perhaps with the exception of some cryptogams, must have arrived since
the retreat of the ice. The vascular flora consists of 35 species, all except 4
occurring in the New Zealand subantarctic area—the 3 species with a claim
to be regarded as endemic should be reinvestigated—while those 4 species are
found in subantarctic South America, from where they are derived. All Mac-

1 According to TAYLOR (p. 570) the two truly Antarctic phanerogams, Deschampsia ant-
arctica and Colobanthus crasstfolius, are very rare in the Antarctic and reproduce only vegeta-
tively. I do not know where he obtained this information. They are scarce but have been
reported from many localities along the coast and adjacent islands of Palmer Land between
lat. 62 and 68 and, in favourable situations at least, both of them flower and produce ripe
seeds—see my paper in Botan. Tidsskrift vol. 51, 1954.

22 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

338 C. SKOTTSBERG

quarie plants have, TAYLOR states, propagules suited to bird transport. It is sur-
prising that WERTH (377), who made a detailed study of the Kerguelen flora,
asserts that not one of the flowering plants possesses any special dispersal mech-
anism for either wind, water or bird carriage. Still, the two islands have some 9
species in common (54). TAYLor’s conclusion that, “if long-distance dispersal
has occurred on Macquarie Island, then it could well have occurred elsewhere”
is certainly correct; we know, for one thing, many wide-spread sea-side plants
and a number of widely dispersed aquatic species, possibly transported by
migratory birds. South Georgia is in much the same situation as Macquarie,
but still rather heavily glaciated thanks to its great altitude, and the possibility
that any higher plants survived the Glacial epoch is very small indeed, whereas
indications that many mosses and lichens date from preglacial times are strong.
The vascular flora is poorer than on Macquarie, and there are no endemic species.
When TAYLOR accuses me of having argued against all overseas migration also
n this case he must have misunderstood me. I expressly took this possibility
into account in the paper he quotes (226).

WALLACE calls attention to sea birds breeding on islands in the tropics;
Phaeton makes its nests on the Hawaiian Islands in 4000 ft. altitude and also in
the highland of Tahiti, and such birds would account for the similarity of the
mountain floras. In reality these floras have practically nothing in common.
Miss GIBBS (327), discussing the origin of the montane flora of Fiji, refused to
regard birds as capable agents; wind may have been more efficient.

No modern zoologist has tried to defend the theory of unlimited overseas
migration with greater zeal than ZIMMERMAN. In his Introduction to “Insects of
Hawaii” we read:

There is no evidence whatsoever to support the contention that they (i.e. the Ha-
waiian Is.) are of continental origin or character, or that they were ever joined together
in an elongate subcontinental land-mass or even in a continuous subaérial mountain
range (298. 6).

And, in opposition to certain other biologists he refuses to regard the islands as
old, they are at most Pliocene and no part of them older than five million years;
most of the lava is younger, the bulk of the land Pleistocene. He is opposed to
my ideas but he thinks that the explanation offered by him will, partially at least,
reconcile the differences between us. In an earlier paper (Amer. Naturalist 76,
1942), to which he refers, he spoke of former high islands, other than those found
on maps, which once existed; once more the “‘stepping-stones routes” are called
to life. Atolls are the remnant of many of them, or reefs like among the Leeward
islands of Hawaii, and such preexisting islands would account for the immigration
from all directions. He does not call for jumping of thousands of miles of open
sea, but rather for series of shorter over-water steps. I am afraid that we need
some substantial refurnishing of the Pacific basin to supply a sufficient number
of intermediate stations. Not all of these routes were, he says, available at the
same time, and this would explain the apparent difference in age of various
sections of the biota (pp. 51, 52). Most of the roads were cut off in Pliocene and

DERIVATION OF THE FLORA AND FAUNA 339

early Pleistocene, some before Pliocene—when no land existed where stands the
Hawaiian chain, if geologists are right. USINGER (273), describing the distribu-
tion of Heteroptera, also looks for convenient stepping-stones.

Divergent opinions on the means of transport.

The dispersal agents universally recognized as important are air currents,
especially monsoon and trade winds, and cyclonic storms, ocean currents, birds,
and last not least man, who with his domestic animals and goods has become
more and more important, whether he brings plants, seeds and animals to extend

their range—and many of them become naturalized—or carries diaspores on his
body and his belongings unaware. It is as a rule not difficult to find out where
we have to do with human action, but we shall limit ourselves here to a discus-
sion of natural factors of distribution. This subject has been treated by innumer-
able writers in biology and a wealth of material was compiled by RIDLEy (205),
a firm believer in the great value of all kinds of dispersal mechanisms, some of
which are, of course, very wonderful. WULFF (297), in bis chapter ‘“‘Natural factors
for distribution” is more critical. Of animals only the birds deserve to be men-
tioned, but the plumage is no good for carrying diaspores any considerable dis-
tance, especially over the sea, and the extreme marine birds, the strongest flyers,
have no contact with land outside their own breeding-places. WULFF reduces the
part taken by birds to almost nothing, but I believe that we have good reason
to count with the migratory birds in certain cases. Water transport is responsible
for the diffusion of litoral halophytes but rarely for migrations of inland species.
Wind is important only on short distances, at least for seed-plants; special devices
do not help very much. Even RIDLEY concluded that winged or plumed diaspores
are not carried very far; spore-plants are more easily spread. WULFF remarks
that according to BENTHAM Leguminosae and Labiatae hold their ground just
as well if not better than Compositae. Altogether, if dispersal by natural factors
had the significance ascribed to it, the vegetation of the globe, within a certain
climatic zone, would be homogeneous and the sporeplants at least ought to be
cosmopolitan, but they are “localized in definite areas, their distribution paralleling
that of flowering plants” (p. 128). Goop observes (zog) that we have no proofs
that species equipped with special dispersal mechanisms are more widely distri-
buted than others. That certain plants with such devices show very wide areas
whereas others without them are rare and local means nothing, because the reverse
is also true; climatic and edaphic factors should always be taken into account.

The relative value of the dispersal agents is put to the test when we deal
with oceanic islands. SETCHELL (277) was inclined to give considerable credit to
migratory birds. The occurrence of identical species of flowering plants in Arizona
and Argentina and in California and Chile could be explained by bird transport,
and bipolarity had originated in the same way. Bird transport helped him to
understand the remarkable disjunct areas of arborescent Compositae and Lobe-
liaceae; their birth place was in the Antarctic, and they had been carried by
birds to New Zealand, Australia, Malaysia, Polynesia, Hawaii, South America and
the high African mountains—we can add Saint Helena, Juan Fernandez and Des-

340 C. SKOTTSBERG

venturadas. In this case it is not the question of identical species and rarely of
genera. It is not improbable that the secret of their origin and early history lies
hidden in the far south, but this is all we can say. HEMSLEY (727.66) when dis-
cussing the endemic Compositae of Saint Helena and Juan Fernandez said that
‘wind seems at first to be the most probable agent”; still he doubted its efficiency.
In SETCHELL’s view storms seem to offer more than a possibility in many in-
stances, particularly tropical cyclones and vertical thrombs able to carry even
heavy diaspores to a great altitude. In another paper (278) he points out that
plant distribution in the Pacific has been from west to east against the prevailing
winds and currents and, in the case of Hawaii, has given much better results
than the expected east-west route; the ‘frequent cyclonic storms” are responsible
for this anomaly together with adverse biotic factors, but I fail to see why they
shouldn't offer the same difficulties for diaspores coming from the west.
ANDREWS (6.615) paid special attention to the occurrence of scattered “‘Ant-
arctic’ genera and species in and around the Pacific and combines their distri-

bution with the direction of ocean currents:

In the South Pacific the westerly current sweeps by Australia, New Zealand, Tahiti,
and the whole of the west coast of South America, where it 1s joined by the cold
uprising water along the South American coast. This gives rise to the north-moving
Peruvian Current which sweeps by Juan Fernandez, Peru, Central America, and Mexico,
whence there is a deflection westerly toward the Hawaiian Islands and the tropics.
This knowledge of the general circulation within the Pacific appears to throw a flood
of light on the occurrence of the Australian, New Zealand and western South American
elements in the Hawaiian flora, such as...(25 genera are enumerated). The influence
of ocean currents is suggested particularly in the peculhar distribution of Acaena, Gunnera,
Nertera, Oreobolus, Santalum, Sophora, and so on.

Juan Fernandez lies outside the Peruvian current, but also the outer island, Masa-
fuera, is reached by drift-wood. Its origin has not been investigated.

A look at a current-chart shows that ANDREWS’ reasoning has its weak
points; besides, I cannot see that the genera he mentions are thalassochorous. |
would recommend the reader to take a look at the many distribution maps pub-
lished in a paper read on the same occasion when ANDREWS presented his argu-
mentation (373).

With regard to the cyclonic storms several authors have, as we have seen,
emphasized their prominent rdle in the violent dispersal of both plants and ani-
mals; they are, in fact, considered to be the only imaginable force by which
larger objects are transported, and it is useless to deny that such events have
taken place and still take place, even if it is difficult to find definite proofs that
the transport did lead to the establishment of an immigrant from*afar. Most
authors who have taken refuge in cyclones have, however, expressed themselves
in general terms without a clear idea of the extension of the cyclonic belts and
the trend of the cyclones.

In two papers VISHER has summarized his studies on cyclonic storms in the
Pacific. Three chief centres of origin are distinguished (322): (1) Western N. Pacific,
originating some distance east of the Philippines in lat. 8° to 25°; (2) Western

DERIVATION OF THE FLORA AND FAUNA 341

S. Pacific, particularly between Australia and Samoa in lat. 10° to 25°; (3) Eastern
N. Pacific off the west coast of Mexico and Central America. Occasionally tropical
storms develop near Hawaii and over Australia. The normal course of (1) is WNW,
recurring NE, and of (2) WSW, recurring SE. To what extent plant distribution
runs parallel to cyclone tracks has, as far as | am aware, not been investigated.
As VISHER says, most of the cyclones originate over the sea “well out in the
ocean” (p. 87). They hit many of the Polynesian islands, and possibly collect
diaspores on one and deliver them on another, but a frequent dispersal of species
in this way does not appear very probable. I wonder whether there is in Hawaii,
with its 90% endemics, a single flowering plant likely to have been borne there
by a cyclonic storm. VISHER is, however, opposed to land connections in the
Pacific, with one exception: “it is known that Australia was formerly connected
with Asia by way of the East Indies and New Caledonia” (323.77). With regard
to Polynesia he points to the west-east hurricanes and their colonizing power.
Sea carriage also comes into the picture, violent cloudbursts may accompany the
storm, brooks are transformed into swift rivers carrying plant material, tree trunks
and soil, forming rafts, which are washed out into the ocean.

A recent paper by BERGERON (324) gives a somewhat different aspect. His
map shows the two areas in the Pacific north and south of the equator, where
hurricanes arise, and how they move. The Hawaiian islands lie, as a rule, outside
the tracks. The direction north of the equator is NW or WNW all through the
hurricane belt, and there is no sign of an easterly direction enabling plants and
animals to be carried all over Polynesia as far north and east as Hawaii, as VISHER
supposed. South of the equator the trend is S and SE; the east of the Pacific
is not reached and Juan Fernandez lies, in longitude as well as in latitude, away
from any cyclonic belt.

It has often been stated that spore-plants, theoretically at least, have much
greater facilities to colonize on long distances, but it has been shown that in
reality their capacity is more limited than was formerly assumed.

COPELAND (68. 165-166) expressed his opinion on the diffusion of fern spores:

Fern spores are carried across water by the wind—ten miles of water is no barrier
at all to their spread. One hundred miles may be one hundred times as great a barrier,
because the spores must hit a target, a suitable place to germinate and grow. Still,
ferns spread readily across seas this wide. A thousand miles makes the obstacle again
one hundred times as great... the limited viability of the spores, the chance of falling
or being washed out of the air, and the chance of very different climate at such dis-
tance, increase it materially. Ferns rarely jump a thousand miles of ocean. Still a number
of species are believed to have crossed the south Atlantic, and I believe that three
‘genera, Plagiogyria, Coniogramme and Loxoegramme, have flown the north Pacific trom
Japan to Mexico, each in one single instance. It is not exactly impossible that direct
colonization has occurred between Chile, New Zealand, Tasmania, the Cape, and Tristan
da Cunha.

It should be mentioned that CHRIST (59), with his unique knowledge of the
distribution of ferns, pointed to the insignificance of spore dispersal as an
explanation of the origin of widely disjunct areas.

342 C. SKOTTSBERG

The fact that IRMSCHER tried to prove that the distribution of plants strongly
supports WEGENER's hypothesis does not lessen the value of what he says about
the limited capacity of plants to migrate. It is small in the flowering plants (743. 291):

Dass bei den Bliitenpflanzen die Beforderung der Samen und Friichte durch Wind,
Wasser und Tiere ganz wesentlich eingeschrénkt werden muss und fiir geschlossene
Formationen auf grossen Entfernungen hin nicht in Frage kommt, ist heute von den
Pflanzengeographen allgemein anerkannt.

IRMSCHER is too optimistic, we have seen that there are phytogeographers
to whom overseas migration is not only possible and undoubtedly happens, on
rare occasions at least, but rather of quite common occurrence. And with regard
to spore-plants their distribution should, if this be correct, show quite different
distribution patterns than they actually do. The bryophytes are no exception to
the rule. IRMSCHER remarks that already in 1903 STEPHANI denied that liverworts
are able to make long and successful jumps. Attention should be paid to DOMIN’s
valuable paper (76) in which he brings together numerous facts illustrating the same
definite distribution patterns in this as in other groups, and Miss FULFORD (e.g.
103) has arrived at the same conclusion. We know that the spores, in many cases
at least, are extremely sensitive to changed conditions and lose their viability
very rapidly when exposed to the air—a promising field for experiments. Regard-
ing mosses I refer to HERZOG’s work (729) where he speaks against the belief in
the importance and great range of dispersal through the air. We find, IRMSCHER
says, the same disjunctions, the same part areas (Teilareale) in angiosperms,
gymnosperms, ferns and bryophytes, and he continues p. 292:

Dass diese vier in ihren Verbreitungsmitteln so verschiedenen Pflanzengruppen die-
selben Verbreitungsziige ihrer Disjunktelemente ergeben, zeigt woh] einwandfrei, dass
hierfiir der “Wind” ebenso wie andere dussere Krifte als Ursache abzulehnen sind.
Waren sie in ausschlaggebender Weise an der Ausbreitung beteiligt, miisste die Besie-
delung entsprechend der Verschiedenheit der Friichte, Samen und Sporen ebenso ver-
schiedenartig ausgefallen sein, d.h. in den einzelnen Gruppen dieser biologischen Ver-
schiedenheit entsprechende charakteristische Merkmale zeigen. Dies ist aber nicht der
Fall. Der allen vier Gruppen gemeinsamen hochdisjunkten Ausbildung so vieler Areale
muss vielmehr eine andere Ursache zu Grunde liegen. .

This common cause was, in IRMSCHER’s opinion, continental displacement in the
sense of WEGENER. Those who disagree with him will have to look for vertical
movements, emergence and submergence of land.

Lichens, fungi etc. were not included in IRMSCHER'’s discussion. A survey of
their distribution patterns is something to be asked for. Lichens are said to de-
pend on their vegetative reproduction bodies more than on spores.

It is maintained that, with certain exceptions, terrestrial animals spread
less easily than plants. I have consulted a number of zoogeographers in order to
learn their opinion on the mode of transport likely to be used by invertebrates
in their supposed ocean voyages. Birds etc. are left aside here.

Numerous insects, butterflies, moths, flies, hymenoptera, grashoppers, cock-
roaches, as well as spiders, myriapods, etc. follow man from land to land, from
island to island. This is, I daresay, the only safe way for such animals to get

DERIVATION OF THE FLORA AND FAUNA 343

abroad. For those, and they are in overwhelming majority, which are not an-
thropochorous, the chance to cover large distances ought to be very small. How-
ever, ZIMMERMAN, in his admirable introduction to the Insects of Hawaii, surveys
one order after the other and finds nothing that speaks against his belief in the
permanent isolation of the Hawaiian as well as all other oceanic islands, and
consequently concludes that, man-borne species excepted, all the ancestors of the
Hawaiian insects were carried there by natural agents. Dispersal with the aid of
birds is of slight importance, but he remarks that sea birds nest in forests on
the islands; I do not think that this means very much, because they are stationary
(comp. above, p. 337). Marine drift is probably the least successful of all meth-
ods, he thinks. Thus, the bulk of the insect fauna was and is wind-borne, a
traffic going on without interruption. ZIMMERMAN refers to experiments clearly
showing that both winged and unwinged insects are carried by air currents to
great heights, 14,000-15,000 feet (p. 58). These are largely abnormal conditions
and due to cyclonic storms, which account for the dispersal all over the mid-
Pacific. The result is, as expected, a disharmonious fauna, where large groups
common to all continents are lacking: “‘they have been eliminated by the selective
agents of oversea dispersal’. It would be interesting to know why all represent-
atives of large and otherwise widely distributed insect groups are excluded from
the passenger list. Besides, would not disharmony result even if land connections
once existed?—the islands have remained isolated for a long time, perhaps
millions of years, while migration, favoured by climatic and edaphic changes
affecting the general character and composition of the vegetation has continued
over land on the continents.

HINTON (65) who, as we have seen, was opposed against both continental
drift and land bridges, believed that wind-borne and raft-borne transportation
across the oceans must have been of common occurrence. If this be so, why did
no snakes, frogs or gymnosperm cones ever get aboard the rafts and arrive at
distant islands? Rafts formed by large, uprooted trees are observed in big rivers
like the Amazon or Orinoco, and I guess that an analysis of their composition
would reveal the presence of a rather varied fauna. WALLACE’S Boa constrictor
was referred to above (p. 335). Snakes and giant spiders are often found hidden
in banana trunks imported to Europe from tropical America. On the other hand,
the chances for the formation of substantial rafts are small within the tropical
Pacific, where no big rivers empty.

What kind of invertebrates are likely to withstand transoceantc migration
HINTON has an answer ready:

The chance of accidental dispersal varies according to the nature of the group.
Colonization of the Hawaiian and other islands, always far removed from any conti-
nent, provides us with absolute proof of the kind of animals and plants that can with-
stand long distance wind or raft transportation across the oceans.

This sounds quite simple, but really is a very complicated problem, to
which sufficient attention has not been paid. The possibilities vary according to

i PTSBERG
344 Cc. SKOTTS

size, flight capacity, habits, mode of reproduction, sensitiveness to changes of
milieu. and so forth, and I am afraid that we have little knowledge, founded on
facts, in most cases.

Earth-worms have been carried all over the world with the human traffic.
It is noteworthy that as a rule no truly indigenous species are found on oceanic
islands, where, if land connections had existed, they could be expected, and even
if the transport of eggs or living animals were effected only by means of rafts
stocked with earth and plant material, as some believe, they ought to be present,
but, as far as I am aware, nobody has witnessed such a transport. The presence
of endemic /eeches on Samoa and Juan Fernandez (Masafuera) can be understood
only if the leeches are carried about on birds acting as hosts, otherwise I cannot
see how they would be able to survive; they are very sensitive to exposure. To
these in particular I should like to apply what GULICK, without referring to any
special group of animals, wrote (z79. 405):

How is it possible at all for creatures that would die almost at touch of sea water to
precede man by a million years on islands standing solitary in mid-ocean? Are their
remote homes really the left-over fragments of ancient inter-continental land bridges, or
are these creatures prima facie evidence that their ancestors possessed an almost incon-
ceivable capacity for passing uninjured over vast stretches of open ocean: The extremes
of hypothesis that have been proposed in response to this dilemma show us how difficult
it has been to find a solution.

Freshwater crustaceavs occur on many islands, both traffic-borne and indigenous
species; only the raft theory would account for their spread.

It is supposed that cocoons of spzders are transported by wind, webs are
torn loose with cocoons attached and carried up into the air, where a storm takes
care of them. Theoretically this is not impossible, but whether the contents stand
a journey of thousands of miles is doubtful. Adventitious species are found on
Pacific islands, but the bulk of the spider faunas is indigenous and endemism is
high. BERLAND, pointing to the general distribution of genera and the high spe-
cific endemism, is in favour of former land connections (23. 1052).

Liisolation doit étre assez ancien pour ce que cette fauna ait pu acquérir les carac-
teres d’endémisme qu'elle présente. Un botaniste a fixé vers le Pliocéne cet isolement,

mais je serais porté a croire qu/il est plutdt plus ancien, en me basant sur la lenteur
de | évolution des Araignées. :

Mayr, who rejects all land connections, remarks (179. 214):

Considering the haphazard manner by which these oceanic islands receive their popu-
lations, it is rather astonishing how similar the faunas of the various islands are. BER-
LAND, on the basis of the distribution of spiders, has come to the conclusion that the
fauna of all Polynesia is so uniform as to suggest that these islands are but fragments
of a single land mass. This view is similar to Pilsbry’s, founded on Mollusca. Actually,
this paradox of the similarity of the faunas of oceanic islands is solved in quite dif-
ferent manner. Of all the possible families, genera and species of the Papuan Region
that are theoretically in a position to colonize, only a small fraction will eventually
avail themselves of the opportunity.

DERIVATION OF THE FLORA AND FAUNA 345

This is quite true, but will not the result become much the same with land
connections? The main source of the fauna is the same, the similarity is a con-
sequence, and local endemism is a result of isolation.

Acarids present the same problem. MUMFORD (783) remarks that is is very
difficult to compile lists of species for the Pacific islands and that, at present,
no safe conclusion can be drawn with regard to the distribution of genera. This
is true, but we know that numerous indigenous species occur and that local en-
demism seems to be high. How these extremely delicate little animals would be
able to stand long overseas voyages is difficult to imagine. They are plentiful in
humid forest soils on Juan Fernandez and all the species except two adventitious
ones are endemic. Little is known of their relationships. If washed down from
the hill-sides and carried out into the sea, they will die—only wind transport
remains. Pseudoscorpions may be more resistant to both salt water and desicca-
tion, but their pronounced endemism bears witness of long isolation. All the false
scorpions recorded from Juan Fernandez are endemic, and there is one endemic
genus. They were unknown when MUMFORD (I.c. 246) wrote:

As Chamberlain points out, it is doubtful whether anything like true insular endemism
occurs in most species of false scorpions because of the ease with which they are distrib-
uted.

They are well adapted to be carried about by man. Among the J/y77a-
poda are many local endemic species. These creatures are, according to my own
experience, rather tough and might be able to spread by the same methods as
earth-worms and land crustaceans. It is not very probable that they are blown
from continents to distant islands. Cosmopolitan forms are probably adventitious.
If this applies also to Collemébola 1 do not know; very wide-spread species may
be so ancient that they have attained their distribution when the map of the globe
was quite different from the present one. It is unlikely that they are able to
migrate overseas. LINDSAY (767.719) writes:

The primitively wingless Collembola seem to constitute better material for distribution
studies than any other insect order, because migration by flight is impossible and the

delicate integument makes it very unlikely that the insect could be carried any appre-
ciable distance by the sea.

We can safely add that, if blown out over the ocean, they would soon perish.
ZIMMERMAN suspects that none of the 32 species recorded for Hawaii is indige-
nous there; thus, all are supposed to have been imported by the traffic. How-
ever, 3 species are supposed to be endemic in Juan Fernandez. Two species of
the Hawaiian 7hysanura are “possibly endemic’ (ZIMMERMAN) but perhaps adven-
titious. These creatures do not appear to be fit for long-distance dispersal, and
the two species known from Juan Fernandez are endemic. One of them belongs
to a monotypical genus with Australian affinities, a disjunction not easily bridged
over without land connections.

Getting to the true zzsects, their mode of dispersal certainly varies a great
deal. Whereas butterflies, moths, flies, hymenoptera etc. are known to be storm-
driven and eventually carried far, heavy beetles, even if they be properly winged,

340 C. SKOTTSBERG

are unfit for long journeys; whether they are transported by waves and currents and
able to stand immersion in salt water during weeks and months I do not know.
The endemic flightless insects have, just as the flightless birds, given cause to
much speculation. They are supposed to descend from winged species; arrived, it
is said, on an oceanic island, they had the choice of losing their wings or being
blown off the island and lost altogether. ZIMMERMAN does not favour this view,
for they may as well have lost their power of flight on the mother continent,
which did not prevent them to be carried off by a hurricane. MAYR agrees with
him, comp. quotation p. 336. Very well, but is it possible to imagine a flightless
rail carried a thousand miles across an ocean?

There are numerous Ovthoptera on Hawaii, most of them endemic species or
even belonging to endemic genera. So far only 4 species have been reported from
Juan Fernandez, two of them endemic. I presume that ZIMMERMAN regards these
insects as normally wind-borne. The relations of the Hawaiian Gryllidae are with
Indo-Pacific forms, so they have had a long way to go. The four species of
termites found in Hawaii are adventitious, which should indicate that they are
unable to reach oceanic islands without human assistance, but the single species
discovered in Juan Fernandez (Masatierra) is endemic. Either did it, or its ancestor,
arrive over land, or a colony was carried in a floating log, which may seem un-
likely, or a storm brought a winged, swarming couple which founded a new colony
—even less probable. The termites are a very ancient order and date, it is said,
back to the Mesozoic at least.

Mallophaga are spread with their bird hosts. All the endemic Hawaiian species
live on the Drepanididae, and the marine birds here and on Juan Fernandez are
infested with widespread forms. 7hysanoptera seem to be easily spread with human
traffic. Most of the 99 species found in Hawaii are adventitious, few indigenous.
Two of the 4 species in Juan Fernandez are endemic. How these delicate insects
manage to get about and to reach oceanic islands I cannot tell.

The Hawaiian islands have a rich and peculiar fauna of Weuroptera, some
“among the most aberrant of all’? (ZIMMERMAN p. 76). Of the five species re-
corded for Juan Fernandez 4 are endemic, one of them belonging to an endemic
genus. Wind drift must be taken into account, but it cannot be very effective. The
situation remains the same when we get to the Lep7doptera, about 1000 species
in Hawaii, of which 85 % have not been found elsewhere. Wind drift or immature
stages (eggs, etc.) carried with plant material are the only possibilities, but they
cannot be very great. So far only 26 indigenous species are known from Juan
Fernandez, 70% of them endemic; when the list of Dr. KUSCHEL’s collection, which
contains over 50 species, has been published, the figures will undergo alteration.
In his survey of the Pacific lepidoptera SWEZEY (262) includes the Galapagos
Islands, but does not mention Juan Fernandez. The Pacific islands were, he says
p. 319, populated from the Malayan and Oriental regions, and the fauna arrived
in the main by accident, winds, typhoons etc. or with plant material brought by
currents. A comparable numerical development of species per genus has taken
place in no other islands than the Hawaiian, and the author infers that they have
a more ancient fauna, descended from ancestors that arrived at a more remote

DERIVATION OF THE FLORA AND FAUNA 347

time. This does not accord with ZIMMERMAN’s statement that no primitive family
is represented in Hawaii. Probably the Hawaiian Dzp/era have been less collected;
ZIMMERMAN, who is responsible for the figures quoted, indicates about 400 species,
of which 60% are endemic. In relation to their small size the Juan Fernandez
Islands seem to be richer with about 150 indigenous species (64 % endemic). Our
knowledge of the dipterous fauna is in the main due to Dr. KUSCHEL’s collec-
tions; the 25—30 species not yet reported on will raise the total number.

Over 1600 species of Coleoptera are reported from Hawaii, of which about
76% are endemic, and there are numerous endemic genera. Numerous species
found elsewhere are adventitious. In relation to its size, Juan Fernandez cannot
be called poor (see above p. 307, etc.), and endemism is just as high here. I presume
that ZIMMERMAN and others regard wind as the principal dispersal agent, though
not for all kinds of beetles, because we have to do with many different types
of animals and of habits and habitats. It is difficult to imagine how a flightless
beetle would be able to keep afloat in the air for thousands of miles; he must
have had a great need of the numerous “stepping stones’ postulated by ZIMMER-
MAN. It should perhaps be mentioned that GULICK (zzg. 414) wrote that “‘it can
hardly be doubted that some carrion-feeding insects have been distributed by
adhering to sea birds’. I doubt that this ever happened, but as he had just dis-
cussed the transport of “invertebrate eggs” in the digestive tract of birds, I sup-
pose that he means eggs of necrophilous flies or beetles which, brought across
the sea, were deposited on another carrion and thus became established on an
oceanic island.

The weevils, a most important and interesting feature in isolated island
faunas, are often dependent on definite host plants, and are thought to sail along
on logs as stowaways. Evidently the Curculionidae have become something of a
stumbling-block. USINGER (273) came to the conclusion that they must possess
some unknown special means of dispersal. BRINCK, who discussed the coleopterous
fauna of Tristan da Cunha (376. 97—104), another isolated volcanic group of islands
where geologists failed to discover any traces of land connections, states that
the fauna contains endemic elements and offers examples of remarkable disjunc-
tions. It must have originated from extinct faunas of neighbouring continents if
not of submerged lands. The only natural agent capable of transportation is the
wind, but BRINCK is convinced that “‘at present no beetles are invading the islands
by natural means” (p. 103), and the reason is not adverse conditions, for several
species, introduced with the human traffic, have become naturalized. The unavoid-
able conclusion is that dispersal agents, man excepted, are insufficient—and they
were the same in the past. The original beetle fauna has survived from pregla-
cial time, an hypothesis that nobody would feel inclined to reject, but it does
not help us to understand by what means it was able ever to arrive. The weevils
of Hawaii have had a long way to come; according to ZIMMERMAN the ancestors
as a rule came from the south Pacific or Indo-Pacific regions. Generally he regards
also the peculiar genera to have originated in Hawaii or, eventually, in one of
the lost islands serving as intermediate stations, but among the weevils are some
that defy all explanations:

348 C. SKOTTSBERG

Nesotocus is evidently a relict endemic genus of four closely allied species, and
there appears to be nothing like it elsewhere... Oodemus with its 58 species. 1S
the largest genus of the Hawaiian Cossoninae .. . together with its close ally Anothes

(3 species) endemic, and I know of no genus or group of genera from any region from

which is might have come. It is an anomaly.

Such cases bring us back to times long before the formation of the present
Hawaiian chain, and similar cases are found also in Juan Fernandez, e.g. the
endemic tribe ¥vanorhint of AURIVILLIUS. Other examples are offered by many
other insect groups. And, leaving them aside for a moment, is not the endemic
Hawaiian bird family Drepanididae another anomaly? Whereas BRYAN (39. 188)
finds a Malayan origin most acceptable, GULICK writes (77g. 420):

The history of Hawaiian land birds must have begun with the arrival of some form

of tropical American honey creeper, which became in due time the progenitor of all 18
genera and 4o species of the Drepanididae.

He seems to have forgotten that the islands are claimed to have dived out of
the ocean in late Pliocene and Pleistocene times.

Hymenoptera are plentiful in Hawaii, about 600 native species, among which
endemics are numerous, and the ancestors are supposed to have come from the
south and southwest Pacific, in exceptional cases from Asia and the Orient. This
order is as yet little known in Juan Fernandez, see above p. 315. Wind drift seems
to be the only possible mode of transport unless infected larvae of butterflies etc.
arrived with drift-wood, which does not seem very probable. Ants are easily spread
with the traffic. Of the 3 species known in Juan Fernandez only one appears to
have arrived without human assistance.

Among the Heteroptera in Hawaii, over 200 species and 80 % endemic, ZIM-
MERMAN pays special attention to the genus Vyszws, which has its greatest known
diversity in those islands. All are endemic and include the most divergent of all
Nystus species. The ancestors are supposed to have come from the south and
west Pacific. The genus is, according to USINGER (273) common in the Australian
and Oriental regions, extending through Melanesia to Fiji and Samoa without a
single representative east of this line, but important in the Hawaiian chain; the
author seems to have overlooked its occurrence in Juan Fernandez. Myszus is
supposed to have reached Hawaii by a circuitous route over open water and the
Leeward Hawaiian islands. This route is indicated by a submarine ridge of con-
siderable depth and may once have been interrupted by island peaks such as
Wake Island; thence it is followed to the Marianas and Caroline Islands and even-
tually to the rich Papuan and Australian regions. But in other cases it is less
easy to construct a suitable route: “The presence of twenty very unique genera
in Hawaii and their absence from old, high islands along the very route they are
said to have traveled is inexplicable by present theories” (l.c. 315). Why not
presume that all related genera have died out?, which is the easiest explanation.

Nysius is a widespread genus, well developed also in New Zealand, and the
single Juan Fernandez species is claimed to be related to another from New Zea-
land. We know numerous examples of the same kind in other animal groups and
particularly among the plants we have called Antarcto-tertiary.

DERIVATION OF THE FLORA AND FAUNA 349

The Homoptera present the same picture in Hawaii, about 350 native spe-
cies, most of them endemic and suggesting a southwest Pacific ancestry. Of the
species reported from Juan Fernandez only one-third have been described, all
endemic and as far as we know with their relatives in the south and central Pacific,

No order characteristic of insular faunas has aroused greater interest among
biogeographers than the Pacific /aza molluscs, nowhere more wonderfully devel-
oped than in the Hawaiian chain. In the discussion of land connections they
occupy a central position.

WALLACE found that the wide distribution of the land ‘snails is “by no
means so easy to explain as that of the insects’; the chances have been ‘rare
and exceptional’, possibly eggs stuck to the feet of aquatic birds, or the animals
themselves were storm-carried, “attached to leaves and twigs’’—this would be
the only means by which viviparous forms could be transported.

GULICK (rg) treats the land-shell problem at some length. Speaking of Easter
Island he asserts that “hurricanes spread gravid land-snails as dust over almost
as great distance as plant seeds can be blown’, but he gives no facts to support
this very positive statement. The land-shells of Juan Fernandez and Saint Helena
are then remarked upon.

At least three elements can have derived their ancestry only from Polynesia, fully
3400 miles away, unless Easter Island served as a way station.

The archaic complexion of the snail fauna is not necessarily very significant, as
younger continental forms do not for the most part yield minute, easily wind-blown
species.

Archaic—exactly, malacologists emphasize that more modern types do not occur
on oceanic islands unless brought by man, and this has, naturally enough, been
used as an argument in favour of early land connections before the modern types
existed. In GULICK’s view the size, not the age, decides.

Large, softskinned creatures invariably make a poor showing... large helices ex-
amplify this disability so excellently that their failure to arrive is a sort of negative
criterium for insularity (p. 414).

But neither are all continental species large, nor all insular ones small, and the
oceanic snail fauna includes many forms that cannot spread like dust. GULICK
remarks on the genus Partu/a that “‘its 120 geographically restricted species mostly
weigh too much and are too tender to fit easily into theories of transport by
air or sea’, and this makes him take into account the possibilities of “land ridges”
to facilitate transport.

“It is evident’, he says p. 419, “that the vast diversification is a proof of
the great local antiquity of these families, and hence of a considerable antiquity
of their island habitat.’ This is not true of the islands as they appear now and
as they have stood for probably millions of years.

Land snails, just as weevils, are dispersed by some unknown method, USIN-
GER thinks (lc. 315), while MAYR, as already quoted, regarded the presence of
small molluscs on oceanic islands as a proof of the efficiency of hurricanes, but
later on expressed himself as follows.

350 C. SKOTTSBERG

It seems to me that the wide acceptance of land bridges by conchologists is chiefly
due to three reasons: (1) our almost complete ignorance of the means of dispersal of
snails, (2) our lack of knowledge of the speed of speciation in snails, and (3) faulty classi-
fication, particularly generic classification. A. Gulick has already directed attention to the
presence of snails on most oceanic islands. They were unquestionably carried there by
some unknown means of transportation.... To me it seems incomparably simpler to
assume a still unknown method of transportation than a land bridge that is unsupported
by any other fact.

Simple, no doubt, but we cannot get away from the problem by an “ignoramus”’.

It is easy to understand that the presence of fairly large forms of land-shells
on distant islands has caused a good deal of trouble. ZIMMERMAN tried to find
a way out of the difficulties.

It has been said that large snails such at the Hawaiian achatinellids and amastrids
are particularly unsuited for overseas distribution. However, it we approach the problem
differently, different conclusions may be reached. If, as I believe, the large Hawaiian
snails have evolved from small or minute ancestors, then the argument based upon
their large size loses its weight. However, if small snails can be distributed overseas,
then what is to prevent eggs or tiny, immature specimens of large species from being
similarly transported? (I.c. 61).

In passing, ZIMMERMAN quotes H. B. BAKER, who thought that land-shells
are carried along by migratory birds. After these speculations it is refreshing to
read BRYAN (go. 9):

The presence of certain kinds of plants and animals found in Hawaii and related
to species in the southwest Pacific is hard to explain by any known means of drift,
either over the sea or through the air. Land snails constitute one such group. These
mollusks, which breathe directly from the air, would drown in water, particularly salt
water; yet they must have moisture. They cannot stand long exposure to the sun, but
live on the leaves and trunks of forest plants or beneath fallen leaves and trash on the
ground. How did their ancestors reach Hawaii if they could neither swim nor drift?

Opponents against the dogma of colonization across the oceans.

Many biogeographers have arrived at the conclusion that the natural dispersal
agents cannot be made responsible for the distribution of all kinds of biota across
very wide expanses of open water. Most of the authors are botanists, which is sur-
prising because the chances should be greater for seeds and spores than for eggs
or individuals of delicate creatures. To ask for land bridges, or for extension of
continental margins later submerged but leaving behind land fragments, is to refuse
to accept overseas migration as the only possible means of colonization. Conse-
quently, a number of authors have already been quoted in the chapter on the
history of the Pacific basin, GOOD, IRMSCHER, WULFF, CAMPBELL; etc., as well
as my own contributions to the discussion, the latest in 1951 (248). I shall add
here that I never disclaimed every possibility of migration over wide expanses
of water; see 226 and 232. Various writers have compared isolated peaks on
continents with oceanic islands and have stated that under present conditions an
exchange of biota is improbable. VAN STEENIS expressed his opinion when dealing
with the Malaysian mountain flora, in very plain terms (258); too many facts

DERIVATION OF THE FLORA AND FAUNA Soe

show the impossibility of attributing any importance worth mentioning to long-
distance dispersal; the theory is, when it comes to migrations of floras, “not
worth a straw’, and REICHE (203), referring to the history of the Chilean flora,
called it “‘eine Kette von Unwahrscheinlichkeiten”. GORDON (773) was strongly
inclined do deny ist value and believed that plants, and more so plant associa-
tions, advance slowly over land and do not jump thousands of miles; he could
not, however, help paying some attention to “Nature’s great Krakatau experi-
ment’, of which enough has been said.

CAIN (42) thinks that “‘migration is usually not a random matter” (p. 162)
and that long-distance dispersal rarely has resulted in migration and establishment,
nor does it explain the discontinuous areas. In reality he belongs to the trust-
worthy opponents. One of the reasons advanced by him is, however, not con-
clusive:

The phenomenon of local races (subspecific endemics) is entirely opposed to the

idea of long-distance dispersal, for such variation depends upon isolation which would
not exist if long-distance dispersal were generally effective (p. 161).

Those who are in favour of the theory emphasize the haphazard character
of the procedure; success may follow once and never more. If the immigrant
belongs to a polymorphous species, with intermediate forms between the sub-
species or varieties, it may, isolated as it remains from the rest of the population,
stand out as a separate insular taxon.

Among the zoologists few, mainly malacologists, are in favour of land con-
nections, but some entomologists agree with them. BERLAND was already quoted.
As a rule a connection South America—Antarctica—Australia (or New Zealand)
is asked for, but to ENDERLEIN this bridge was not sufficient:

Die zahlreichen endemischen Gattungen (9) und Arten (2) zeigen, dass die Juan
Fernandez-Inseln ein Refugium fiir die Reste der Faunen unfangreicherer untergegan-

gener Gebiete darstellen, die nicht mit dem neotropischen Gebiet in Verbindung gestan-
den haben (ENDERLEIN 87. 643).

Did ENDERLEIN dream of a submerged Pacific continent? Many of the island
flies are also native of the mainland of Chile, others have their relations there,
and if the islands were formerly connected with some other land, it was with
South America. Besides, the majority of ENDERLEIN’s new genera have been
reduced to synonymy by later authors, his speculations have little weight, and
WYGODZINSKY (294. 81), referring to Gigantodax kuscheli, arrived at a different
conclusion. G7zgantodax is an exclusively South American genus.

Chapter VII.

Biological characteristics of isolated islands.

In his classical “Lecture on insular floras’ (z38) J. D. HOOKER formulated,
in very lucid terms, the special features of island floras and his opinion on their
evolution. As examples he chose Macaronesia, St. Helena, Ascension and Ker-
guelen Island. Their peculiarities were stated under five items.

352 C. SKOTTSBERG

1. In all cases considered floristic relations exist between the tsland and one
mother continent.

2. The floras of all the tslands in question are more temperate in character
than that of the mother continent on the same latitude.

This may be true in most cases thanks to the influence of the surrounding
ocean; it holds good for Juan Fernandez.

3. All these islands show many biological peculiarities by which they are
distinguished.

The distinguishing characteristics are mainly expressed in endemism. HOOKER
referred the endemics to two categories, such as do not show affinity to the plants
on the mother continent, and such as, even if belonging to endemic genera, are
related to continental ones.

If we turn to the Pacific where, for obvious reasons, only the high volcanic,
well-watered islands are considered, we find that all of them are distinguished by
numerous, in many cases also very remarkable endemics. Hawaii stand out above
the others and so do Juan Fernandez and Desventuradas; the floras of Tahiti,
Samoa, Marquesas etc., as well as of Micronesia, are less independent. The degre
of spatial isolation is not conclusive; the flora of Juan Fernandez is more peculiar
than that of Marquesas which are situated much farther away from any continent,
and this holds good not only for the angiosperms but also for the ferns. En-
demism in angiosperms is 69 % in the former and 50 in the latter; of the ferns
about 30% in each, but only Juan Fernandez has an endemic genus.

4. The general rule is that the species also found in the mother continent are
the most abundant, the peculiar spectes are rarer, the peculiar genera of continental
affinity rarer still, but the plants with no affinity elsewhere are often very common,

This is, I suppose, true of the islands examined by HOOKER, with the
exception of St. Helena before the arrival of man, but not of Hawaii, nor of
Juan Fernandez. Some of the continental species—Lzbertia and a few grasses—
are abundant, while others are rare, all according to the supply of suitable
habitats; among the peculiar species of continental affinity are many quite
common ones, e.g. the endemic species of Acaena, Drimys, Dysopsis, Es-
callonia, Gunnera, Myrceugenia (the leading forest tree on Masafuera), Pernettya,
Rhaphithamnus, Ugni, Evigeron fruticosus, Uncinta Douglasi; and of the peculiar
genera allied to South American ones, Nothomyrcia is the leading forest tree
on Masatierra, where Ochagavza is also common. To these may be added such
common endemics as Loehmeria and the species of Hagara and Coprosma, under
the assumption that related species of Boehmeria, and representatives of Coprosma
and /agara, once belonged to the neotropical flora. The plants of no affinity in
the mother country, that is the nearest continent, are as a rule very local, few
are common and many extremely rare.

Possibly I have misunderstood HOOKER here. When we say that a species
is abundant in a country we mean that it is copious; if we call it common, it
is widely spread; if we call it rare, it has been reported from a small number
of localities only; and if we use “species” in plural we mean the same thing.
But what if HOOKER with “‘abundant’’ and “common” wanted to say that these

DERIVATION OF THE FLORA AND FAUNA 353

species were the most numerous in the flora, and that the “‘rare’’ ones were few
in number? If this was what HOOKER meant, the result will be: Species also found
on the continent: 46; endemic species allied to South American or other conti-
nental species: 71; endemic genera, related to continental genera: 5 (6 species);
endemic genera not related to continental ones: 12 (24 species). Still, this is per-
haps to give a wrong interpretation to HOOKER’s words. There are very few
native non-endemics on St. Helena, and they are of course not in the majority
in the Canary Islands, nor are, in the latter place, the isolated endemic genera
and species very many. In this case HOOKER says “‘plants’’, not “‘genera’’ and
“species”. If, to this group, as represented in Juan Fernandez, we add isolated
endemic species, not related to any species in the mother continent, although the
genera occur there, this group comprises 54 plants, or more than 1/, of the angio-
sperms, while group 2 is reduced from 71 to 40. The figures would be: 46, 40,
6, 54, thus conforming much better to HOOKER’s rule.

5. [Indigenous annual plants are extremely rare or altogether absent.

Here “rare’’ must mean few species, and Therophytes are very few in Juan
cernandez and some of the registered species perhaps not originally native.

How were plants transported to distant islands? HOOKER’s answer is: either
across the sea or over submerged bridges, and he adds: “‘the naturalist, who takes
nothing for granted, finds insuperable obstacles to the ready acceptance of either’’.
The situation is still the same 90 years after HOOKER.

HOOKER regarded the isolated island plants as “relics of a far more ancient
vegetation than now prevails on the mother continent’, but he most certainly
never wanted to say that the continental flora was altogether younger than the
insular but that species, now restricted to the island, formerly occurred on the
mother continent, having become replaced there by younger species. He based
his opinion on the fact that Macaronesian relicts had been found as fossils in
Tertiary deposits on the continent. Time has not permitted me to collect modern
data, and I can only suppose that some of the old determinations still hold good.
The vegetation of Europe has undergone great changes “within the lifetime of
these Atlantic island species’; they once grew in Europe, but were driven out
from there to be preserved on the islands, which they had reached “when condi-
tions may have been very different from what they are now”.

The theory of a continental Macaronesia, including Madeira and the Azores,
goes back to FORBES’ theory of the former connection between the British Islands
and the mainland, definitely proved ages ago. FORBES went further and revived
the old idea of a lost Atlantis, still favoured by many.

It is interesting to follow HOOKER’s discussion with his friend DARWIN on
island problems. DARWIN believed in the efficiency of dispersal agents to carry
plants and animals across wide expanses of sea, and his arguments made such
a deep impression on HOOKER that he became almost convinced. Still, he hesi-
tated, and certain serious difficulties prevented him from fully accepting DARWIN’s
ideas. The composition of the flora of the Azores was not what we had reason
to expect from the direction of winds and currents. The Macaronesian Ornis is
almost the same as in Europe and undoubtedly came from there, but the flora

23 —557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

354 C. SKOTTSBERG

is considerably different; thus it may be argued that “the birds and plants do
not come under the same category’. DARWIN replied that

the migration of birds is continuous and frequent, and the individuals surviving and
breeding, they keep up the specific type, and do not give origin to local varieties;
whilst the transport of seeds is casual and rare, and very few surviving, these not being
crossed by the original stock, in the process of time give rise to varieties, etc., and
do not perpetuate the continental races (py 10):

This is the situation in a nut-shell, and DARWIN’s arguments are repeated by
scores of biogeographers to this very day.
Also St. Helena, Ascension and Kerguelen made HOOKER hesitate:

They [St. Helena and Ascension] have no land birds, but an African vegetation;
and though nearly midway between Africa and America, they have scarcely a single
American type of flowering plants; and Kerguelen’s Land has a flora of whose ele-
ments most have emigrated not from the nearest land, but from the most distant (p. 10).

HEMSLEY (727. 59) remarks that HOOKER seems to have forgotten the
Compositae in St. Helena, most of them showing American affinity.

Kerguelen’s nearest land is Antarctica, but not a single flowering plant is
known from the coast south of Kerguelen. Africa as a mother country—it goes
without saying that subtropical or warm temperate plants cannot endure a
subantarctic climate, and only the most distant lands, Tierra del Fuego and the
Falkland Islands, were, thanks to the strong and constant west wind drift, re-
garded as a mother country. Even if South Georgia served as an intermediate
station, the distances are very great; besides, we do not look west for the an-
cestors of the peculiar endemics in the Kerguelen area. Where the capacity of
the dispersal agents appeared to be inadequate, HOOKER was strongly inclined
to look for better land connections. The existence of identical Macaronesian
species on Madeira and the Canary Islands can, he says, hardly be explained
without the help of

intermediate masses of land, as the Salvages (supposing them to have been larger) . .
the only conceivable means of interisland transport... and if intermediate islands are
granted (and Mr. Darwin freely admits these), why not continents?

He must have found that the distance between Madeira and the Canaries is too
large to permit direct transport of diaspores under present wind and current
conditions; in the Kerguelen case they are, at least, favourable. Nevertheless,
later on HOOKER’s faith in transoceanic migration was not as steadfast as before;
the case of Kerguelen troubled him (339):

Turning to the natural agents of dispersion, winds are no doubt the most power-
ful, and sufficient to account for the transport of Cryptogamic spores; these, almost
throughout the year, blow from Fuegia to Kerguelen Island, and in the opposite
direction only tor very short periods, but appear quite insufficient to transport seeds
over 4000 miles (p. 13).

Various phenomenons ... common to... Kerguelen, the Crozets and Marion,
favour the supposition of these all having been peopled with land plants from South
America by intermediate tracts of land that have now disappeared; in other words,

DERIVATION OF THE FLORA AND FAUNA 355

that those islands constitute the wrecks of either an ancient continent or an archipelago
which formerly extended further westwards... (p. 15).

GULICK’s paper “Biological peculiarities of oceanic islands’ does not, con-
trary to its title, contain a review of the special characteristics of island biota;
his object was to expound and defend the theory of permanent isolation of is-
lands like the Hawaiian, Galapagos, Juan Fernandez, St. Helena, etc., which are
said to offer irrefutable proofs of true oceanity. Of this enough has been said
already; I shall return to HOOKER’s five points, to which others may be added.

Endemism.—Yhe occurrence of numerous genera and species restricted to
oceanic islands has caused much discussion. ‘‘Reichtum an Endemismen ist tber-
haupt der hervorragendste Charakterzug der Insel-Floren’’, HAYEK wrote (304). It
is, however, equally pronounced in continental districts like the Cape region, south-
western Australia, western China, California or Chile, where local concentrations of
endemics are found.

If insular endemics show distant affinity only or, in extreme cases, no af-
finity at all, to continental taxa, they are looked upon as relicts; as the islands
are geologically young, the endemics have not evolved there but must have im-
migrated from some mother country, where they have become extinct. They may,
however, have undergone some change after their arrival to the island. There
is also a possibility that the continental progenitor has, in its turn, changed in
a different direction, making its descendants so unlike that their relations are
obscured. Species only slightly different from continental ones are much more
numerous than the relicts; they are supposed to have originated in the islands
and give examples of so-called progressive endemism. As CHRISTENSEN (60. 149)
pointed out, another alternative leading to the establishment of endemic species
should be considered. On the continent, from where a plant found its way to
an isolated island, opportunities for crossing with other species often exist,
eventually leading to the disappearance of the original taxon with its special
characteristics. Its island offshoot does not share its fate but remains true to the
original type. The island form did not originate through a genetic change of
the continental species: it represents the surviving species and is, as it were, a relict.
This does not apply to the pteridophytes. Crosses are extremely rare, the fern
species represent, in a high degree, pure lines, whence it follows that insular en-
demics are much rarer than among the phanerogams. This is true, but it is
usually explained as a result of the enormous spore production and the facility
with which they spread.

Opinions about the true nature of systematically isolated taxa vary. GUPPY
(722) regarded them as either highly specialized products of the islands, “the first
of their race’, or modified forms of allied continental genera, the majority of which
had passed away, “‘the last of their race’ and probably doomed; to him the
islands appealed “more as registers of past floral conditions in the continents
than as representing their present state’—this in accordance with HOOKER’s
views. The Age-and-area theory of WILLIS (286) claims that wides are older
than endemics, a rule with few exceptions; in another paper (285) he states that
“insular endemic genera are as a rule young beginners, not relics’. I have dis-

356 C. SKOTTSBERG

cussed his theories in an earlier paper (2377) to which I refer. RIDLEY (205) called
the island endemics “newborn species’, admitting that all did not fall within this
category but were “‘epibiotics, relics at the end of their species life . . . unable to
reach another suitable spot for their growth”. It is noteworthy that so many of the
“epibiotics’ are Composites, famous among diffusionists for their alleged effective
dispersal mechanisms; in RIDLEY’s eyes they are, perhaps, pseudo-relicts. CAIN (42)
takes more or less the same position: “the relic nature of an endemic should
never be accepted without some form of positive evidence” (p. 227); proofs are hard
to find, no island cases as clear as Gzzkgo or Metasequota are known. An endemic
inhabiting a strikingly limited area may be a young species that had no time to spread,
or it may be too stenotopic, but others are what CAIN (p. 230) calls ‘“senescent’’; such
species occupy a small area, are relatively constant, ecologically of narrow amplitude
and show low competitive ability. They are unable to “penetrate the prevailing
habitats that are dominated by the typical vegetation of the region’ but behave
just as stenotopic young beginners. If indeed old and senescent they ought to
show some primitive characteristics. According to CAIN senescent species con-
stitute ‘‘an anomalous element in the flora of a given region’’; this may be true,
for in many instances they survive from an earlier climatic period and are barely
able to hold their ground under the changed conditions. Nevertheless there are
cases when such anomalous species form the typical vegetation of a certain habi-
tat and where nobody would dream of regarding them as young beginners; the
“Robinsonia assemblage” in Masatierra offers a good example (see 257).
Number of species per genus.—We know that island floras contain a fair num-
ber of monotypical genera, many genera that are large elsewhere but repre-
sented on a given island by a single species, and few with many species, so that
the numerical relation between species and genus approaches 1 and does not
exceed 2, and this has been regarded as a good proof that the island is truly
oceanic and has been peopled accidentally by waifs and strays. This rule is not
without exceptions, among which the Hawaiian flora is the most striking. FOs-
BERG, who contributed a chapter on the higher flora to ZIMMERMAN’s book (298),
indicates 83 families, 216 genera and 1729 species of angiosperms. The figure
for the genera may be a little too low. With regard to species all depends on
the species concept. HILLEBRAND (307) was conservative; from his Flora DRUDE
(305. 136) got the ratio 6.2: 1, but scores of well-marked species have been de-
scribed since 1888, and in addition, particularly during the last two or three decades,
a large number of taxa that are little more than microspecies; unfortunately
no case of apomixis has been found as yet.! It is difficult to know which way
to take out of this maze; neither the role played by hybridism nor the existence
of modifications due to environment has been duly considered. Based on FOs-
BERG's figures, the relation species: genus is 8:1, and this is not at all what one
expects to find in an oceanic flora. Nobody is, I think, likely to disclaim New

+ Among taxonomists particularly responsible for this alarming increase E. E. SHERFF and
H. Sr. JOHN should be mentioned first, but some others have also contributed and the writer

cannot plead innocent. SHERFF, in addition to numerous new species, has described an endless
number of varieties.

DERIVATION OF THE FLORA AND FAUNA 357

Zealand's continentality. According to CHEESEMAN the number of families is 97,
of genera 382 and of species 1415; the relation in question is 3.7: 1. I cannot
attach much importance to such figures. If an island is a remnant of a sub-
merged land-mass upon which during the process of sinking lava was ejected until
the old foundation disappeared, only a small part of the flora and fauna will sur-
vive the catastrophe, and unless progressive endemism comes to play a role, the
living world will present a picture of disharmony, with a reduced number of
families and a low species: genus ratio. Climatic changes will create a similar
situation, if a portion of a continental flora has lost its connection with the con-
tinent. The Falkland Islands rest on the continental shelf, they are formed by old
sediments on a granitic foundation and are, universally I think, classified as con-
tinental islands, but their angiospermic flora is disharmenic and fragmentary: 38
families, 97 genera and 143 species; the ratio is 1.47: 1. There is a single endemic
genus and a small number of endemic species. The poverty is due to a severe
climate and to losses suffered during the pleistocene period of solifluction, con-
temporaneous with the glaciation on the mainland. The constitution of a flora, con-
tinental or insular, depends on a combination of many factors, geological, his-
torical, climatic, genetic and so forth.

Absence of large, widespread and “successful” families. —The lack of conifers
in islands demonstrates, it is said, that such islands are not continental, for in
all continents gymnosperms are plentiful; ““cones do not float’, and the seeds,
winged or unwinged, have no chances to be carried very far, but it is surprising
that also Taxads and many Podocarps, adapted, as it were, to endozoochorous
bird dispersal, are conspicuous by their absence. Consequently, islands where cone-
bearing species exist are regarded as continental: New Zealand, Tasmania, Nor-
folk Island, New Caledonia, Fiji, etc.

Exceptions occur, islands looked upon as permanently isolated, Bermuda,
the Azores, Madeira and the Canaries, have at least some species of Funiperus;
perhaps junipers possess some kind of dispersal capacity and do not count, but
we cannot get away from /7uus canariensis.

Other large and wide-spread families very poorly represented on isolated is-
lands are Leguminosae, Araceae and Orchidaceae. Peas are a staple food of many
birds but few if any able to pass their digestive tract unharmed; they are too
heavy to be carried any distance by wind and, notorious beach plants excepted,
will sink in water. The dry pods would float but they do not fall off but open
on the plant to discharge the seeds. Nevertheless Sophora sect. Edwardsza has a
number of closely related species scattered over the south hemisphere on islands
as remote as Diego Alvarez, Rapa, Easter and Marquesas, with one species, more
well-marked than the rest, on Hawaii. This is a very puzzling case of disjunct
distribution, but it is not expedient to draw any far-reaching conclusions from
this unique case. The seeds are of the size of a small pea and not equipped with
a capacity to travel greater than in hundreds of leguminous plants, which are
within reach but never crossed any water barriers, nor used any land bridges.
Did they not yet exist, were they not within reach when connection was estab-
lished or have they all died out in the islands? Each alternative seems equally

358 C. SKOTTSBERG

improbable. Araceae are plentiful in humid tropical climates, and some islands,
the Hawaiian for instance, ought to offer a suitable environment; still there is not
a single native species. And it is strange that orchids, with their dust-like seeds,
should be, if not altogether absent, so few, becoming fewer and fewer as we proceed
east from Malaysia and Melanesia; cf. 375, map 21. However, the seeds, even
if carried far, rarely are able to retain their viability long enough to become
established, should they happen to strike a spot where they can germinate. I be-
lieve that also where we find a very rich orchid flora most of the species occupy
restricted areas; their advance over land is slow. A species introduced on pur-
pose will, in rare instances, become naturalized, but I cannot remember ever
having heard of an adventitious orchid. Taking all these circumstances into account,
the poverty of distant islands, even if temporarily connected with other lands,
is perhaps not altogether incomprehensible. Besides, the submerged links may,
for all we know, have been poor in legumes, arum lilies and orchids. CAMPBELL,
admitting that the total absence of Avaucaria, Agathis, Podocarpus, Ficus and Ara-
ceae in oceanic islands of the Pacific is a valid objection, not easy to explain,
suggests that perchance they once did exist there but were destroyed by volcanic
eruptions (g6. 181).

The preponderance of woody plants in oceanic islands—HEMSLEY (227. 31)
emphasized the prominent part taken by arboreous and shrubby species in many
islands and pointed out that in some cases they belong to otherwise herbaceous
families. Many peculiar genera of Compositae are confined to islands such as
Hawaii, Galapagos, and Juan Fernandez, and are scattered over Polynesia (see
318, map 20), and in the Atlantic (Macaronesia, St. Helena) and Indian oceans
(Socotra, the Seychelles, Mascarene Islands); arboreous Compositae are, as HEMS-
LEY justly remarks, by no means restricted to island habitats but numerous in
tropical regions of the continents, particularly in America. Both Hawaii and Juan
Fernandez offer good examples of woody plants belonging to otherwise herba-
ceous families or genera, but HEMSLEY’s statement p. 31 about Guznera in Juan
Fernandez—“‘caulescent species unknown elsewhere’’—is erroneous, for they are
just as caulescent in Hawaii; besides, they are not woody, but herbaceous.

Scarcity of herbaceous species, especially of therophytes.—Native annuals and
biennials are rare in islands. They cannot have been less capable of migrating
across the sea than trees or shrubs, nor could soil or climate prevent them from
getting established; this is at once disproved by the countless herbaceous weeds
brought by man which threaten to overrun so many islands. The reason must
be historical. If it is true that life forms with a woody stem are the most ancient,
and if the original stock making up island floras dates back to before the rise of
herbs, a period during which the distribution of land and sea was another than
now, we can explain the high proportion of arborescent species in islands iso-
lated since millions of years.

Lvidence for a greater antiquity of lignified angiosperms.—The angiosperms
originated in the Mesozoic and are traced back to the Lower Cretaceous. The
tree form was the response to a warm and humid climate. These problems were
subjected to a comprehensive analysis by SINNOTT and I. W. BaILEy (224), whom

DERIVATION OF THE FLORA AND FAUNA 359

I shall quote at some length. They state that evidence for a more recent origin
of herbaceous plants is furnished by palaeontology, anatomy, phylogeny and phyto-
geography. Evidence from palaeobotany is, however, not conclusive, because herbs
are much less fit to be preserved as fossils, but it is true that they increase in
number in the younger formations. Evidence from anatomy and phylogeny are
said to be positive; within a group including both ligneous and herbaceous spe-
cies, the former show other primitive characters; of the Leguminosae the more
primitive members are all woody (Mimosaceae, Caesalpiniaceae), the proportion
of herbs being vastly greater among the Papilionaceae. The authors continue, p. 572:

It is generally admitted that endemic species of a flora... are for the most part
more ancient than the non-endemic element, for they must either have had their origin in
the region ...a process usually requiring a long time—or else they must be remnants
of an older vegetation which has elsewhere become extinct. Endemic genera and
finally endemic families are in this way regarded as progressively more ancient portions
of the flora.

I doubt that this statement is of general application; we cannot argue that,
in a given flora, all endemics are more ancient than the wides, because a widely
distributed species may have remained unchanged for millions of years and be
older than another which, for various reasons, happens to have become greatly
restricted in range and endemic in a small area. WILLIS and his school are dia-
metrically opposed to the opinion of SINNOTT and BAILEY and neither is in pos-
session of the absolute truth.

Proceeding to an analysis of certain insular floras deserving special attention
the authors assume that, if woody types are more ancient than herbaceous, a
flora which has been for a long time isolated ought to contain a large proportion
of woody endemics, and this is what they do find. Their analysis of the Juan
Fernandez flora was based on JOHOW’s work, but this has long been out of date
and the figures are incorrect (new ones will be found in Chapter IX), so I shall
leave this subject aside here. The general conclusions in the chapter “Discussion
of Isolated Insular Floras’” deserve to be quoted in full (p. 579).

It is thus very clear that woody plants constitute a more conspicuous element in
the flora of isolated oceanic islands than in the flora of adjacent continental areas from
which their vegetation has possibly been derived, and also that the most ancient por-
tion of the island floras, if endemism is to be regarded as a criterion of antiquity, is
much more woody than the recently acquired elements. Annual herbs, which seem to
be the last step in reduction, are almost entirely absent from insular floras, as has
been noted by Darwin, Hooker, and others. Since the vegetation of these isolated
oceanic islands is to be regarded as more ancient in its composition than that of larger
land areas, it may be looked upon as a vestige of an earlier and much more uni-
form flora which flourished over the earth during the middle or latter part of the Ter-
tiary, and before the great flood of herbaceous vegetation, developed chiefly in the
north temperate lands, had spread over the globe. This conclusion is strengthened by
the many similarities which these widely separated island floras bear to one another.

Some of the statements made by these authors are questionable, but in the
main I agree; only I prefer to say “‘early to middle Tertiary’. DARWIN, who was

360 Cc. SKOTTSBERG

the first to note the predominance of trees on oceanic islands, tried to explain
it as a result of natural selection. SINNOTT and BAILEyY’s criticism is absolutely

convincing.
The authors also paid attention to The Ancient Flora of Antarctica, p. 592:

It seems to be a reasonably safe conclusion that all genera commonly designated
as ‘Antarctic’ from their confinement to the temperate region of the southern hemi-
sphere, were inhabitants of the ancient Antarctic continent.

The authors made an attempt to reconstruct this flora and came to the con-
clusion that ‘two thirds of its endemic dicotyledons were woody plants” (p. 599).

In the authors’ opinion most of the herbaceous vegetation originated in the
north; this may be so because in the latter part of the Tertiary period, with the
increasing differentiation of climatic regions, the temperate and cold-resistant flora
is supposed to have taken possession of large areas, but these were concentrated
in the north; except the Antarctic continent there was not much land in the
far south.

Several authors have expressed the same opinion as SINNOTT and BAILEY.
IRMSCHER (7743) was convinced that most of the primitive angiosperms were trees
and that the relation between tropical and-temperate genera and species point in
this direction; to take one example: it is generally acknowledged that the her-
baceous, mainly temperate family Cruciferae descends from woody tropical Cap-
paridaceae (IRMSCHER might have called attention to the miniature Lepzdium trees
in Hawaii):

Auch sonst ist die Abstammung gemissigter Sippen von tropischen nachgewiesen
worden. ... Diese Gegeniiberstellung der gemdssigten und tropischen Sippen lehrt aber
auch ohne weiteres, dass mit der Anpassung von Formen an die extratropischen Zonen
zugleich die Umprigung des Typus des Holzgewdachses in den des Krautes
vor sich ging, somit letztere als die jingeren Formen die Abkomm-
linge von Holzgewichsen sind (I. 209).

Dass die Theorie der Abstammung der iibrigen Wuchsformen von der Gestalt des
tropischen Baumes immer mehr an Boden gewinnt, geht zum Beispiel aus dem Buche

von Bews hervor: “The megatherm hygrophilous forest of the tropics is probably the

most ancient type of habitat... and most recent of all (life-forms) is the annual type”’
(IL 327).

The investigation of JEFFREY and TORREY (744) led to the conclusion that
“the origin of the herbaceous type in Dicotyledons is from woody or arboreal
forms”, and STOCKWELL, in his monograph of Chaenactis (259; Compositae, 33
sp.), states that, ‘‘within a family or genus, woody perennial species are more
primitive than herbaceous annual species’. An evolution in the reversed direction
is, however, postulated by HUTCHINSON (zzo. I. 4) in the Polycarpicae. He regards
the herbaceous Ranunculaceae as the most primitive; a woody structure is, in this
order, secondary. The current opinion is that the woody Magnoliaceae are among
the oldest living angiosperms.

The rosette trees.—Yhe common type of this interesting growth-form is char-
acterized by a candelabrum-like mode of branching, the two or three (rarely more)
innovations situated on practically the same level at the base of a terminal in-

DERIVATION OF THE FLORA AND FAUNA 301

florescence; in some cases cauliflory is observed. A much more uncommon type
is the unbranched stem terminated by a tuft of large leaves (if entire = GRISE-
BACH’s Clavija form, 325.1. 11), either pollacanthic with lateral inflorescences
or hapaxanthic when, after a number of years, a terminal inflorescence ends the
life of the individual; this type is the most unusual of all. All kinds have in com-
mon the very short internodes and the short-lived leaves, a self-evident condition
for the formation of a compact tuft. A reduction of the rosette tree to herbaceous
state would result in a compact caudex multiceps or a single basal rosette; an
hapaxanthic tree would become a therophyte.

These growth forms are by no means restricted to island habitats but are
also found in all continents, they belong to many different genera and families
and are characteristic of such dicotylous families as Araliaceae, Caricaceae, Theo-
phrastaceae etc.; typical examples are found in Compositae (e.g. Espeletia, species
of Senecio), Epacridaceae (Dracophyllum), Lecythidaceae (Gras), Meliaceae (Ca-
rapa), Rutaceae (Spathelia), Sapindaceae, and so forth. Among the monocotyle-
dons the vast majority of palms belong here, the Pandanaceae should of course
be mentioned, and well-known examples are scattered through the Liliiflorae (spe-
cies of Aloe, Cordyline, species of Yucca, Dracaena, Fourcroya etc.), Ravenala in
the Musaceae, and Puya Raimond in Bromeliaceae. Hapaxanthic trees are few;
KRAUSE (762) called attention to the interesting rutaceous Sohureya excelsa Krause
from Amazonas.

Thus, even if rosette-trees are more or less widely distributed over the globe
in warmer regions, it is a fact that they are a particularly conspicuous feature in
island floras. In the Pacific they are very plentiful in the Hawaiian Islands, espe-
cially among the Compositae and Lobeliaceae, also the hapaxanthic type repre-
sented; they occur in the Galapagos Islands (GRISEBACH II. 512), we find many
in New Caledonia, they constitute a large proportion of the poor Juan Fernandez
flora (see Chapter IX), and they are scattered over Oceania. In the Atlantic they are
numerous in Macaronesia (e.g. Campanula Vidalii on the Azores, Musschia on Ma-
deira, species of Aeontum, Dracaena, Echium, Melanoselinum, Sonchus, Sinapiden-
drum, etc.). | suppose that some are found also on the islands of the Indian Ocean,
but I have no reliable information. I may be entirely wrong, but I have the idea
that this is an old-fashioned growth form, a relict element in island floras, and that
the high volcanic islands of the Pacific and the Atlantic, which are so like each
other in geology and topography, are, in their present shape, of approximately the
same age and date back to a period of land submergence and great volcanic
activity. Perhaps also the “inland islands’, the high volcanic mountains of Africa,
deserve to be mentioned in this connection, famous as they are for their magni-
ficent tree Lobelias and Senecios.

Rosette trees seem to favour open, sunny situations; this is certainly the case
in Juan Fernandez, where they are definitely adapted to such habitats. Single
specimens of a few species are sometimes found growing in the shade, with the
result that the internodes become much longer, the dense tuft dissolves and the
formation of flowers is suppressed. SCHIMPER believed that they are adapted to
a very windy climate and fit to withstand the pressure of strong winds better

362 C. SKOTTSBERG

than trees of a more ordinary type, but this theory has been refuted (227), and
recent observations in the field have not made me alter my opinion.

Absence of quadrupeds. The only mammals regarded as possibly native in
isolated islands are bats, but many islands do not have any. Storm-drift has been
postulated, but if the distance be very great, the animals might be without food
too long unless Nature provides a fair supply of air-born insects. To other land
mammals wide stretches of open water are an absolute obstacle and their absence
is regarded as one of the safest proofs of the permanent isolation of oceanic is-
lands, just as the presence of endemic foxes in the Falkland Islands (now extinct)
supports the theory of an earlier connection with South America.

However, we cannot know that indigenous mammals never existed on islands,
gradually having become extinct when the submergence of land had proceeded
and grazing grounds became smaller and smaller. It would appear that this theory
is contradicted by the fact that herbivorous mammals have been introduced by
man to many islands and do make a living there also when not tended, but if
allowed to naturalize and multiply unrestrainedly in virgin surroundings, their
ravages would perhaps prove catastrophal to the native vegetation and, as a con-
sequence thereof, to themselves. They would die out and leave the flora to re-
cover. We must remember that the islands are small, most of them very small,
that herbivorous animals need space and that pasture lands as they exist now
are a result of cleared forest soil and introduction of innumerable alien weeds or
cultigens. Another explanation of the absence of mammals is that the islands were
cut off so early that mammals had not yet taken possession of the earth or were
not universally distributed and perhaps not within reach. But these are mere wild
speculations. No fossil remains have been discovered, nor can they be expected
on purely volcanic islands. We cannot attack this problem with a hope of success
as long as we know little or nothing of the geographical history of islands and
archipelagoes. But we have better remember that continental islands such as New
Zealand, New Caledonia, etc. are in the same precarious position with regard to
mammals as the Hawaiian Islands.

Native reptiles and amphibians are also absent. CAMPBELL (46) mentions in
passing that there are half a dozen lizards in Hawaii, but all are species wide-
spread in the South Pacific; most likely they were introduced with the early human
immigrants or perhaps later. That there are no frogs or toads would indicate,
CAMPBELL thinks, that the archipelago became isolated before the modern kinds
of these animals had been developed. But, as MUMFORD points out, many con-
tinental islands lack all lower vertebrates (783. 248).

How can we explain all these peculiarities in oceanig islands? HOOKER’s answer
was this:

Thus, according to the hypothesis of trans-oceanic migration, and the theory of the
derivative origin of species, we can understand why the ancient types ... should have
survived on the islands to which but few of the superior race had penetrated;—we can
understand how it comes about, that so many continental species and genera are represented
on the island by similar but not identical species and genera, and that there is such

DERIVATION OF THE FLORA AND FAUNA 363

a representation of genera and species in the separate islands of a group;—we can under-
stand why we find in the Atlantic island Floras of such a graduated series of forms,
ascending from variety to genus, without those sharp lines of specific distinction that
continental plants exhibit;—why whole tribes are absent in the Islands; why their Floras
are limited, and species few in proportion to genera;—why so many peculiar genera
tend to grotesque or picturesque arborescent forms... (p. 11).

All this is, perhaps, not so easy to understand as HOOKER, impressed by
DARWIN’s theory, thought. Finally I shall quote the summary of WALLACE’s General
remark on Oceanic Islands (278. 329-330), to bring this chapter to a close:

They all agree in the total absence of indigenous mammalia and amphibia, while
their reptiles, when they possess any, do not exhibit indications of extreme isolation and
antiquity [for the moment WaLLAcrE seems to have forgotten the giant tortoises and the
peculiar endemic lizards in the Galapagos, typically oceanic islands in his opinion].
Their birds and insects present just the amount of specialisation and diversity from con-
tinental forms which may be well explained by the known means of dispersal acting
through long periods; their land shells indicate greater isolation, owing to their admittedly
less effective means of conveyance across the ocean; while their plants show most clearly
the effects of those changes of conditions which we have reason to believe have occurred
during the Tertiary epoch, and preserve to us in highly specialised and archaic forms
some record of the primeval immigration by which islands were originally clothed with
vegetation.

Chapter VIII.
Evolution in Oceanic islands.

Ever since the high proportion of endemic organisms in oceanic islands and
especially the occurrence of systematically isolated genera were first noticed, an
explanation of this condition has been sought. Most authors have, as we have seen,
assumed that the islands never had been connected with a continent and that,
consequently, their entire living world had developed from a limited number of
ancestors carried across the water. Even if the islands had emerged as late as during
the Pliocene, the time was thought to have been sufficient for immigrants to get
transformed. It was a peculiar island world, the Galapagos Archipelago, which gave
birth to DARWIN’s theory of origin of species through variation and natural selec-
tion, and HOOKER, when dealing with a number of islands, was convinced that
DARWIN had given the solution to their problems and that their status strongly
supported his theory:

And if many of the phenomena of oceanic island Flora are thus well explained by
aid of the theory of the derivative origin of species, and not at all by any other theory,
it surely is a strong corroboration of that theory. Depend upon it, the slow but steady
struggle for existence is taking advantage of every change of form and every change of
circumstance to which plants no less than animals are exposed; and, variation and change
ot form are the rules in organic life.... By a wise ordinance it is ruled, that amongst
the living beings like shall never produce its exact like; as no two circumstances in
time or place are absolutely synchronous, or equal, or similar, so shall no two beings
be born alike; that a variety in the environing conditions in which the progeny of a living
being may be placed, shall be met by variety in the progeny itself. A wise ordinance

364 C. SKOTTSBERG

it is, that ensures the succession of beings, not by multiplying absolutely identical forms,
but by varying these, so that the right form may fill its right place in Nature’s ever
varying economy (Z7é. 11-1 2).

As we have seen already, HOOKER’s first point when summarizing the charac-
teristics of island floras was that also their most peculiar genera had their origin
in a mother continent, that no oceanic island had been a centre for special creative
forces. GRISEBACH (II. 494) may have been the first to express another opinion.
Referring to the systematically isolated endemics in St. Helena, he argued that they
never had had any allies elsewhere:

Nach diesen Thatsachen kann von einem Stammkontinent weder in dem Sinne die
Rede sein, dass die Flora von daher durch natiirliche Einwanderungen bereichert wurde,
noch als ob die endemischen Arten aus Umbildungen von Pflanzen hervorgegangen waren,
die in einer friiheren Periode von auswirts dahin gelangten. Nur von gewissen klima-
tischen Analogieen ist ihre Organisation der Ausdruck. S. Helena verhalt sich demnach
ganz verschieden von den Kap-Verden und liefert den vollgiiltigen Beweis, dass die Entste-
hung der Pflanzen auf Inseln, ebenso wohl wie auf Kontinenten, unabhingig von anderen
Vegetationscentren méglich war. Warum sollte auch der geographische Umfang eines
Gebiets auf die Krifte, welche die Organisationen erzeugt haben, von Einfluss sein? In
dem kleinsten Raume, wie im gréssten, konnten sie in besonderer Weise sich entfalten:
nur werden sie im ersteren Falle weniger zahlreich sein miissen.

In his discussion of the Galapagos flora he admits that what he terms vicarious
species, endemic in the islands, may have arisen through transformation of American
species, but he doubts it.

Diejenigen, welche annehmen, dass die vikariierenden Arten aus Umbildungen von
eingewanderten hervorgegangen sind, kénnen auch unter den endemischen Gewdchsen
der Galaipagos Beispiele genug anfiihren, dass eine nahe Verwandtschaft dieselben mit
Amerika, als ihrem vorausgesetzten Stammkontinent, verbinde. Aber allgemein lasst sich
dieser Gesichtspunkt nicht durchfiihren. Gerade unter den geselligen Holzgewachsen .
finden wir die eigentiimlichsten Erzeugnisse, die Scalesien und andere Synanthereen, die
nach ihrer systematischen Stellung dem Festlande ebenso fremdartig gegeniiber stehen
wie die Lobeliaceen des Sandwich-Archipels. ... / Alle Beredsamkeit, womit die Abstam-
mung der Vegetation ozeanischer Inseln von den Kontinenten vertheidigt zu werden
pflegt, kann die Thatsache nicht verdunkeln, dass in solchen Fallen [the genus Scalesza
etc. are mentioned] die Organisationen nicht anzugeben sind, aus deren Variation man
sie hervorgegangen vorstellen m6échte. Die nahe Verwandtschaft hingegen, welche zwischen
vielen endemischen Erzeugnissen des Archipels und denen der amerikanischen Floren
unleugbar besteht, kann aus dem Bildungsgesetz der riumlichen Analogien ebenso wohl,
als aus einem genetischen Zusammenhang abgeleitet werden. Und warum sollte tiberhaupt
das Festland vor den Inseln den Vorzug selbstindig entstandener Organisationen gehabt
haben, deren erste Erzeugung in den friihesten Perioden der Erdgeschichte jeder Méglich-
keit einer Variation vorausging? warum sollte sich nicht spiter und an verschiedenen
Orten sich wiederholt haben, was urspriinglich méglich war und wovon nur die Bedin-
gungen ein noch ungeldstes Rithsel geblieben sind? (II. Ls eels r3)3

Such ideas have little more than historical interest. DRUDE, another leading
authority on plant geography, was more in accordance with modern thought:

... dass die Flora der Inseln nicht nur als Transformationen der jetzt lebenden Kon-
tinentalfloren erfasst werden darf, sondern dass auf vielen Inseln unzweifelhaft eine Weiter-

DERIVATION OF THE FLORA AND FAUNA 365

entwickelung alter, vielleicht den Charakter einer ilteren tertiaéren Periode reprdsentieren-
der Stammfloren stattgefunden hat, welche sich hier im Schutze der Abgeschiedenheit
fern von dem Einfluss kontinentaler Umwalzungserscheinungen sich erhalten konnten

(305. 128).

DRUDE agreed with HOOKER that extinct species, known as fossils in IXurope,
were the ancestors of species now endemic in Macaronesia.

With regard to the Pacific islands several authors have discussed the question
to what extent endemism has been of a progressive kind and if not only species but
also isolated genera have evolved on the islands from a limited number of un-
known ancestors. Most of them, in spite of their firm belief that the islands are, geologi-
cally spoken, very recent, regard the insular biota as a local product; the first arrivals
had had time to give rise to new genera, those that came a little later became
new species, still more recent ones varieties of a continental species, and such as
arrived in our era have not had time to change but are expected to do so, because
isolation in a new and strange environment makes them adapt themselves by changing
their genetic structure. HAYEK (304. 252) expressed this very clearly:

Dass diese eingewanderten Elemente infolge ihrer Isolierung eigene Entwickelungs-
richtungen einschlugen, die sich in einer oft auffallend grossen Zahl von Endemismen
aussern, ist ja selbstverstandlich.

Among botanists of the latest decennia ANDREWS (6) may be chosen as a
representative of the school of “rapid adaptive radiation”’.

If a newcomer belongs to a primary form of a virile genus such as Acacza, Coprosma,
etc. it may be expected rapidly to become differentiated into varieties and species. As
Bentham pointed out long ago, the geographic station of a waif or colonist imposes
variations upon it almost from the moment of its arrival. Eucalypts planted in New
Zealand, California, and other places present marked differences from the forms the same
species possess in Australia. In the second place, if the plant assemblage into which
the waif or colonist arrives be a result of long-continued struggle for existence such
as occurs commonly in Holarctica and the cosmopolitan tropics, then the opportunity
for the development of new forms is remote, unless the new arrival itself is a plastic
form, and a grand example of the survival of the fittest. If, on the other hand, the
newcomer belongs to an agressive species in its own continental setting, then it has,
all other things being equal, an excellent chance of survival and of differentiating into
new forms (p. 617)..

ANDREWS, himself an Australian, speaking on the Hawaiian flora, mentioned
Acacia, Coprosma and other genera of Australian or southwest Pacific origin which
developed new, endemic forms in Hawaii, where they take a prominent part in the
vegetation. But when it comes to Eucalypts planted in California, nobody will, I
suppose, consider this example as a proof of the origin of new taxa under the
influence of new surroundings.

There is, he continues, a difference in the physical character of islands; some
did not encourage the colonist to vary and evolve, others did.

If the island... be very small and of negligible relief, it again has but little op-
portunity for differentiation, and, furthermore, if the island be even large and high, but
the time be short ...the response will have been but slight (l.c.).

366 C. SKOTTSBERG

If, on the other hand, the island group lies in the tropics and is large in area, the
islands of the group being close together, if the vertical relief be very great indeed ...
if the precipitation be very variable ...if the soil be rich, but variable in porosity; if
the plant assemblages into which the waifs or colonists entered are not the end result
of severe plant competition; then the stage is set for the rapid differentiation of primary
types of agressive genera.

The evidence available suggests that these genera never existed as such on the land
from where their immediate predecessors were derived, but that virile types, of the families
concerned, arrived as waifs from Malaysia, Australia, New Zealand, Central and North
America. Once they found themselves removed from their former severe competition
with other plants, they gave rise to the vigorous, endemic Hawaiian genera (p. 618).

What ANDREWS depicts is an island which has reached maturity. It has risen
from the bottom of the ocean to an altitude where the moisture of the trade winds
is condensed, where there are leeward and windward slopes, different habitats and
a rich soil. This island looks back on a very long history, but life begins to arrive
long before the island has come to rest. Aerial plankton will bring microscopic
green and bluegreen algae, bacteria, spores of all kinds and also air-borne seeds,
and some drift may be washed up on the shores. But to begin with there is little
humidity and hardly anything we can call soil, we need water before even the
most primitive organisms can exist, so that the island becomes fit to receive its
first settlers, microscopic algae, then mosses and lichens and mycelia of fungi, to
form soil where the first seeds can germinate and start to form an incipient vege-
tation cover and an abode for a soil fauna and flora. Lava cracks in cooling, some
water may stand in the fissures which form the starting point for further develop-
ment, a spectacle we have before our eyes where streams of lava are still formed.
Of higher plants, ferns are likely to be among the first to get established—GUPPY
even spoke of the “era of ferns’; a halophytic Asplexzum is the only living thing
observed on the far-flung reef Sala y Gomez between Easter Island and South
America.

An island in a comparatively recent stage, where there is plenty space for
new settlers, let it be that little comfort is as yet offered, would, we should think,
offer good opportunities for “virile and agressive’ immigrants to get a foothold
and to become the ancestors of the most ancient element in the flora, but this is
not what ANDREWS says. In order to start an evolution of new species and genera
a very great vertical relief, a variable precipitation and a rich soil are the conditions,
but in order to get a rich soil cover we must have a closed vegetation cover, also
forest. WALLACE (278. 295), speaking of St. Helena, very rightly said that “no
soil could be retained unless protected by the vegetation to which it in great part
owed its origin’, and the same is true everywhere. If the change in environment,
the new living conditions, are the cause of variation, why did they not act until
the island was already more or less stocked with plants? Were there no virile
species among the earliest immigrants which took possession of the land and formed
the oldest element of the flora? Was it lost among the later arrived aggressive
newcomers? It is calculated that about 90 % of the Hawaiian angiosperms are en-
demic, most of them belonging to endemic genera or to species very different from
their continental congeners. Do some of the most remarkable monotypical genera

DERIVATION OF THE FLORA AND FAUNA 367

represent true relics, descendants from the earliest settlers, whereas the other are
examples of progressive endemism? This is often said, but it is not, as far as I
can see, what ANDREWS means.

In the opinion of zoologists still infected with Lamarckian ideas environment
is the direct cause of new hereditary characters. Time after time we are told that
as soon as a ‘germ’ happens to land on an island, it gives rise to something
new. However, this does not, MAYR says, imply that every little island is turned
into a centre of evolution (779. 216).

The small and usually rather isolated islands of Polynesia have not only not been
new centres of evolution, like the Galapagos or Hawaiian Islands, but, on the contrary,
there is good evidence that many of them are “‘traps’’. Species that reach these islands
are doomed to extinction.

This is peculiar. There are many small islands stocked with both non-endemic
and endemic species, many of the latter stenotopic, it is true, but quite able to
hold their own as long the environment remains unchanged. It is a truism that,
if a newcomer lands on an island where, for climatic or other reasons it cannot
live, it is doomed to disappear pretty quickly, but there is no reason why, once
established and able to reproduce itself, it would become extinct as long as the
habitat does not undergo any change for the worse. It is generally known among
ornithologists, MAYR says, that island birds are very vulnerable. He continues:

The recent considerations of Sewall Wright have given us a possible key to this curi-
ous phenomenon. Apparently in these isolated populations there is more gene loss than
gene mutation. The species are therefore adjusted to an exceedingly narrow limit of
environmental conditions. They are unable to respond to any major change of conditions
and must die if such a change occurs, or are crowded out if competitors arrive.

We cannot be sure that mutations, should they be induced, would do them
any good and we need not assume a “gene loss” to explain why organisms unable
to escape to a more favourable habitat are bound to become exterminated as a
result of “major changes of conditions”.

ZIMMERMAN (298) is one of the prominent defenders of the idea that, in oceanic
islands, as exemplified by the Hawaiian chain, a small number of immigrants has
given rise to a comparatively rich fauna and flora; the proportion between genera
and species attains figures expected under continental conditions but certainly not
in oceanic islands. He admits that everything did not necessarily happen on the
present islands as we behold them, for differentiation may have begun on some
distant land and proceeded in the course of migration, with Hawaii as the terminus.
He picked out, as an example, a large curculionid genus ranging from Australia
to Micronesia and east to Marquesas; most islands or archipelagoes have their own
endemic species, which have developed on their respective islands. He thinks that,
if within a varying population, one pregnant female, not carrying the gene consti-
tution of the entire population, gets isolated on an oceanic island, she may stand
out as distinct from the average—and if this sequence of events ‘be accompanied
by conditions conducive to isolation and survival, rapid and diversified speciation
may follow’ (p. 125). This process is repeated on island after island, and ‘‘the

368 C. SKOTTSBERG

intensity of divergence will be increased’. It is not clear to me why the intensity
would be increased. Now, if sufficient time has elapsed and the original sources
have been eroded down and perhaps become sterile atolls, their faunas will have
been exterminated and on the newer islands segregates without obvious ancestral
relation will be left. This development explains why Hawaii has so many isolated
endemic types. The living world in Hawaii is older than the rock—in a way, he
says. Quite true, but it is not true that this possibility has, as he says, been entirely
overlooked in previous discussions; I think that I have, on repeated occasions,
expressed myself very clearly on this point, even if I do not agree with ZIMMERMAN
when it comes to explain why and how it happened.

The biota as we know it today is in part the ultimate product of a progressional
development which has moved and evolved along great insular archipelagos over periods
of time much longer than the ages required for the development of the main Hawaiian
Islands and their contemporary biota. Various genera and stem forms of groups of species
may have evolved in islands—now atolls such as some of the leeward Hawaiian chain,
the great Micronesian archipelagos, the Line Islands—which form the approach to Hawaii.
However, some of the genera and the bulk of species known today have originated on
our present main islands (p. 125) ...in contemporary Hawaii there are preserved rem-
nants of a biota which has in part developed by unique methods and in which are preserved
forms which are the end products of species chains that carry back, through archi-
pelagos now worn away, to geological ages indeterminant (p. 126).

In few words, we have to do with relict as well as progressive endemism—
nobody objects to that. A genus may be an ancient relic, while the actual species
are the result of more recent, progressive differentiation. On the other hand, there
is no reason why not a species could be immensely old without having undergone
any perceptible change.

ZIMMERMAN does not hesitate to conjure up all the sunk archipelagos he needs,
if only land connections are left out of the discussion. Once more he describes
his vision on p. 127 which I shall permit myself to quote, with the obvious risk
of tiring out the reader.

I believe that the great atoll chains of the Pacific may hold some of the now hidden
clues to the stories of the magnificent biological development of Polynesia. Many of the
peculiar endemic groups of the Hawaiian and southeastern Polynesian islands owe their
existence, if not their very origin, to ancient high islands of the one-time splendid archi-
pelagos marked by clusters or coral reefs. Surviving lines of middle Tertiary and of
perhaps even older continental, faunas may have had their germ plasm filtered down
through successively changing generations which have passed successfully through island
maturity and degradation to atoll formation and have carried over to new high islands
in different archipelagos. Thus, some supposedly old types such as certain land molluscs
could have maintained themselves (but evolving) in insular isolation through long periods

of time while their continental progenitors became extinct or restricted under continental
conditions.

I cannot think of what kind of higher organisms would have passed success-
fully through island degradation down to atoll stage; they must have left for new
high islands long before their abode became uninhabitable. Even if, as ZIMMERMAN
thinks, much of the evolution took place during migration from island to island,

DERIVATION OF THE FLORA AND FAUNA 369

specific segregation was mainly effected after arriving at the final station, in this
case Hawaii. This happened yesterday or the day before, geologically spoken:
“the rate of erosion is such that these main islands could not have stood here as
they are longer than from a period late in the Tertiary” (p. 121). “Explosive
speciation’ set in during late Pliocene and must have increased during “the great
Pleistocene erosion which has left such a spectacular and rugged topography in
its wake. New land open to colonization is conducive to speciation’”’ (p. 122).

This late and rapid differentiation is illustrated by ZIMMERMAN for the land
snails pp. 98-101:

Helicidae. 59 species developed from one, or possibly two original immigrant
stocks.

Pupillidae. Possibly 4 ancestral species gave rise to the 86 Hawaiian forms.

Cochliocopidae-Cochliocopinae. One immigrant of Cochliocopa stock could have
given rise to the 142 forms.

C.-Amastrinae. 294 forms apparently developed from one basic stock.

Tornatellinidae-Tornatellininae. 117 forms derived from 4 or fewer ancestral
forms.

T.-Achatinellinae. It appears certain that this subfamily had its origin and
development in the Hawaiian area and all of the 215 forms may have been derived
from a common tornatellinid ancestor.

FOSBERG, in a chapter contributed to ZIMMERMAN’s book, tried to fix the
number of ancestors of the Hawaiian angiosperms. His method is quite simple:
if the species, few or many, of a certain genus present the appearance of a more
or less homogeneous group, only a single ancestor is made responsible for the
segregation; if subgenera or sections are distinguished, we must count with the
same number of ancestors as of taxonomic groups within the genus. The method
seems a little too easy; possibly we are confronted with a rather complicated
question, the solution of which I am not going to attempt.

SETCHELL (278) uttered some sensible words on migration and endemism:

Where endemism of the degree of ordinal or family endemism occurs on oceanic
islands, we may feel strongly inclined to believe that evolution of such degree took
place on the continental area which was the source of the original migration and not
on the island where now found, the original becoming later extinct, leaving the migrant
as an endemic. The same is true of generic endemism or even specific endemism of a
strong type, that is when representing an isolated or aberrant species under the genus
(p. 874).... To assume that insular conditions originate new forms is to overlook what
has taken place on continents (p. 875).

This is, however, what so many authors do. They claim that oceanic islands
follow their special laws, that a plant or an animal which happens to land far away,
will, as it were, lose its balance; hidden factors, repressed as long as they lived
on their fatherland under “‘severe competition’, are set free and allow them to
develop their inherent possibilities, they are not subjected to any struggle for
existence in their new environment. Says ZIMMERMAN: ‘The environment, of course,
plays an all-important part in the development of species .. . it is generally agreed
that profound changes have been effected on organisms by environment” (p. 187).

24 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

370 C. SKOTTSBERG

This is pure Lamarckian language. We may say, of course, that change of environ-
ment may cause mutations, but our experience tends to prove that only a fraction
of 1% Age valuable, the remainder, if not deleterious, at least indifferent. Mutations,
gene losses, hybridization, polyploidy and so forth, all may have their share, but
they are only ripples on the surface. No theory has been able to penetrate to
the nucleus of the problem. Anyhow we have no reason to think that, in its pro-
duction of species and genera, families and orders, Nature has followed other lines
in islands than in continents. A prominent Swedish geneticist, after a life-time’s
speculation on the causes generally accepted as responsible for the “‘origin of species”’,
rejected all of them; unfortunately he threw the egg away with the shell and con-
vinced himself that there had been no evolution at all. He did not, however,
revert to an omnipotent creator but invented a new and entirely revolutionary theory;
if there ever was a stillborn one, it was this (zJ6).

I guess we can take it for granted that no peculiar, outstanding types were
created on young volcanic islands, whatever their present faunistic and floristic
status may be like. Their basic stock is much older than the rocks and inseparably
connected with the great continental faunas and floras. Wallace, however, made a
distinction between island and continental history. He emphasized that evolution
has required an enormously long time to produce the present status, that from
the Cretaceous until now nothing of a revolutionary character had happened, families,
genera and in cases even species still living date from early Tertiary at least—
but on islands the great period of creation was repeated during the last epochs.

Finally I shall quote some selected passages from STEBBINS’ book on variation
and evolution (257).

The differentiation of orders and families of flowering plants through the action of
natural selection under present conditions is well-nigh impossible. ... All the trends
leading to the differentiation of families of flowering plants probably took place simul-
taneously and at a relatively early stage of angiosperm evolution. For instance, both
distributional and paleontological evidence indicates that the Compositae, the most highly
specialized family of dicotyledons, already existed in the latter part of the Cretaceous
period, and distributional evidence indicates a similar age for the most advanced families
of monocotyledons, the Orchidaceae and Gramineae. It is likely, therefore, that the
major part of angiosperm evolution, involving the principal trends in the modification
of the flowers, took place during the Mesozoic era (pp. 501-502).

The gymnosperms prove, he remarks, that neither great antiquity nor rigid
stability necessarily leads to senescence, but here, as in other cases, a stenotopic
character has resulted in restricted areas and also in extinction.

Isolated records excepted—the latest discovery is a palm from a Triassic stratum,
if correctly determined

angiospermic fossils are found in greater quantity from
younger Cretaceous and then in many surprisingly modern types. Evidence is strong,
STEBBINS has found, that evolution was rapid during one period and slower during
another. At the end of the Cretaceous and the beginning of the Eocene the number
of modern types increased rapidly. In eocene deposits in North America and Eurasia
“the majority of the species belonged to or closely approximated modern genera”’
(p. 520). This conclusion, mainly based on leaf impressions, is strengthened by the

DERIVATION OF THE FLORA AND FAUNA 371

famous London clay from lower Eocene, where a wealth of fruits and seeds were
found. The corresponding fossil floras of California include an increasing number
of plants that cannot be distinguished from living genera, and abundant evidence
shows that since late Pliocene “new species of woody plants have been added to
the Californian flora only in such large and complex genera as /:rzogonum, Ceanothus,

Arctostaphylos, and various Compositae”’ (p. 521). The fossil floras of East Asia point

in the same direction, the rate of evolution on woody plants has, since the middle
of this period, been as slow or slower.

The most likely inference on the basis of all available evidence is that most of the
woody species of to-day have existed for five million years or more, and that the evolution
of the genetic isolation mechanism separating them took place largely during the early
and middle parts of the Tertiary period (p. 522).

With regard to the herbaceous floras there is evidence for a rather rapid evo-
lution during the later Tertiary. STEBBINS quotes the results obtained by ELIAS
in fossil caryopses dating from lower Miocene to middle Pliocene and showing a
distinct progression until the most recent ones cannot, to judge from the illustrations
redrawn by STEBBINS, be distinguished from St#pa or Piptochaetium, an evidence
from palaeontology which strengthens the theory of SINNOTT and BAILEY already
referred to.

Tsolation as a cause of evolution.—There is no reason why a species, which
has migrated to an island, should change and give rise to new forms only because
it is spatially isolated. A form of a varying population which gets isolated on an
island, may possess characters making it stand out as a more or less well-marked
form. As JORDAN (257, 752) says, isolation is not the direct cause of the origin.
Spatial relation, STEBBINS remarks,

may persist over long periods of time without causing the isolated populations to diverge
from each other enough to become recognizably distinct or even different. ... the nature
of many distributions strongly suggests that some of these disjunct segments of the same
species have been isolated from each other for millions of years. There is some reason
for believing, therefore, that geographic isolation alone does not result even in-the
formation of subspecies....In small populations, which are particularly frequent on
oceanic islands, spatial isolation is the usual precursor to divergence in non-adaptive
characters by means of genetic drift or random fixation (p. 197).

STEBBINS also points out that spatially isolated races or species usually are
separated also by ecological barriers, because different areas also differ ecologically;
distance may not be the original isolating factor, separation and differentiation may
be due to the selective effect of ecological and climatic factors before geographical
separation occurred. The Hawaiian Islands offer a wide field for a study of isolation
and segregation resulting in local endemism (242). The continental flora inherited
by a Great Hawaii included numerous populations gradually taking possession of
the new soil, a process of very long duration, Volcanic activity progressing from
west to east, and when separation took place, different forms and species became
isolated on different islands. The wonderful systems of valleys, effectively separated
by high and very steep ridges, furthermore promoted segregation. Little has been

272 C. SKOTTSBERG

done to examine the genotypical constitution of this astonishing assemblage of
local forms; some may, for all we know, be ecotypes, but experimental studies
have not been made. A limited number of hybrids have been described, but we
know nothing of their behaviour. Anyhow, the floristic difference between the islands
is very striking and hardly in favour of the theory of transoceanic migration, for
the majority of organisms seem to have been unable to cross even the straits
separating the islands, and biotic factors preventing establishment can hardly be
made responsible in so many cases.

Chapter IX.

Juan Fernandez—oceanic or continental ?

To WALLACE as well as to the majority of biogeographers the Juan Fer-
nandez Islands were typically oceanic in spite of their moderate distance from
South America. They had the advantage of antiquity, WALLACE remarks, for
the means of transmission had formerly been greater than now, their surface was
varied, soil and climate favourable, “offering many chances for the preservation
and increase of whatever plants and animals had chanced to reach them” (p. 287).
Had the character of Masafuera been known to him he might have been less
optimistic. The land-shell fauna, entirely endemic, testified to the great age of
the islands, for none had been introduced for so long a period that all which
did come had given rise to new forms—or were the last of a fauna extinct
on the continent.

JoHow (z50) based his opinion on WALLACE; when discussing the dispersal
agencies and the morphology of the diaspores his starting-point was /wo islands,
Masatierra plus Santa Clara and Masafuera, which has risen separately from the
deep sea. It is strange that he never thought of another possibility, because the
geologist who went with him and who wrote a chapter on the geology of Masa-
tierra, claimed to have discovered, in one place, a fundament of rocks older than
the omnipresent young basalt—JOHOW could not know that the interpretation
of this stratum was false, as later shown by QUENSEL (302). After his visit to
the Desventuradas JOHOW modified his opinion; the submarine ridge uniting these
little islands with Juan Fernandez had then been discovered:

Die unter gleicher Breite mit dem Hafen Caldera und in derselben Entfernung
vom Kontinent wie Juan Fernandez gelegene Inselgruppe ist vulkanischen Ursprungs
und stellt, wie die von dem Mitglied der Expedition, Herrn Chaigneau, ausgefiihrten
Lotungen ergaben, die iiber Wasser befindlichen hdéchsten Gipfel einer im Ubrigen
unterseeisch verlaufenden Bergkette dar, welcher auch die Inseln der Juan-Fernandez-
Gruppe als siidlichste Gipfel angehéren. Aus dem Vergleiche der Floren und Faunen
beider Archipele, welche trotz der grossen klimatischen Verschiedenheiten frappante
Verwandtschaft aufweisen, ergiebt sich mit zwingender Notwendigkeit die Hypothese,
dass die zwei Inselgruppen in der Vorzeit mit einander in Landverbindung gestanden
haben und dass ihre Isolierung die Folge einer stattgehabten Senkung jener Bergkette
ist (330. 259).

DERIVATION OF THE FLORA AND FAUNA 3/3

In this assumption he may be quite right, but it finds, as already pointed
out by REICHE (203. 269), little support in the flora, for the two groups have
only a single species in common (247), and the endemic genera and species are
very different. JOHOW’s belief in their permanent isolation from the continent
remained unchanged.

In his tables, Continjente A and B, pp. 218-220, is a column indicating
(not always with sufficient accuracy) the nature of the fruits or seeds, and another
stating the probable dispersal agent. Of the 143 vascular plants known at that
time, all the ferns and 34 phanerogams were supposed to have been wind-borne,
61 had been transported by birds, either inside or adhering to their feet or
plumage; only one, Sophora “tetraptera’, with winged pods, had drifted with
the current. Five |Aryxgium bupleuroides and sarcophyllum, Apium fernandeszianum,
Colletia spartoides and Fagara (Zanthoxylum) mayu\ offered too serious difficul-
ties to make their presence in the islands explicable, but I cannot find that they
are more ‘impossible’? than many of the others. In an earlier paper (235) I sur-
veyed the nature of the diaspores; I shall not return to this question here but
only repeat that almost one third of the flowering plants show no special adap-
tation to any particular mode of dispersal across the water, and I shall add a
few remarks on some knotty cases. Among the island Compositae, a family known
to be well adapted to wind dispersal, are several species which lose their pappus
when the achenes are still enclosed in the involucre, or where it is reduced to
uselessness. Only one species of Sophora, called tetrapfera, was recognized by
JOHOW and recorded from Chile, Juan Fernandez, Easter Island and New Zealand.
Genuine /e/rapiera is restricted to New Zealand, but this is of less importance,
the pods have narrow wings (poorly developed in some forms) which were sup-
posed to help the pods to keep afloat, but as they open on the tree and drop
their rather heavy seeds to the ground, the wings serve no purpose. Halorrhagis
is a somewhat similar case. JOHOW, and others before and after him, identified
the plant found on Masatierra with //. erecta (alata), a New Zealand species with
four narrow wings on the fruit, and he did not hesitate to regard it as wind-
borne. His evecta is, however, an endemic species; there are two more on Masa-
fuera (see 229, with illustrations), and their fruits are quite or almost unwinged.
There can be no question of an adaptation to wind carriage. The same applies
to Selkirkia, another of JOHOW’s anemochorous plants. Edible fruits, if not too
big, will be swallowed by birds, but the indigestable stones and hard-coated
seeds will be dropped before the islands are reached, and the diet of the wide-
ranging and fast-flying sea birds is another. Epizoic transport is possible in a
small number of cases, but otherwise the dispersal mechanisms, many of them
quite wonderful, serve to maintain the population within its range, and not to
stock distant islands.

Other advocates of the permanent isolation of Juan Fernandez are e.g. PLATE
(799), GOURLAY (3oz), BURGER (gz), and GOETSCH (724). PLATE drew his con-
clusions from a comparison between the littoral faunas of Juan Fernandez and
the opposite coast:

C. SKOTTSBERG

Os
~I
aS

Die Seichtwasser-Fauna weist deutlicher als die geologischen Verhaltnisse darauf
hin, dass Masatierra in der That als eine ozeanische Insel anzusehen ist, die nie in
Zusammenhang mit dem Festland gestanden haben kann... (p. 228),

but this did not prevent him from believing that Masatierra and Masafuera once
formed a single large island, in spite of the distance (92 miles) and the deep
water that separates them (p. 222). No more will be said here about a connection
with the mainland; I have already discussed this question from an algologist’s
viewpoint, and with a different result (238).

The reason why GOURLAY rejected all land connections was his conviction
of the easiness with which plants and animals are transported; the under-title of
his article reads “Plants make ocean voyages’. He had little faith in the birds,
wind and sea were the principal means of transportation, and as so many others
he pointed to Krakatau as the classical example. The distance was very small,
it is true, but winds and currents were favourable along the coast region of
South America, and the flora was derived from southern Chile. Of the genera and
species quite without relations not only in Chile, but in all America, he said
nothing.

BURGER, who had visited the islands, followed JouNOW; they were typically
oceanic. After telling us about some of the most remarkable endemics, he exclaims:

Wer brachte diesen entlegenen Stitten solch késtliches Geschenk? Die Strémung,
die Vogel und vor allem die Winde. Sie beluden sich mit Sporen und Samen... .
selbst vom Feuerlande empfangenes gedieh. Doch aus viel weiteren Fernen kamen die
Einwanderer, auch vom tropischen Amerika, Polynesien, ja sogar von Australien und
Neuseeland und den Inseln des Indischen Ozeans (pp. 17-18).

The majority of the newcomers remained true to their stock in spite of the
changed conditions, but others changed and some took such a fancy to the
climate that from being herbs they became trees. I shall leave these speculations
without comment; we have better reason to observe GOETSCH’s paper which,
in spite of being fairly recent, contains many amazing statements not supported
by facts. Ignoring QUENSEL’s report on the geology, he tells us that the islands
rest “auf einem Sockel von griinlichem Andesit, der auch die Hauptmasse der
Anden bildet’. He had seen my writings on the flora; the use he made of them
may be illustrated by a couple of examples. Among the plants introduced from
Europe he mentions ‘‘“Aromo aus Castilien’’—this is a local name for the en-
demic Azara fernandeztana! He rejects all land connections, the sandal-wood
came from the East Indies; he had not observed that it belongs to a quite dif-
ferent section than Santalum album. The arborescent Compositae and Plantago had
originated in the islands, “Plantago fernandesia und Skottsbergi7, 1-2 m hoch mit
20 cm langen und 3 !/, cm breiten, an der Spitze des Stammes stehenden Blat-
tern’. He overlooked that the so-called P. Skottsbergii is a modest annual and
a form of the common Chilean ¢7wzcata, a variable species. But GOETSCH knows
what happened in the islands: ‘Dass der europaische Wegerich auf Juan Fernandez
meterhohe Bliitenahren und fusslange Blatter tragt, weist auf die Entwickelung
hin, die seine Verwandten einstmal nahmen” (p. 29). This refers to P. lanceolata,

DERIVATION OF THE FLORA AND FAUNA 375

which’ luxuriates in the islands, but cannot prove that the arborescent P. fervan-
dezia evolved from one of the many Chilean species with which, from a system-
atic viewpoint, it has nothing to do.

GOETSCH was, perhaps, more familiar with zoology, but where he got the
impression that the lower fauna was extremely poor, | cannot tell, unless JOHOW’'s
meagre list was his only source. A considerable number of land shells and in-
sects had been described, and everything tended to show that they represented
but fragments of the fauna. The most efficient dispersal agent was, in this case,
the current.

Wir wissen, dass die warmen StrOmungen Ozeaniens bis an die Insel reichen
und australische Pflanzen und ‘Tiere verfrachten ko6nnen. Aus diesem Grunde muss
zwar die Insel auch ausserhalb des sog. Humboldtstroms liegen, der von der Antarctis
bis zum Aequator und der siidamerikanischen Westkiiste lauft. Wir wissen aber nun
seit neueren Untersuchungen, dass dieser sog. Humboldtstrom keineswegs ein conti-
nuierliches Fliessen in noérdlicher Richtung ist, sondern dass Oberflachen-Wasser vom

Lande wegstromt und durch kaltes aus tieferen Schichten ersetzt wird. Der chilenische
und patagonische Einfluss ist dadurch gesichert.

From what he just said we learn that the Humboldt current, whatever it is,
and the upwelling cold water do not reach as far as Masatierra. On the other
hand, it is not true that, as GOOD says (zog. 220), Juan Fernandez lies entirely
outside the Pacific beach drift. Drift-wood is found on the western side of Masa-
fuera, but it shows very rough handling and must have been months or rather
years under way, and if one has seen the place one feels convinced that not a
single plant owes its presence in the island to this mode of conveyance, not to
speak of the fauna. But GOETSCH believes that also the tropical element in the
flora has arrived by sea; he continues:

Wir wissen endlich, dass vom Norden her dann dem Humboldtstrom eine warme
MeeresstrO6mung entgegenwirkt, der sogenannte Nifio, der sich periodisch in einzelnen
Jahren so geltend machen kann, dass seine Wirkung bis Valparaiso gespiirt wird... .
Damit findet auch der tropische Einschlag in Fauna und Flora seine Erklarung (p. 38).

This cannot refer to the warm currents of Oceania mentioned before, because
they came from Australia and did not bring the neotropical element. Does he
think of the equatorial counter current which, occasionally, would extend its
influence to the South American coast and become deflected south, thereby
fetching new passengers? This current does not reach the coast.

Other authors have acted as spokesmen for the bridge-builders. W. A. BRYAN—
not having access to his book I quote a newspaper article written by him (300)—
believes that a “prehistoric continent’’ once embraced the Pacific islands, a theory
based on the distribution of the land-snails. Juan Fernandez belonged to the
same geological period as Hawaii and their living world proves that at some
time they were united. The general biological character is, he says, the same,
but there is a slight difference in the species. For details I refer the reader to
261, where I have reviewed his book. BRYAN was professor of zoology and geology
in the University of Hawaii and had visited Juan Fernandez, but he does not
appear to have profited very much by his visit.

376 C. SKOTTSBERG

Where BRANCHI (303) got his idea of the island flora I cannot tell, in any
case not from facts, in spite of having visited Masatierra:

Como curiosidad cientifica se puede decir que Masatierra no seria sino una de
las tantas vetas del gran continente sumergido en el Océano Pacifico en €pocas en
que el continente sudamericano era un lecho de mar, y por consiguiente la isla serfa
mds vieja que el suelo de la madre patria. Como comprobacién esta la ausencia de
los volcanes, los esarpados farallones que muestran las capas geolégicas, el desliza-
miento de las rocas de la época glacial, y sobre todo la flora unica, diversa del conti-
nente y que puede sdlo acercarse a la flora polinésica y australiana (p. 1a)

I guess we can agree with him that this is all very curious, but it is not
scientific.

To complete the picture we have better consult ARLDT’s great work (8)
in which the author draws his conclusions from the land fauna. ARLDT was one
of the great bridge-builders, but his Pacific bridges were of old date, and he
thought that Juan Fernandez had been isolated since late Cretaceous times, so
that only the most ancient element was supposed to have arrived over land (p. 322).

Wie Tristan da Cunha einen Rest des Siidatlantis darstellt, so die Juan Fernandez-
Gruppe einen der Ozeanis der siidpazifischen Landbriicke. Das beweisen nicht sicher
die Vogel... auch nicht die Schmetterlinge, von denen die Nymphaliden durch die
chilenische Pyrameis carye vertreten sind, die Pyraliden durch Scoparia ragonoti,
neben denen noch weitere chilenische Formen genannt werden. Alle hatten auch trans-
marin die Inseln erreichen kénnen. Wichtiger sind die Landschnecken, deren Gattungen
fast alle auch in Chile vorkommen. Vertreten sind z. B. die Succineiden und Ferrusa-
ciden, sowie die Endodontiden. Diese sind hier mit AmpAzdoxa vertreten. Deren typische
Untergattung ist auf Juan Fernandez endemisch. Stephanoda besitzt 6 endemische Arten
auf den Inseln, andere auf Chiloé, in Chile, Patagonien, Feuerland, Argentinien, Para-
guay, Brasilien und eine auch auf Kerguelen. Dagegen gehort die auf Juan Fernandez
endemische /ernandezia zu den sonst hawaiischen Amastriden, die auf Hawaii auch
ihre einzigen néiheren Verwandten in den Achatinelliden besitzen. Hier kann nur eine
Ausbreitung iiber Land angenommen werden. Von anderen Landtieren findet sich auf
Juan Fernandez z. B. noch der Tausendfiissler Geophilus laticollis. Endlich sind sie auch
von einem Oligochaeten erreicht worden, der Ocnerodrilinen Kerria saltensis, die auch
in Chile vorkommt. Hier wire eine jiingere, iiberseeische Einwanderung denkbar. Aber
Kerria ist auch alt genug, dass sie auf dem Landwege nach Juan Fernandez gekommen
sein kann. Auch die Flora der Inseln zeigt interessante Beziehungen (pp. 324-325).

As could be expected, ARLDT’s south Pacific bridge was supported mainly by
the land molluscs, and they made him extend his bridge across the equator to
Hawaii. There is in the Juan Fernandez flora some perplexing affinities with
Hawaii that seem to defy all attempts to an explanation. The occurrence of two (not
only one) endemic species of Vesogeophilus (formerly a subgenus of Geophilus) is in-
teresting, Aevrza is, perhaps, less important. Other striking cases of disjunction were
already referred to in the lists of evertebrates above. It is a pity that ARLDT over-
looked the leech NVesophilaemon, but also from what he says about the butterflies it is
evident that the numerous zoological papers forming vol. II of “The Natural History
of Juan Fernandez and Easter Island’’ had escaped his notice. The leech would,
I suppose, have furnished him with one of the most eloquent evidences of
former land connections. Nor did he know the botanical volume of this work,

DERIVATION OF THE FLORA AND FAUNA 377

for he gives a summary of my 1914 paper (227). He agrees on the great age
of the ‘‘Palaeopacific element’; second comes the smal] neotropical, third the
large Chilean group: “hier liegt also wohl zum grossen Teil spate, iiberseeische
Einwanderung vor’. This was, he found, even more true of the subantarctic
element, ‘das iiberhaupt keine endemische Arten aufzuweisen hat’—very few
were known then, but later some endemic species were discovered.

The question to which extent Juan Fernandez presents the biological pecu-
liarities regarded as characteristic of oceanic islands—see Chapter VIJ—will now
be answered. We have seen that exdemzsm is very high among the phanerogams
and that the various kinds distinguished by HOOKER are represented; we have
even a primitive endemic family, Lactor7daceae, we have a proportionately large
number of peculiar genera, some of them quite isolated, particularly among the
Compositae (Cextaurodendron, Dendroseris, Hesperoseris, Phoentcoseris, Rea, Rhe-
tinodendron, Robinsonta, Symphyochaeta, YVunguea) but also in other families,
Cuminia (Labiatae), Fuanza (Palmae), IWegalachne and Podophorus (Gramineae);
further there are a few endemic genera closely related to South American ones,
Nothomyrcia (Myrtaceae), Ochagavia (Bromeliaceae), and Se/kirkia (Boraginaceae).
As among the genera, so we find species of a strong character in Chenopodium,
Coprosma, Eryngium, Euphrasia, Fagara, Peperomia, Plantago, Santalum, Urtica,
Wahlenbergia, well-marked but not very aberrant species in Lerberzs, Boehmeria,
Carex, Cladium, Colletia, Evigeron, Escallonia, Gunnera, Halorrhagis, Hespero-
greigia, Ranunculus, Rhaphithamnus, Solanum and Ugni, and many not very
different from their continental congeners in Adrotanella, Acaena, Apium, Azara,
Cardamine, Chusquea, Drimys, Dysopsis, Galium, Luzula, Margyricarpus, Myrc-
eugenta, Pernettya, Phrygilanthus, Sophora, Spergularia and Uncinia. Species
undoubtedly native but also found elsewhere do not number more than 46,
and in several cases their citizenship is open to question. Endemism among
the ferns is not so high, but there is one very aberrant genus (7/yrsopieris) and
several peculiar species. Bryophytes and lichens will not be considered; they were
not included in HOOKER’s paper and I have not had occasion to compare
them with other island floras.

There are no conifers, a single leguminous genus (Sephora) with two species,
and no orchids. The proportion genus: species is 1: 1.65. Mammals, batrachians,
reptiles and fresh-water fishes are absent. The earth-worms are supposed to be
adventitious with the possible exception of Aevrza saltensis. In all these respects
Juan Fernandez agrees with the character attributed to oceanic islands.

The high percentage of woody plants was emphasized by SINNOTT and
BAILEY, but as the literature on which they based their figures is quite out of
date, a new table was prepared. .

The object of SINNOTT and BAILEY was, as we have seen above, to show
that the woody plants increase in number with the rising degree of endemism,
and that the endemic genera were trees or shrubs and formed a more ancient
element than the herbaceous plants which during former epochs were few in

378 C. SKOTTSBERG

Table V7.

Percentage of woody and herbaceous species in Juan Fernandez.

| Total} Woody % |Herbaceous %

| Indigenousrspecies: 7. rants. cal) LAY, 68 46.3 79 53-7
| Not endemic ........ | 46 5 10.9 Ae | OO
| Endemic, genus not endemic. . 71 Bi 52.1 34* | 47.9

Also the genusendemic . .. . 30 27 90.0 3 10.0

* Four species suffruticose.

comparison, and the Juan Fernandez Islands were regarded as supporting their
hypothesis; this, to judge from the table, they certainly do. The five woody
non-endemic species are Lmpetrum rubrum and Salicornia fruticosa (low, erect
shrubs), and JMyrteola nummularia, Rubus geoides and Calystegia tuguriorum
(trailing).

The rosette tree form is observed in 16 genera with together 31 species,
belonging to six families, Boraginaceae, Bromeliaceae, Chenopodiaceae, Compositae,
Plantaginaceae and Umbelliferae; the palm ¥uwanza is of course excluded. As |
have paid special attention to them in another paper (257), where they were
well illustrated, no more will be said here.

Annual and biennial herbs, the therophytes of RAUNKLAR, are not completely
lacking, but they are very few: Cardamine chenopodiifolia, Chaetotropis (2 species),
Partetaria, Plantago truncata, Tetragonia and Urtica Masafuerae. In 257 1 listed
Chaetotropis among the hemicryptophytes; they give the impression of lasting
more than one year and are found green at all seasons, and the same may be
true of Paretaria. | am not at all sure that Plantago truncata is native. The
Cardamine has been seen twice, last time in 1872, the Uréca not since 1854,
when it was discovered. It may be that it is an ephemerous plant and disappears
in early spring, a season when very few botanists have visited the islands.

Thus, the “oceanic peculiarities’ are all there, but possibly some of them
can be explained otherwise. Endemism of a very high degree within a small
area is no monopoly of isolated islands; it will be sufficient to mention the Cape
flora or southwestern Australia. The large proportion of woody plants can be
understood if the islands became isolated before the myriads of herbs, particularly
the annuals of Central Chile, had evolved. GRISEBACH's ‘‘Clavija’’ and related
life forms are not confined to oceanic islands; if my interpretation of this mor-
phological type as an evidence of antiquity is correct, we can understand why
it takes such a prominent part in old island floras. The reason why therophytes
are almost wanting is not climatic. This is amply proved by the innumerable
annual weeds introduced with the traffic and thriving only too well. The climate
is of a modified Mediterranean type, and from a purely climatic viewpoint we
should expect a large percentage of native annuals and biennials; this question
was discussed at some length in 257. 827-830. Chile has hundreds of endemic

DERIVATION OF THE FLORA AND FAUNA 379

therophytes, but they have not spread to the islands although many have special
dispersal mechanisms, and they were not available at the time when the supposed
connection with the mainland existed before the final uplift of the Andes.

The existence of many aberrant genera and species and, above all, the
marked difference between the phanerogams of Masatierra and Masafuera shows
that there is no exchange between the two islands in spite of the very moderate
distance; even the 360 miles separating Masatierra from Chile should, in the
eyes of the diffusionists, amount to little if sufficient time be granted. The Mar-
quesas Islands, situated much farther away from the continent, have a less peculiar
angiospermous flora than Juan Fernandez; the very opposite ought to be expected
if overseas migration had played the dominant rédle. The absence of the flora
of Central Chile speaks against the efficiency of the natural dispersal agents.
These circumstances are in favour of the opinion that the volcanic islands arose,
not from the depths of the ocean, but on a piece of land formerly connected
with South America and not sunk until the newborn islands, now reduced to
ruins, had become a refuge for the ancient continental fauna and flora.

It is also true that several large and widespread families are lacking, such
as leguminous plants (with one exception) and lilies, well developed in Chile,
but they belong to the modern Chilean flora.

The plant world of oceanic islands is described as a haphazard collection
of waifs and strays, and this is said to explain why so few genera contain more
than a couple of species. But would not the result be the same if the actual
islands originated through volcanic activity on a sinking land? Chance would
decide what took possession of the new soil, and different sets find a refuge
on Masatierra and Masafuera.

We know that of the mammals introduced by man the goat thrives and
multiplies since 400 years and quickly became naturalized. There were no goats
on the mainland when the land-bridge existed, but there may of course have
been some primitive mammals; if any of them reached the islands, they have
disappeared long ago. The islands, as we see them, appear never to have offered
great possibilities for the subsistence of a mammalian fauna. They are very small
and as there was little open land there cannot have been any grazing grounds
worth mentioning before man altered the landscape. Even after the ground has
been cleared on all the lower slopes, pasture is miserable, and one valley after the
other has been turned into a desert by the ravages of sheep and cattle. If left
to run wild and multiply, the final result can be foreseen. Carnivorous animals
need a prey, and there was none. The same is true of snakes which should
thrive well now since the domestic rats and mice have been introduced.

Chile’s mammalian fauna is poor and nobody knows if the huemul, the pudu,
the Chilean rodents and small marsupials would be able to make a living on
Juan Fernandez. Reptiles are poorly represented in Chile, there is not a single
tortoise, very few snakes and a dozen lizards. Amphibians are few, but toads
and frogs occur. Among the invertebrates are several orders, the chances of which
successfully to get transported across the sea are very doubtful or, as far as we
can see, none at all.

380 C. SKOTTSBERG

The insular peculiarities as displayed in Juan Fernandez (and in other similar
cases) do not, I think, permit us to take a definite position against the hypo-

thesis of former land connections.

Chapter XI.

The Chilean coast line and the history of the Andes.

From a look at the map we easily get the impression that the trend of the
South American west coast is a product of the rise of the Andes, because this
enormous uplift must have been compensated by the submergence of old border
lands and by the formation of a deep trench, and that these movements, which
certainly were of very great magnitude, may have extended its effects west as
far as to the region where we find the Juan Fernandez and Desventuradas Islands
forming the exposed summits of a submarine ridge. I shall call this ridge the
Chaigneau Ridge after the Chilean navy officer who was the first to survey it.

The Chaigneau Ridge.—The two archipelagoes lie within the 2000 m line,
San Felix and San Ambrosio on a plateau rising above the 400 m curve and
extending a long way toward Juan Fernandez, as seen from CHAIGNEAU's table
(52) which is reproduced here with the soundings rearranged from N. to S. accord-
ing to latitude and with the addition of some figures from the latest chart.

The Merriam Ridge.—i160 miles NW. of San Felix-San Ambrosio another
ridge, called the Merriam Ridge, was discovered during the U.S. “Carnegie”
campaign 1928-29 (zz). It extends between 25° 3’.2 S., 82° 20’ W., and 24 54’S.,
82°13’ W.; the depths found were 1445 and 1260 m, respectively. The bank
rises 3000 m above the bottom and is only 10 miles wide. Along the most
elevated part 1186, 1188 and 1168 m were found. From the latter spot a series
of soundings was taken SE. of the ridge, showing the rapid increase of the depth:
3 miles 1260 m, 9 miles 2751 m, 20 miles 3620 m, and 32 miles 4115 m; the
Merriam ridge is separated from the Chaigneau ridge by deep water. Toward
WNW. the slope is more gradual until a depth of 3000 m is reached.

North of the Merriam ridge, in 21° 4o’ S., 81° 40’ W., approximately, a sudden
rise, bounded by the 2000 m curve and surrounded by deep water, has been
discovered (see map). Here the bottom rises to 972 m below the surface. I do
not know if this remarkable place has a name.

The Carnegie Ridge.—During the cruise of the ‘‘Carnegie’”’ two soundings
about 100 miles off the coast of Ecuador, lat. 1°32’ S., long. 82° 16’, gave 1515
and 1454 m, respectively, indicating a rise of 1800 m above the bottom, but
before the entire distance along the coast has been surveyed we do not know if
a series of ridges, probably much less well-marked than the Chaigneau-Merriam
ridge, can be traced all the way between lat. 35° and the equator.

Between these ridges and the coast is the deep trench (greatest depth
7635 m), and south of the latitude of Juan Fernandez depths exceeding 5000 m
are still found, but farther south the 4000 m curve is soon reached and the trench
disappears. Older maps, two of which were reproduced in 227. 44, 45, show

DERIVATION OF THE FLORA AND FAUNA 381

Table VI.

The Chaigneau Ridge.

r=rock, s=sand.

eee, im Bottom S. lat. W. long. | Remarks
38 — 26° Lay 79° 56 14 km NE. San Felix
80 — 26° 15- 80° 4°.5 | 4.5 km N. San Felix
69 aes 26° 17.5 | 80° 8'.5 | 4 km W. San Felix
179 =a 2OP TS, 5 Ory Sr 5 km W. San Felix
220 r 26° 18.5 79°57" Between San Felix and San Ambrosio
400 s 26° 18.5 79° 49.3. | 5 km NE. San Ambrosio
250 s 26° 19 79° 52 3 km N. San Ambrosio
150 s 26° 19 7Q° 51, 3 km N. San Ambrosio
215 Ss 26° 19 Oia 49 .2 4.5 km NE. San Ambrosio |
182 = 26° 25. 79° 56.5 | 6km S. San Ambrosio
148 = 26° 26° 79° 52) 8 km S. San Ambrosio
105 Ss 26° 30° 79° 49 15.5 km S. San Ambrosio
400 r 26° 36.5 | 80° 8'.5 | 33 kmS. San Felix
550 r 27 AGiaae || Boo TA" c. 165 km S. San Felix
675 r 20 Eea. ciel *GOlc LG). c. 200 km S. San Felix
660 r 28° 33.5 80° 11 c. 260 km S. San Felix
660 r 29° 14 Soo a5" c. 340 km S. San Felix
1300 r 30° 49° 80° 24'.3
1430 r ayn ey 80° 10°
1800 — 32 CNA G KG 80° 18’ Between Masatierra and Masafuera, about 1° N.
1800 o Balan 80° 36° About too km SSE. Masafuera

Juan Fernandez situated on a “lobe” with somewhat shallower water, less than
3000 m deep, extending NW. from the coast and suggesting a possible former
connection, but recent charts are less unambiguous. The 2000 m curve makes,
however, a bulge around lat. 38°, where a depth of only 1238 m is indicated
while deeper water, 1400-1500 m, is met with near the coast. Soundings be-
tween the islands and the line where the trough stops are too few to be of much
value, but in all probability the connection, if it did exist, should be looked
for farther south. This is, as we shall see, also the opinion of the geologists.
They do not hesitate to regard the banks just described as an extension from
the continent. Nobody will, I suppose, argue that every marked rise of the ocean
floor is a sign of sunken land; the majority of oceanographers prefer to call
them independent products of volcanic action. Some, perhaps most of them,
never reached the surface, some have done so, but were broken down, but many
are still above the water and form the Pacific islands. But even the advocates
of the permanence of this largest of basins admit that its margins are zones of
considerable disturbance. If it can be proved, or at least be made probable, that

382 C. SKOTTSBERG

the deep trench following the trend of the Andes, is a consequence of the gigan-
tic mountain-building processes, the submarine ridges west of the trench can-
not help to get implicated. This idea is by no means new, it has been expressed
by many: “the width of South America may well be a good deal less now than
before the Andes were uplifted’, as GOOD says (z0g. 349), but some of these
writers did not try to penetrate the complicated geographical-geological history
of this region of great tectonic disturbance. This is true of myself, when I tried
to describe what I imagined having occurred (227. 43), but it does not apply to
IRMSCHER (743) who took pains to inform himself of the history of the Andes
as told by geologists. That they arose in a geosyncline is proved by the Jurassic
and Cretaceous beds now elevated thousands of meters and covering the older
eruptives. To the east of the depression land had existed since the Permian, to
the west was a Pacific land mass of hypothetical width; one opinion regarded
the Coast Range as belonging to this land. IRMSCHER, who took WEGENER’s
side, did not ask for any large-scale subsidence correlated with the uplift of the
Andes, because the resistance of the sialic crust to the westward drift of South
America was sufficient to account for uplift and folding. Consequently, he was
unwilling to accept PENCK’s intrusion theory; if, in the future, it should appear
essential to accept a land mass, it could be nothing more than a narrow strip
which, perhaps, had been connected with California (p. 45).

PeNcK stellte die Ausbildung der ozeanischen Tiefen am Rande des siidamerika-
nischen Kontinentes, also das Versinken angrenzender Teile des Pazifiks, der aufwarts
bewegten andinen Scholle gegeniiber und schliesst, dass die niachstliegende Erklarung
fiir die Volumeninderungen unter der festen Kruste in dort stattfindenden Massenver-
schiebungen zu suchen ist. Wenn Massen aus der pazifischen Region in die andine
iibertreten, so muss in ersterer die Kruste nachsinken (p. 51).

The cause of the uplift and folding was, according to PENCK, a result of
the intrusion of the andesitic magma, which lifted the mountains but, IRMSCHER
remarks, this could not be the only source of the tangential pressure:

Die mechanischen Ursachen des Faltenvorganges sind zweifellos anderer Natur, und
die Magmaintrusion ist genau so eine Wirkung derselben wie die Faltung. Denn es steht
fest, dass die Gebirgsbildung mit dem fenecdaneen des Magmas synchron ist und die
Intrusion somit gleichzeitig mit der angenommenen allméahlichen Lostrennung Siidamerikas
von Afrika.

Where does the Chaigneau-Merriam bank come in? WEGENER left it unexplained,
according to Du Toir it was an “advance fold”.

BERRY and SINGEWALD (29), in their review of the tectonic history of South
America, describe the development in the following terms.

Some students regard the Cordillera de la Costa in Chile as remnants of an an-
cient massif, the bulk of which has been downfaulted beneath the waters of the Pa-
cific. It consists of crystalline rocks both igneous and metamorphic and these have
commonly been assumed to be of great age—even Archean. This inference of antiquity
rests, not upon their known relations, but upon the geosynclinal nature of the Andean
seas which are clearly epicontinental and not shelf seas.

DERIVATION OF THE FLORA AND FAUNA 383

The sediments in this geosyncline are compressed between the Brazilian
massif and another massif in the west, the vestiges of which should be looked
for in the coast range. Anyhow, ‘“‘that there was land to the west of the Western
Andes cannot be doubted’. These authors date the Concepcion-Arauco series
to the older Miocene and they do not regard the flora as a coast flora.

Let us now turn to BRUGGEN, author of a modern handbook on the geology
of Chile (33z). To begin with I shall allow myself to quote FLORIN’s summary
(95.4-6) of BRUGGEN’s earlier writings.

Until Middle Tertiary times the great Andes and the Coastal Range were... a
continuous upfolded mountain chain subjected to powerful denudation. ... The principal
uplift of the Andes in Chile occurred in the Middle Cretaceous, and altered the
palaeogeographical features of the Andean region considerably. The old geosynclinal had
been turned into a continental area, at the western verge of which the border of
the Pacific Ocean at the end of the Cretaceous occupied approximately the same line
as to-day. Marine deposits of Danian... as well as Palaeocene age are lacking, and
at the beginning of the Tertiary period the continent probably extended further to
the west.

In the Eocene and Oligocene, respectively, subsidences took place and the ocean
encroached more and more on the land. The Concepcién-Arauco coal measures
are coastal deposits, which have been called the Concepcion Series by BRUGGEN.
This series, about 400 m. thick, rests unconformably on marine strata of Upper
Cretaceous (Senonian) age. The shales containing fossil plants occur in conjunction
with intercalated coal seems.... The base of this section is of marine origin, and
in addition marine layers are intercalated here and there in its middle part, which
is otherwise generally built up of freshwater deposits. According to BRUGGEN the
Concepcion Series is overlain by the deposits of the marine Navidad Series, which
is upper Oligocene or Lower Miocene in age. BrRUGGEN came to the conclusion that
the Concepcién Series belongs to the Eocene, basing this on stratigraphical as well
as on zoo-palaeontological evidence. The sediments of this series were according to
him deposited on a broad, slowly sinking coastal plain, and subsequently subjected
to considerable tilting and faulting, probably in the Miocene.

BERRY regarded both the Concepcién and the Navidad series as belonging to
the Lower Miocene or possibly Upper Oligocene, but according to BRUGGEN
this dating holds good for the latter only, while the former is much older, and
whereas BRUGGEN thinks that the Eocene coal flora was deposited in extensive
coastal swamps, BERRY regarded it as neither limnic nor littoral, but inhabiting
a lowland area away from the coast. FLORIN found that the plant remains were
laid down in the vicinity of the sea and that they are too well preserved to
have been transported any great distance.

If it is true that the coast-line, at the end of the Cretaceous, occupied the
same position as to-day, one is inclined to believe that the palaeogeography
was different before the great uplift occurred in the Middle Cretaceous, when a
large scale subsidence ought to have taken place. The oscillations along the
coast of central Chile during later times could hardly have involved the area
where the submarine ridges are found, so that the possibility to link them to
that part of the continent as late as that is small. Consequently attention has
been directed farther south, as already suggested. West Patagonia is a region
of considerable and late subsidence; the longitudinal valley of central Chile

384 C. SKOTTSBERG

disappeared under water to form the long series of the Patagonian channels, the
Andean valleys became fiords and the broad dissected fringe of islands and
skerries also give evidence to what has happened. The weight of the inland ice
during the periods of glaciation must, however, also be taken into account.

The development of the coastal region, as told by BRUGGEN 1950, is ex-
plained by facts which, if they have been correctly interpreted, open wide per-
spectives to the biologist, even if serious difficulties still have to be overcome.

BRUGGEN begins by stating that ‘“‘el mar del Eoceno’ ended somewhere
in the latitude of Arauco (38°), because a continental mass, “la Tierra de Juan
Fernandez’’ still existed (p. 50), and pp. 56-59 he relates the history of this land.
North of Rio Maullin (about 42°) is a zone of dislocations foreign to the struc-
ture of the Andes, and this zone coincides with the direction of a broad sub-
marine ridge which branches off from the continent; on this ridge are situated
the Juan Fernandez Islands and, farther north, San Ambrosio and San Felix.
Taking the 2000 m curve as a boundary, the ridge extends south to the Magellan
Straits; we observe e.g. in the island Diego de Almagro the same northwest
direction that we find in the Tertiary deposits of Parga and other places in the
zone north of Rio Maullin. To this must be remarked that the 2000 m line sur-
rounds the Chaigneau ridge and that in order to unite it with West Patagonia
the 3000 m curve has to be used. This is also seen from BRUGGEN’s map, prob-
ably copied from SupAN. The absence of marine sediments of Eocene age
shows (p. 59) that the Juan Fernandez land was, at that time, united with the
continent, but that, during the Oligocene, subsidence set in is evident from the
extension of the marine Navidad series south to 45°, and this was, as we have
heard, referred to Upper Oligocene or Lower Miocene. Also after the separation
the Juan Fernandez land continued to exist until finally, presumably with the late
Tertiary uplift of the Andes, the last rest disappeared, but not before considerable
magma ejections had given birth to the two archipelagoes. To judge from the
degree of denudation and in view of the recent volcanic activity close to Masa-
tierra and on San Felix! the islands are young, probably Pliocene,

cuando existia todavia un resto de la antigua Tierra de Juan Fernandez, de /a cual
inmigré la flora del Eoceno.2 Cuando mas tarde se hundié tambien este resto, sobre-
salian solamente las partes volcdnicas, constituyendo las islas actuales de Juan Fernandez,
que servian para refugio de la flora.

This is the process as I have described it (227. 43) and BRUGGEN is of the
same opinion from the geologist’s viewpoint. And if we go back to HOOKER’s

lecture, we shall find that the same idea, applied to a different region, was famil-
iar with him.

1 This refers to the submarine eruption in 1835 near the coast of Masatierra; whether
any signs are left I cannot tell, for the place has not been sounded. This eruption was simulta-
neous with an earthquake in Concepcidén. Possibly there was a connection with the tsunami of
Vallenar in 1922 and the eruption at San Felix three months later, when gas was ejected on
the island, killing a great number of sea-birds.

2 My italics.

DERIVATION OF THE FLORA AND FAUNA 385

With regard to the objection that oceanic islands are volcanic, and hence prob-
ably not the mountain-tops of sunk continents and that they contain no fossil mam-
mals, we have in the Malay Archipelago, vast areas of land which if submerged (and
they are exposed to constant subsidences and risings) would leave only isolated vol-
canic peaks, such as oceanic islands present. Were such an area to be submerged,
leaving exposed the volcanic peaks of Java and the Moluccas Xc, &c, should we expect
to find either recent or fossil terrestrial mammals upon them? Nor should it be over-
looked that, as a general rule, islands diminish in size and numbers toward the
centres of the great oceans, which, taken with the admission, that the great islands
adjacent to the continents were previously united to them, would favour the hypo-
thesis that all may have been so. And finally, we have instances of continental distri-
bution, presenting facts so analogous to oceanic, and hitherto so utterly unexplicable,
on any hypothesis of migration that does not embrace immense geological changes,
that we can scarcely avoid coupling the phenomena they present with those of oceanic
islands (p. 11).

However, HOOKER did not stop here. When he threw the pros and cons
into the balance, the scale of scruples sank:

On the other hand, to my mind, the great objection to the continental extension
hypothesis is, that it may be said to account for everything, but to explain nothing;
it proves too much, whilst the hypothesis of trans-oceanic migration, though it leaves
a multitude of facts unexplained, offers a rational solution of many of the most puzz-
ling phenomena that oceanic islands present: phenomena which, under the hypothesis
of intermediate continents, are barren facts, literally of no scientific interest—are
curiosities of science, no doubt, but are not scientific curiosities.

This was, I am afraid, to say too much—but are not HOOKER’s words a
good expression of the biogeographer’s dilemma?

The latest brief outline of the history of the Andes is found in GOODSPEED’s
monograph of Vcotéana (rr2); it was, partly at least, based on WEEKS’ paper
(279). The sequence of events especially refers to the central Andes, with which
the history of the southern Andes is said to agree. Four major periods of uplift
are recognized, the final one during the Pliocene—Pleistocene, with an average
elevation of 3500 m. Coincident with the uplift was a compensory coastal sink-
ing. The troughing along the coast ceased with the foundering of the Pacific
fore-lands at the close of the Miocene and early Pliocene. These lands are sup-
posed to have extended an unknown distance westward (p. 31). It is possible
that the border-lands extended all the way north along Central America, Mexico
and California, where are found islands in a position corresponding to that of
the Galapagos and Juan Fernandez Islands.

From what has been said above I can only find that there is good geo-
logical evidence for a former extension west of the continent, uniting southern
Chile with a “Juan Fernandez land’’. Before attempting to trace the sequence
of immigration of the various floristic elements and of their fate we ought to
know something of the fossil floras of Chile and, further, to devote a chapter
to the supposed Antarctic migration routes.

25 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

380 C. SKOTTSBERG
Chapter XI.

The Tertiary floras of Chile and Patagonia.

Tertiary plant fossils have been discovered in the coast region of south Chile,
in Patagonia along the Andes south to the Magellan Straits, and in Tierra del
Fuego. The richest localities are on the coast of the province of Arauco, 39 3 7ae.
s. lat.. in combination with the coal seams of Lota, Lebu etc., and east of Lake
Nahuelhuapi on the Argentine side.

The Concepcton-Arauco series.—The fossil flora was examined by ENGELHARDT,
who described more than one hundred species of angiosperms, and later by BERRY
(27). The determinations are, just as in all the other cases, based on leaf impres-
sions, but although BERRY found that ENGELHARDT’s determinations “are usually
to be relied upon” (p. 75), very many of them are, as well as his own, open to
doubt. The species were as a rule referred to living genera, belonging to many
more or less important tropical-subtropical families, Annonaceae, Apocynaceae,
Bignoniaceae, Bombacaceae, Caesalpiniaceae, Combretaceae, Erythroxylaceae, Lau-
raceae, Lecythidaceae, Myrtaceae, Palmae, Sapindaceae, Styracaceae, Vochysiaceae,
and so forth. In general character the flora-approaches that of the Amazon basin,
extended, the relief of the Andes being low at that time, to the west coast of
the continent and reaching south at least to 40° s. lat. The flora contains “no
elements of the flora of Central Chile’ (p. 106)—Cassza is, however, one of the
genera mentioned, another is Myrceugenia, represented by several species in cen-
tral and south Chile (to know Myrtaceae without flower and fruit is well-nigh
impossible). BERRY referred the flora to the Lower Miocene, “‘it is surely not so
old as Eocene’, and also younger than the Vothofagus flora in the Magellanian
zone (p. 115); this he considered to be of Lower Oligocene age.

There are two gymnosperm genera in BERRY’s list which seem to disturb
the impression of an otherwise homogeneous neotropical assemblage, Araucaria
and Seguoia. The material was reexamined by FLORIN (95) who showed that
Araucavia araucoensts Berry is a species of Podocarpus, and Seguota chilensis
Engelhardt p.p. another. These, together with a fern described by HALLE (Lygo-
dium, 312) are important additions to the Arauco flora. FLORIN, in accordance
with BRUGGEN, refers it to the Eocene and characterizes it as follows.

The composition of the fossil conifer vegetation, and the distributional aspects, of
its constituents, indicate that it derives from a warm-temperate or subtropical rain-forest,
more particularly a lowland podocarp-evergreen dicotylous broad-leaved tree forest,
growing on the coastal plain or perhaps partly on low hills not far from the coast. The
climate was probably characterized by great humidity and rather uniform temperature.
It was frostless, and warmer than the present climate of the same district (p. 26).

The possibility that plant material from the uplands had been carried down
and become mixed with material from the coastal plain is contradicted by the
state of preservation which is the same in all cases (p. 26).

The Pichileufu flora.—The fossiliferous beds of Rio Pichileufi are situated
in 41° s. lat. about 30 miles east of Lake Nahuelhuapi in a treeless steppe country.

DERIVATION OF THE FLORA AND FAUNA 387

BERRY (28) distinguished more than 130 species, most of them referred to still
living genera and belonging to 48 families and 21 orders. The general character
is subtropical, and the following families may be mentioned: Anacardiaceae, An-
nonaceae, Apocynaceae, Asclepiadaceae, Bignoniaceae, Burseraceae, Caesalpinia-
ceae, Celastraceae, Cochlospermaceae, Erythroxylaceae, Euphorbiaceae, Flacourt-
iaceae, Icacinaceae, Lauraceae, Loganiaceae, Meliaceae, Mimosaceae, Monimiaceae,
Moraceae, Myristicaceae, Myrtaceae, Nyctaginiaceae, Rubiaceae, Rutaceae, Sapinda-
ceae, Sapotaceae, Sterculiaeae, Styracaceae, Symplocaceae, Vitaceae. Of conifers
we find Avaucaria pichileufuensis, Filzroya tertiarta, Libocedrus prechilensis and 2
Podocarpus; further, there is a species of Zamza and Ginkgo patagonica, and of
ferns 3 species, one of them a Lcksonta. Araucaria and Libocedrus are, according to
FLORIN, correctly named (95), Gizkgo should be called Gixkgoites; Fitzroya belongs
to Podocarpus, and one of the Podocarpus sp. belongs to Acmopyle of PILGER.

There is no trace of /agaceae and BERRY referred the flora to Lower Mio-
cene and regarded it as contemporaneous with the Arauco flora; they have 20
species in common, the general character is the same and is said to bear witness
of the same climate. The relief of the Andes was low, no rain-shadow existed,
prevailing westerly winds carried abundant moisture across the country, there
was rain forest where now we have dry grass-land. Still, there is a difference
between Arauco and Pichileufi. BERRY (27) pointed out that the present South
Chilean rain forest flora is not represented in the Arauco flora whereas the
Pichileufi’ beds contain such Chilean genera as Asara, Berberis, Maytenus and
Myrceugenia and, in addition, the following Antarcto-tertiary genera: Dvzmys,
Embothrium, Eucryphta, Laurelta, Libocedrus and Lomatia—provided that the
determinations are correct. Nevertheless the age is supposed to be the same,
Lower Miocene according to BERRY, Eocene according to FLORIN, thus older
than the <Avaucaria-Nothofagus beds of Magallanes. It is surprising that, if the
two floras are of exactly the same age, the advancing Antarctic flora had not
found its way to the coast of Chile; Pichileufu ought to be younger, but perhaps
still Eocene, a period of very great length.

The Chalia flora.—Of considerable interest was the discovery, in Santa Cruz
Territory in the valley of Rio Chalia about 51° s. lat., of a fossil flora similar
to the Arauco and Pichileufu floras and proving that the subtropical vegetation
had extended far south. Avaucaria and Nothofagus are absent, the only conifer
found, /7tzroya tertiaria, is, as shown by FLORIN, a Podocarpus. Of angiosperm
families Anacardiaceae, Annonaceae, Bignoniaceae, Lauraceae, Monimiaceae, Myr-
taceae, Sterculiaceae etc. are represented, of Chilean genera Laurelza and Peumus
may be mentioned. The age is early Miocene according to BERRY (334), Eocene
according to FRENGUELLI (337), the climate warm temperate.

BERRY (3725) regarded all the fossil floras containing an abundance of Faga-
ceae (Vothofagus, according to DUSEN also Fagus, which is questionable) as of
approximately the same age and older than the Concepcion-Arauco series.

The Nirthuao flora.—Three localities close together on Nirihuao river near
Lake Nahuelhuapi. Some ferns, among them A/sophila australis, also known from
Seymour Island, further Zamza, Araucaria Nathorstii, Fagus (?) and one species

388 C. SKOTTSBERG

of Nothofagus. Age proposed by BERRY (334) Lowest Miocene or Upper Oligocene,
then younger than the Magallanes flora.

The Magallanes flora.—DUSEN (79) distinguished two plant-bearing horizons,
an upper Avaucaria horizon and a lower Nothofagus horizon. Of the remaining
dicotylous genera none were identified with living ones; DUSEN preferred to call
them Escalloniiphyllum, Hydrangeiphyllum etc. There is no obviously tropical
element, it is a temperate flora. DUSEN regarded the two horizons as distinctly
different in age, dating the upper to Lower Miocene, the lower to, perhaps,
Oligocene. BERRY, who doubted the correctness of this distinction, regarded them
as older than the Concepcién-Arauco flora, which seems improbable.

The Seymour flora——To judge from DUSEN’s description (So) the tropical
element is not conspicuous, whereas the actual South Chilean forest flora is well
represented: Avaucaria (nearly related to A. araucana), Drimys, Nothofagus,
Calacluvia, Laurelia, Lomatia, all supposed to be of Antarctic parentage; I can
see little reason for BERRY’s assertion that the Seymour flora contains “a large
element of subtropical or warm temperate types like those found to-day in south-
ern Brazil’ together with “another large element of forms suggestive of the
existing temperate flora of Southern Chile and Patagonia’; the former was sub-
tropical and coastal, the latter temperate ‘and montane, washed down, DUSEN
thought, from the mountains and embedded together with the leaves of the low-
land trees. The age was estimated to be Upper Eocene. FLORIN’s discovery of a
species of Acmopyle (Phyllites sp., DUSEN) is of particular interest (338).

We have seen that BERRY considered the .Vothofagus beds to be older than
the Arauco-Pichileuft’ deposits. All the local fossil floras of Patagonia are, he
says (335), older than the marine Patagonian transgression and undoubtedly pre-
Miocene. FRENGUELLI distinguished three epochs: (a) Late Cretaceous to early
Eocene, with a tropical flora, known from the Chalia beds; (b) an intermediate
period with subtropical and temperate types (Vothofagus) mixed; to this he would,
I suppose, refer the Seymour flora; (c) the youngest epoch, Miocene-Pliocene,
evidently extending into Pleistocene: a temperate flora, now ranging along both
sides of the southern Andes and characterized by the dominance of WVothofagus
and of a number of conifers. The more or less corresponding development of
the Andes was according to BERRY (28): (1) Eocene-early Miocene: low relief,
no high continuous mountains, followed by (2) a period of great uplift; (3) late
Miocene to early Pliocene: mature erosion, low relief; (4) late Pliocene to Pleis-
tocene: extensive uplift, beginning of the formation of the Chilean and Peruvian
deeps, where earlier there was land; (5) submergence of the coastal plain. Finally,
but not mentioned by BERRY, the series of glacial and interglacial periods, a
most important factor of disturbance.

This is the background against which we have to discuss the history of the
Juan Fernandez flora.

DERIVATION OF THE FLORA AND FAUNA 389

Chapter XII.

Antarctica as a source of the present circumpolar floras.

Much has been written on this subject and it is not necessary to review
the entire literature, which has been done already by several authors, but the
Antarctic problems are so important when it comes to an analysis of Juan Fernan-
dez that they cannot be passed in silence. J. D. HOOKER was the first to survey
all the lands scattered around the Antarctic, Tierra del Fuego and the Falkland
Islands, Kerguelen Island, Tasmania, New Zealand and its subantarctic dependen-
cies; he was struck by the discontinuous distribution of many genera, families
or even species—HEMSLEY, 727 (a), and the author (228) have given lists of
such genera and species—he drew the consequences, although nothing was known
then about the vanished flora of the large, ice-covered continent, nor of the palaeo-
geography of the adjacent zone, and the idea that Antarctica had formerly ex-
tended farther north and that the sporadic southern islands eventually were
remnants of larger land masses entered his mind. Since that, the various subant-
arctic and austral floras have become very well known, and the cases of remark-
able disjunctions have multiplied. To those who adhere strictly to the hypothesis
of long-distance dispersal across the oceans this means nothing more than further
proofs that they are right, for the west-wind drift explains everything. Fortunately,
the land bridge between South America and Antarctica rests on solid foundation,
geographical as well as geological. In his important paper of 1929 (737) HOLTE-
DAHL has shown that the old idea of land connection between Tierra del Fuego
and Graham land (Palmer peninsula) by way of the Burdwood Bank, Shag Rocks,
South Georgia and the South Sandwich and South Orkney Islands, which had
been doubted by some, holds good: we have to do with a mountain range, a
continuation of the South American Andes, bordered by deep water which, on
the Pacific side, has the character of an abysmal trench but which exhibits old
sediments to such an extent that we are forced to postulate land where there is
now deep sea. The South Sandwich Islands, being entirely neovolcanic, have the
appearance of an “‘oceanic’” archipelago, but they were built up during late Ter-
tiary times over an older foundation—a parallel to the history of Juan Fernandez
and of many other islands.

It is ... quite evident that the South Shetland land mass has once had several
times the width that it has to-day. ... With their large amount of terrigenous, clastic
sediments etc., the South Orkneys and also South Georgia agree with the folded ranges of
the continent. In fact, in order to explain these masses of sediments we must necessarily
assume land to have been present where there is now deep sea.

We need not assume that all the links of the Scotia or, as it is also called,
South Antillean Arc were united at the same time, let it be that this is possible
or even probable; if so, there was no communication by water between the At-
lantic and the Pacific, which undoutedly must have had its consequences to the
water circulation, a question I am quite unfit to discuss. To judge from the compo-

390 C. SKOTTSBERG

sition of the local subantarctic floras, rich in herbaceous plants belonging to many
different families and orders, land communication must have persisted to middle
or even late Tertiary times. That the connection goes far back is shown by the
occurrence of a Gondwana flora on both sides of the Drake passage (Graham
land. Falkland Islands), and the Cretaceous rocks of South Georgia prove, as
HOLTEDAHL says, the existence of a land mass of considerable size where there
is now sea. JOYCE (65) remarks that “there is good evidence that the Scotia Arc
with its extension into West Antarctica has persisted as a structural feature since
Lower Palaeozoic times’.

The Scotia passage offers one of the migration routes over land that we are
in need of, but another passage, the Macquarie route between East Antarctica
and Australia-New Zealand is required to make the trans-antarctic route complete.
There are intermediate islands, Macquarie, Auckland and Campbell Islands, with
a subantarctic flora suggesting former connections, and there are tracts with shal-
lower water between Tasmania and the continent, but for want of geological evi-
dence this bridge is hypothetical, and many authors prefer to speak of submerged
intermediate islands sufficiently close to facilitate the spread of organisms able
to cross moderate water barriers. As AXELROD says (r4. 183):

Archipelagoes of only slightly greater extent than those now present could account
for the continuity of the Antarcto-Tertiary Flora in all these regions during the early
and middle Cenozoic.

FLORIN, referring to the present and former distribution of conifers, expressed
himself in similar terms.

Antarctica has played an important role in the development and distribution of
the southern group of conifers. The data related to its distributions considered in this
paper seem most readily explained by assuming land connections, or at least much
closer proximity between Antarctica and the adjacent southern ends of South America,
Australia, New Zealand and South Africa (95. 92),

and after the discovery of a Tertiary species of Acmopyle in Patagonia and a sec-
ond fossil form in the Eocene of Seymour Island, he wrote (3378. 136):

Fiir eine ehemalige Verkniipfung der australischen Region mit der Antarktis spricht
auch die Verbreitung der Gattung Acmopyle. Dass also die Antarktis in diesem Falle
als eine alte Vermittlerin zwischen der australischen Region und Siidamerika gedient
hat, muss meines Erachtens angenommen werden.

However, one of the supposed links, Macquarie Island, does not, it seems,
possess any plants, perhaps not even mosses or lichens, dating back to the height
of the Glacial Period, for this island was, at least during maximum glaciation,
entirely ice-covered and must, TAYLOR says, have received its present plant world,
very poor it is true, in postglacial time from the north and across a considerable
stretch of open sea (263). In Antarctica proper the situation is, with regard to
lichens and mosses, different. DAHL (72. 231), basing his opinion on the discovery
of numerous endemic lichens and of a few mosses not very far from the south
pole, concludes that part of the flora survived the glaciations. Here, where high

DERIVATION OF THE FLORA AND FAUNA 391

mountain ranges are found right along the coast, the inland ice, even during
maximum glaciation, cannot have covered everything. The flora may not have
been as rich in species as it is now but perhaps a little more varied than one
has been inclined to think. Many species are also found in the subantarctic
zone, and further research work will, perhaps, reduce the number of endemic
species.

FLORIN spoke, as we have seen, of a “proximity to South Africa’. This is
where most biogeographers hesitate, in spite of such eloquent facts as the distri-
bution of Restionaceae, Proteaceae and other families, and the occurrence of a
subgenus of Gunnera on African soil. HOOKER, it is true, had a vision of a larger
Kerguelen land, but this was still a long way off from Africa. Perhaps GULICK
(77g) ought to be mentioned here; he opposed the continental nature of isolated
islands, but he was tempted to exempt what he called “continental outsiders’,
“Kerguelen, Crozet, St. Paul and two or three more’’; the sea lacked the deepness
of a typical ocean, and sediments occurred on Kerguelen; for these reasons he
admitted a possible former existence of a “northward lobe of the Antarctic con-
tinent”’. The lichenologist C. W. DODGE has taken up this question; the lichen
flora of ‘““Kerguelia’’ presents features of great antiquity as well as of prolonged
isolation (5 endemic genera), and the angiosperms include such aberrant types as
Pringlea and Lyadlia; it should be remembered that WERTH was opposed to over-
seas dispersal. Kerguelen is volcanic, but old, the oldest lavas dating from late
Mesozoic or early Tertiary times, and on them fluviatile sediments and, on top
of these, Oligocene strata rest. Erosion broke down the island during Miocene-
Pliocene, but renewed volcanic activity followed from the end of Pliocene into
Pleistocene. The Gaussberg-Kerguelen ridge connects Antarctica with Kerguelen +
Heard Island; an elevation of 400 fathoms would be sufficient to unite the two
islands, a rise of 100 fathoms would transform the Crozet group into a single
island, and DODGE supposes that there is a submarine connection between Ker-
guelen and the Crozet swell. Kerguelia in its prime would include all the islands,
also Marion and Prince Edward. The great difficulty, the extension to South
Africa, remains. So much seems to be certain that, if this bridge did exist,
separation took place early, long before the other Antarctic connections were
broken off.

I think that the majority of phytogeographers agree with MERRILL who, in
his last work (306), wrote that

there is no reason whatever to doubt the validity of this ancient Antarctic route of
migration of various families and genera of plants; certainly, no experienced phytogeo-
grapher would question the validity of this route, for it is as thoroughly established
as its more evident equivalent by what is now the Arctic region (p. 178).

Most botanists have drawn their conclusions from the present distribution
of the plants and this is, as a rule, all they can do, because few have left any
traces of their distribution in earlier epochs. Nevertheless we have no good reason
to doubt the important part taken by Antarctica in the history of the south
hemisphere, but the proofs that such was the case, BERRY emphasizes (28. 34),

392 C. SKOTTSBERG

must rest on palaeobotanical evidence, and it happens that the fossil records are
at variance with current ideas. The occurrence of Araucaria, Drimys, Laurelia,
Nothofagus etc. in Tertiary deposits on Seymour Island would lead us to infer
that they are of Antarctic origin and have radiated from there, but Avaucarza
once had a world-wide distribution, Drzmys belongs to an order—Magnoliales—
of Holarctic range, the same is true of Fagaceae, and even Lawrelia is, BERRY
points out, open to doubt, in spite of the fact that the Monimiaceae are a south-
ern family. Many other genera, called Antarctic on the strength of their modern
distribution, are known as fossils in the north temperate and Arctic zones. “ Arau-
cavia stands as a perpetual warning against forgetting that the past is the key
to the present’, BERRY wrote (I.c. 36). A Holarctic genus may have reached New
Zealand or Australia as well as Patagonia from the north, never having used an
Antarctic route, and without leaving a trace of its wanderings. On the other hand,
the little we know about the preglacial vegetation of Antarctica is sufficient to
prove that this large land mass, just as every other part of the globe, was inhab-
ited by a rich and varied flora, that it may have been a primary centre of
evolution, that, in other instances, it served as a secondary centre and that it was
a much-trodden road between America and Australia-New Zealand.

Miss GIBBS appears to have been one of the very few experienced phyto-
geographers who refused to regard Antarctica either as a centre or as a migra-
tion route over land; it had always been surrounded on all sides by water and
no other agents than “the wild west wind” (z06. 103) and a pole-ward north-west
wind, coming from Asia, were needed to explain every distribution pattern. The
southern focus of development was not Antarctica but the mountains of New
Guinea. The highland of Tasmania, the subject of her survey, had received
nothing from the south, all the so-called Antarctic plants, genera lika Adrodanella,
Astelia, Carpha, Colobanthus, Coprosma, Drimys, Gaimardia, Gunnera, Lageno-
phora, Nothofagus, Oreobolus and so on, had come from New Guinea, and from
there they had radiated to Polynesia, Hawaii, Juan Fernandez, Tierra del Fuego
and, I presume, Antarctica. Had she lived to hear of the discovery of Vothofagus
in New Guinea, where more species have been found than anywhere else, and
in New Caledonia, and of the rich development of the Winteraceae in New Guinea,
she would have regarded such finds as a forcible proof of the correctness of her
opinion.

Many zoogeographers have looked with much suspicion at the Antarctic
continent as a centre of radiation. SIMPSON, in his review of the theories involving
Antarctica in the distribution of vertebrates (223), concluded that dispersal had
been, in all cases, from north to south; not even the Scotia Arc had ever been
used as a route of migration. His reasoning is logical and often conclusive. The
invertebrates are, however, left aside. To quote part of his summary (p. 767):

There is no known biotic fact that demands an Antarctic land-migration route for
its explanation and there is none that it more simply explained by that hypothesis
than by any other. The affinities of the southern faunas as a whole are what would
be expected from the present northern connection known, or with considerably prob-
ability inferred, to have existed at appropriate times in the past. There are certain

DERIVATION OF THE FLORA AND FAUNA 393

troublesome anomalies and exceptions in the evidence, but none of these can be ade-
quately explained by postulating an Antarctic connection. The general weight of evi-
dence is against such a connection.

In scientific theory the best-supported and most nearly self-sufficient hypothesis
should be preferred and unnecessary additional hypotheses should be rejected or held
in abeyance. On this basis the Antarctic migration route hypothesis remains simply a
hypothesis with no proper place in scientific thinking.

To this I shall make a few remarks. If SIMPSON had said “no fact involving
the vertebrates’ instead of “no biotic fact’—very well, let us assume that the
routes across from and to either America or New Zealand were impassable to
mammals, reptiles, amphibia and flightless land-birds, birds with good flight capa-
city would have found little difficulty to cross, and many biotic facts are known
that clearly speak in favour of an Antarctic migration route for invertebrates and
plants. SIMPSON must have thought that either did the Antarctic continent never
possess a fauna of land vertebrates, or, if it did have one, it had evolved inde-
pendently of all other faunas and disappeared without leaving a single trace.
Only penguins are known in a fossil state in Antarctic Tertiary deposits. Until
fossil land vertebrates are discovered, the question of the former existence of an
Antarctic fauna of terrestrial vertebrates must be left open.

Among the invertebrates are many examples of a discontinuous distribution
most readily understood if Antarctica is taken into account. Several were men-
tioned in the chapter devoted to the composition of the fauna of Juan Fernandez,
a few more may be quoted here. BERLAND, dealing with the Pacific spider fauna (23):

Nous avons tiré de notre étude cette notion importante que la liaison entre |’ Aus-
tralie et l’Amérique a eu leu par une terre antarctique dont les témoins restent ac-
tuellement, mais ni par la Nouvelle Calédonie, ni par la Nouvelle Zélande, ni par le
groupe Samoa—Tonga-Fiji, et, par voie de conséquence, qu'elle n’a pas eu lieu par le
centre du Pacifique (p. 1053).

BERLIOZ (24), with examples offered by the distribution and affinities of beetles,
states that the group of Buprestidae, forming “le noyau essentiel’’ in the Bupres-
tid fauna of Australia, has mainly South American affinities (several genera),
and that

la faune des Lucanides d’Australie et de Papouasie présente avec celles de l’Amérique
du Sud surtout de la région andine et patagonienne, des affinités aussi étroites que
curieuses.

Finally LINpsAy (767), calling attention to the Subantarctic Collembola,
extremely delicate creatures “supposed not to be carried any appreciable distance
either by wind or sea, thus being important proofs of former land connections’,
mentions a genus of 3 species of which one is Fuegian, one recorded from

the Scotia Arc, and one found in New Zealand. And other similar examples may
be found.

ERG
394 Cc. SKOTTSB

Chapter XIII.
The history of Juan Fernandez—a tentative sketch.

In my first paper on the Botany of Juan Fernandez (227), where only the
vascular plants, as known at that time, were included, I expressed my view on

the history of the flora in the following words:

Freilich haben wir keine Ahnung davon, wie schnell Arten oder Gattungen ent-
stehen, aber wir k6nnen uns kaum denken, dass in der kurzen Zeit, die seit der Ent
stehung der jetzigen Inseln verflossen ist, sich Typen wie Lactoris oder Robinsonia aus
“Keimen’”’ entwickelten; die nach den Inseln gebracht wurden, um sich in ungest6rter
Isolierung umzuformen ... Ich bin der Meinung, dass das alte Element nicht auf Masa-
tierra oder Masafuera entstand, sondern Alter ist als die jetzigen Inseln, und dass es
wenig wahrscheinlich ist, dass die alten, endemischen (oder andere, eng verwandte)
Gattungen und Arten von Juan Fernandez, noch nachdem die Inseln gebildet waren,
die vielen vermeintlichen Ursprungsorte bewohnten, und dass Verdnderungen in der
Pflanzenwelt von Polynesien, Neuseeland, Chile u.s.w. in quartérer Zeit die isolierte Stel-
lung bewirkt haben. Ich glaube also, dass in vor- und friihtertidrer Zeit gréssere Ent-
wickelungszentra existierten, und dass ihre Flora nunmehr als ein altpazifischer Rest
fortlebt. Auch eine Restflora wird sich aus vielen Familien und Gattungen aber ver-
haltnismadssig wenigen Arten zusammensetzen.

At that time our knowledge of the flora was incomplete; much fresh and new
material was added in 1916-17. I had fixed my attention on what I called the
“Old Pacific element’, but did not venture to look for an exact site of an evolu-
tion centre, though the possibility of Antarctica as an important source of genera
and families had been pointed out before; the first sign of the presence of there-
tofore unknown, subantarctic flora in Masafuera had been observed, but I had no
reason to link it with the Old Pacific types. During the 1916-17 survey a rather
strong Antarctic-bicentric group took shape, and in my 1925 sketch of the history
of the flora (23z) not only was this, but also the Old Pacific plants claimed to
have “reached Juan Fernandez over South America, where they have disappeared”
(p. 31). My object this time is to see if we can approach these problems in other
than general terms.

Our starting point is the “Tierra de Juan Fernandez’ of BRUGGEN, forming
a westward extension or lobe of South America, reaching the actual site of Juan
Fernandez and Desventuradas Islands, as indicated by the bathymetrical conditions,
something in keeping with the peninsula of Lower California and separated from
the coast by a broad, toward the south gradually narrowed bay. However, if we
remember that the deep trough is supposed to have originated with the final
uplift of the Andes, the present coast line ought to be recent all along, but this
is not in accordance with the opinion that the Eocene Arauco flora was a coast
flora. The conclusion would be that the great depths were initiated already during
Cretaceous times, getting deeper and deeper with the successive periods of uplift.
The Juan Fernandez land or peninsula formed part of the neotropical Eocene flora
region which extended from Venezuela and Brazil to south Chile and east across
the mountains, during this era of low relief, as shown by the fossiliferous beds

DERIVATION OF THE FLORA AND FAUNA 395

of Pichileufu. As the Concepcién-Arauco flora it was a subtropical rain forest
flora with podocarps, tree ferns, evergreen dicotyledonous trees and lianas, be-
longing to some 30 tropical families; the species were, with two exceptions, referred
by BERRY to still existing genera—if the determinations are reliable. This is
perhaps more than we can expect; all we can say is that BERRY was a man with
a wide experience of both fossil and living tropical plants and that undoubtedly
many of the families listed and perhaps also a fair number of the genera are
correctly placed. Two are found in the present flora of Juan Fernandez. To Azara
celastriniformis and tertzarza—and the fossil does suggest Azara—BERRY remarks
(28. 107):

As a recent form occurs on the island of Juan Fernandez one can predicate a con-
siderable antiquity for the genus, which is more than verified by the present fossil forms.

His Berberis corymbosifiora is of still greater interest:

I have seen leaves of all the South American species and the most similaris Berberis
corymbosa Hook. et Arn. of Juan Fernandez (l.c. 75).

I have compared his illustrations with the island species and I am willing to
testify to the striking similarity between them. A revision of these most important
fossil floras, with application of modern technique, is eagerly longed for.

Anyhow, the neotropical character of the old flora has been safely estab-
lished, and Arauco and Pichileuft' agree in their general composition; 20 species
occur in both. If this flora extended to Juan Fernandez, this land must have been
sufficiently high to force the prevailing westerly winds to unload part of their
moisture and to give rise to altitudinal belts. The flora most likely had its special
distinctive marks. In view of the very large area it inhabits, it cannot have been
uniform, and different floristic provinces showed special features and had their
own endemics. If anything still survives in identical or very similar form can only
be a subject of conjecture and is not demonstrated by leaf impressions. Two of
the genera reported from the mainland, Azara and Myrceugenza, still occur in
Chile and Juan Fernandez, and I am inclined to believe that the endemic element
in the insular forest flora dates back to early Tertiary times, genera like Podo-
phorus, Megalachne, Fuania, Ochagavia, Nothomyrcia and Selkirkia, and species
of Chusquea, Hesperogreigia, Urtica, Phrygilanthus, Chenopodium, Colletia, Dys-
opsis, Ugni, Eryngium, Rhaphithamnus, Solanum and Nicotiana.

Centaurodendron, Yunquea and the four endemic Cichoriaceous genera stand
apart. They are montane and we have no clue at all to their history, but we can
take it for granted that they are not “new beginners’, but old relicts, without
any near relatives anywhere. Whether we assume that they arose in the islands
and have left no marks in the continent, or derive the four dendroseroid genera,
which form a natural group, from Antarctic ancestors, a possibility certainly not
offered by Cextaurodendron and Yunguea, we are victims of wild speculation.

Few endemic ferns belong to the neotropical element, 7richomanes Ingae,
Dryopteris inaequalifolia, Asplenium macrosorum and stellaitum, Pellaea chilensis,

396 Cc. SKOTTSBERG

Polypodium intermedium and Ophioglossum fernandezianum, but Pellaca and Ophio-
glossum do not grow in the forest, the former inhabiting the dry coast cliffs, the
latter the open grass land, and they may have their own history.

How and when the non-endemic South American species reached Juan Fer-
nandez is hard to tell. They are temperate and, with the exception of Wyrteola
numimulavia, herbaceous. Myrteola is the only member of this group that extends
south to the subantarctic zone. Of the others Danthonia, Koeleria, Stipa, Piptochae-
tium, Eleocharis, Funcus procerus (also dombeyanus and imbricatus?), Libertia,

eperomia fernandesziana, Partetaria and Mimulus show, in their mode of occur-

rence, every sign of being indigenous. Advocates of transoceanic dispersal would
not hesitate to call them “‘late arrivals which have not had time to change’, and
a direct transport is not altogether impossible. It would be interesting to know
if a grass land existed when Great Juan Fernandez was connected with the mainland,
but unfortunately we do not even know the extension of the S¢t7pe/um when the
early voyagers reported on the vegetation and already found the introduced Avena
barbata in dominance in the treeless western part of Masatierra. A remark made
by BRUGGEN deserves to be quoted in this connection. A current coming from
the south swept past the shore of Great Juan Fernandez. When, during the Na-
vidad transgression, separation from Chile occurred,

.el mar del polo sur entro en comunicaci6n con el mar que bafiaba las costas de Chile
Central y el primer antecesor de la corriente de Humboldt llev6é las aguas mas frescas hacia
el norte, dando principio a la gran zona desértica.

BRUGGEN seems to have forgotten that at that time Antarctica was covered,
not by an inland ice, but by luxuriant vegetation, and that the sea cannot have
been cold; however, there must have been an uppwelling of cold water, and the
dry climate of the basal belt may have been as unfavourable for tree growth as
it is now. Nevertheless I cannot believe that the present steppe-like communities
date back to early or even middle Tertiary time. On the other hand it seems
quite unlikely that the species of Stipa, Piptochaetium, Danthonia, etc. were in-
troduced with the traffic while, in this respect, Chaetotrop7s (the endemic nature
of Ch. mberbis questionable), the two Cyperus, Funcus capillaceus, Paronychia,
Centella, Hedyotis and Plantago truncata are under suspicion.

Of the ferns found elsewhere Polypodium lanceolatum is pantropical and old
enough to have belonged to the ancient flora, and this may be true also of
Lrichomanes exsectum, Hlymenophyllum spp., Adiantum chilense, Pteris chilensis and
semiadnata, and Elaphoglossum; | have suggested that the latter was carried
directly to Masatierra by a northerly storm, and this could have been the case
also with Polypodium Masafuerae, observed a single time 100 years ago and never
again. P. trichomanoides remains doubtful in spite of the specimens still extant
and labelled Juan Fernandez (279. 766).

The presence of a large, presumably boreal element is not difficult to explain,
for it extends all along the Andes to the far south. Many of the species are
endemic, Agrostis masafuerana, 2 sp. of Spergularia, 2 Berberis, Cardamine Krues-

DERIVATION OF THE FLORA AND FAUNA 397

selit, Galium masafueranum and 6 Lyrigeron. The Boreal character of these genera
is recognized. This element extended south along the precursors of the late Tertiary
mountains; the species of Agrostis (?), Spergularia, Berberis, Cardamine and Ga-
‘ium have their closest relatives either in the tropical Andes (erber7s) or in Chile.
Evigeron deserves special attention on account of the large number of Andean
species nearly related to each other but less so to the insular group, which shows
a remarkable differentiation: /. fruticosus and its cognate /uteoviridis, the three
herbaceous rosette herbs, and the peculiar /. 7wpzco/a of the coast rocks; of these
fruticosus is found on both islands and the other species endemic on Masafuera.

Nine non-endemic species, all found in Chile, also belong here, 777setum, Carex
Banksi, Paronychia, Cardamine flaccida, Callitriche, Rubus, Empetrum, Calystegta
and Guaphalium, and, among the ferns, perhaps Cys/oplerzs. Rubus geoides forms
together with A. radzcans an isolated section but has a more southerly distribution,
but all may have reached Juan Fernandez from South Chile.

Finally, Czmznza remains to be accounted for. Whether we link it with the
palaeotropical Prasioideae, which seems to be the best way, or with Bystropogon,
it stands out as an isolated relict genus.

I have distinguished a large Antarcto-tertiary element, over 40% of the
angiosperms and 60% of the ferns. Among the former there are three or, if Lac-
toris is kept aside, two groups; one of them (1) is still represented in South
America, the other (2) not. In (1) two types can be distinguished, (a) not confined
to subantarctic or alpine habitats and demanding a milder climate; to this lot I
refer Uncinta Douglasi and costata, Drimys, Phrygilanthus, Escallonia, Margyrt-
carpus, Sophora, Gunnera, Apium, Pernettya, and possibly Plantago fernandezia.
All have relatives in Chile. To these are added the species also occurring on the
mainland: Danthonia, Koeleria, Funcus, Libertia, Acaena ovalifolia (indigenous),
Centella and Nertera; the two grasses are, however, only tentatively referred to
this element.

The occurrence of an Antarctic element in the Eocene flora of the mainland
has been demonstrated. BERRY lists Avaucaria, Libocedrus, Drimys, Embothrium,
Laurelia and Eucryphia from the Pichileufu beds; even if the ‘magnolia stock’’
is of Boreal origin this does not exclude the possibility that the Winteraceae
radiated from Antarctica, a parallel case to Fagaceae and Nothofagus.

If we follow COPELAND many of the ferns also belong to 1 a: Hymenoglossum,
five species of Hymenophyllum, Lophosoria, Polystichum, Blechnum, Hypolepzis,
Flistiopterits, Gleichenta pedalis (indigenous?) and Lycopodium scariosum.

Group 1 b includes the so-called Subantarctic-Magellanian element inhabiting
Fuegia, the Falkland Islands, etc. ranging north along the Andes and belonging
to a well-known circumpolar assemblage of genera and species. Here we find
Oreobolus, Uncinia brevicaulis, phleoides and tenuis, and Lagenophora hirsuta,
further three endemic species, Acaena masafuerana, Abrotanella crassipes, both
with near relatives in West Patagonia—Fuegia, finally Agrosts masafuerana, if its
relation to the bicentric A. magellanica is confirmed after monographic treat-
ment. I brought it to the boreal group. The following pteridophytes are attached

398 C. SKOTTSBERG

here: Serpyllopsis, Hymenophyllum falklandicum, Polypodium (Grammitis) magel-
lanicum, Gleichenta quadripartita, and Lycopodium magellanicum.

Did this Antarctic element, mostly not endemic, extend to Great Juan Fer-
nandez or did it arrive after the separation from the mainland took place, per-
haps even after that the volcanic islands had been formed? The same question
was raised when we discussed the non-endemic neotropical-temperate Stipa, Pipto-
chaetium, Myrteola, Rubus, and so forth; the Boreal group is, as we have seen,
also involved. Is it probable that also this flora dates back to early or middle
Tertiary times? This seems unlikely. The subantarctic species are, with the excep-
tion of Grammits, restricted to the highland of Masafuera, but may have occurred
also on Masatierra when the islands stood higher. Either we must assume that
the land connection with the continent persisted much longer than is otherwise
probable, or those species have immigrated across the water in late Pliocene or
in postglacial time. I have suggested this on repeated occasions (237, 232, 340).
However, we must not forget that we have to do, not with stray colonists, but
with plant communities composed by flowering plants, ferns, bryophytes and
lichens. Unfortunately nobody beheld the vegetation before introduced species,
Anthoxanthum odoratum and Rumex acetosella, had invaded the highland and
changed the entire aspect beyond recognition.

To distinguish, among the bryophytes, the old element which undoubtedly
must have formed an important part of the subtropical forest flora is more than
I can undertake; we do not know if the many endemic species are relicts or not;
in fact, we do not even know if they are endemic until the opposite mainland
has been well explored. So much can be said that species with a pronounced
tropical distribution are few, but if we add the Chilean species extending through
the Valdivian and Magellanian forest zones, this South American group makes
up about 40% of the mosses and 30 % of the hepatics, endemic species of Ameri-
can affinity included. Perhaps half a dozen mosses, not counting the few that
accompany man wherever he goes, and some liverworts, have a wide distribution
outside America. The dominant element is Antarctic; this was emphasized by
such authorities as CARDOT and HERZOG: 56% of the mosses and 68 % of the
hepatics were referred to the Antarcto-tertiary element.

When it comes to distinguishing corresponding groups among the lichens we
move on very unsafe ground, but there are indications that, beside a large South
American element, we also have an Antarctic group to which no less than 58
species were referred. They are austral- or subantarctic bicentric or tricentric,
but many of them range north into lower latitudes. A conspicuous part is formed
by Stictaceae. The great bulk of the family is by no means southern, Loédaria,
Sticta and Pseudocyphellaria are frequent in tropical, subtropical and temperate-
oceanic climates throughout, but the austral-circumpolar species are so many that
we cannot exclude Antarctica as a possible source.

We shall proceed to group 2. It comprises the genera or species which are
foreign to the South American flora and have their relations in Australasia. JoHow
recognized very few; he had, just as several later authors, no other explanation
to offer than that they had arrived across the Pacific from Australia, New Zealand,

DERIVATION OF THE FLORA AND FAUNA 399

the East Indies, etc. without reaching the coast of Chile. They are much more
numerous than JOHOW thought, 28 species: Cladium, Carex berteroniana, Pepero-
mia berteroana, margaritifera and Skottsbergu, Boehmerta, Santalum, Ranunculus,
Fagara (2), Halorrhagis (3), Euphrasia, Coprosma (2), Wahlenbergia (5), Robinso-
nia (5), Symphyochaeta and Rhetinodendron, the three last genera endemic. Objec-
tions may be raised against including Cares and /uphrasia; the section to which
Carex berteroniana was referred by KUKENTHAL is almost confined to New Zealand
and barely represented in Australia, Tasmania and Norfolk Island, but one little
known Chilean species is included, and Luphrasia formosissima is distantly related
to £. perpusilla of South Chile. The species of Wahlenbergia are puzzling, but
I have given my reasons for bringing them here as representing an African sector.
The most eloquent members are, perhaps, Saxtalum, Ranunculus, Halorrhagis and
Coprosma.

This element is conspicuous also among the ferns: the extremely old 7/yr-
sopleris, Arthropteris, entirely unfamiliar with the neotropical flora, Dicksonia,
Blechnum Schottit and Pteris berteroana.

I never looked in earnest for a direct road across the south Pacific from
Australasia to Juan Fernandez, a route which ought to have had South America
as its terminus. I preferred to think that the group in question reached the islands
over the Scotia bridge and South America where, however, it had become extinct.
To prove this we must turn to palaeontological evidence. The Eocene beds on
the mainland contain leaves of many different plants, and it is not impossible
that a revision of the material will contribute to a solution of the problem. In
the lists published by BERRY two items call for attention, Cyatheordes thyrsopt-
eroides in the Arauco flora, and Coprosma from Pichileufu, but the material is
sterile. It is true that, to judge from BERRyY’s illustrations, Cyatheoides suggests
Thyrsopterts, but the author later (28.57) compared it with his Decksonza patago-
nica, which was found with sori and undoubtedly belongs to the Cyatheaceae.
Thyrsopteris-like fossils have been reported from various places in the north
hemisphere. He described 2 species of Coprosma, based on leaf impressions
which, as far as I can see, tell us little about their systematic position. To
C. spathulifolia he remarks:

These tiny leaves have occasioned a good deal of trouble, as the South American
representatives of the genus are not similar to the fossil. ... The Chilean species are
not closely similar... .

and to C. zcerta, a most appropriate name:

... they are so much like the endemic species of Coprosma of the Juan Fernandez
Islands and several forms from the Hawaiian Islands that I feel constrained so to
identify them, at least tentatively... .

I cannot find that they agree better with Cofrosma than with many other
genera. When BERRY gives the distribution of the genus as “from the Malayan
archipelago through the Pacific islands to Chile’ he includes Juan Fernandez under
Chile where, politically, the islands belong, for there are no species on the mainland.

400 C. SKOTTSBERG

It is very easy to construct a hypothetical passage by which the “Coprosma
group’ reached Juan Fernandez without crossing the Antarctic or encroaching very
much upon the surface of the -acific Ocean. If we have reason to think that South
America extended farther west I cannot see why this wasn’t the case also with West
Patagonia and Tierra del Fuego, a region which undoubtedly has undergone con-
siderable submergence; we have to count with a wide Scotia bridge and an exten-
sion of Palmer (Graham) Land, where the geographical-geological situation is the
same as in South America and where the uplift of the mighty “Antarctandes”’
ought to have been accompanied by submergence of the fore-land. Plants and
animals could have travelled by a circuitous route from the New Zealand region
over West Antarctica to Juan Fernandez without finding their way east to what
is now Chile. This south Pacific path was suggested above when I tried to divide
the angiosperms according to their supposed primary sources; the species involved
are enumerated under I: 3, forming a group ‘‘as far as known without continental
American affinities, either suggesting an ancient Antarcto-Pacific track east from
Australasia without reaching America, or having arrived along the road over the
Scotia Arc without leaving any traces in the present American flora” (p. 2609).

This idea of a South Pacific track is not new. It was postulated by ARLDT
as a South Pacific bridge and it finds an expression in CROIZAT’s South Pacific
base-line, which, however, if it existed, hardly permits us to draw such far-reaching
conclusions as he did. In the case of Juan Fernandez only 13 genera are concerned,
belonging to 11 families, the ferns not included, but they make up 20% of the
angiosperms, and others may have existed that disappeared later.

It remains to see if, among the cryptogams, a “Coprosma group’ can be
recognized. In HERZOG’s paper on the Hepaticae (730) a single species is indicated
as restricted to New Zealand and Juan Fernandez, Pallavicinia xiphoides, and two
endemic species are said to have their nearest allies not in South America but
in New Zealand. No case equal to Pallavicinza is found among the mosses, but
several endemic species are considered to be related, not to American ones, but
to species inhabiting the south-west Pacific region.

With regard to the fauza I shall confine myself to some general remarks.

The few land-birds are of neotropical origin. Of the seven species, three,
Eustephanus galeritus, the owl and the thrush, occur in identical forms on the
mainland; the remainder are either endemic varieties or endemic species. The
most divergent is Lustephanus fernandensts. GOETSCH’s idea that it originated in
the islands as a mutation of /. gaderitus is contradicted by the fact that they are
not at all closely related but even brought to different genera by some ornitho-
logists. The former is a relict, the latter perhaps a late immigrant. Of the breeding
sea-birds the genus /verodroma forms an austral-circumpolar element and may, in
preglacial time, have inhabited the coasts of Antarctica and adjacent islands.

Little can be said as yet about the invertebrates. The endemic leech, Veso-
philaemon, is an important case of non-American ancestry, and the only terrestrial
amphipod is bicentric. The spider fauna is an appendix to the fauna of South
America, but with special features; there is no endemic genus, but specific endem-

DERIVATION OF THE FLORA AND FAUNA 401

ism is high. In the fauna of the mainland Antarctic affinities have been stated
to occur (BERLAND 23. 1044):

The New Caledonia, New Zealand group has affinities with the Malaysian region
and still more with Australia. But the small islands situated south of New Zealand,
namely the Campbell, Auckland and Macquarie Islands, are different; they present,
rather abundantly, a group of spiders, Cybaeinae, relatives of which are found in the
extreme south of South America; these spiders are not present in New Zealand, but
are allied to Australian and ‘Tasmanian species.

This is an interesting observation, for it is known that connections between
East Antarctica and lands to the north have been looked for both with New Zea-
land and over Tasmania with East Australia.

Among the millipedes Aulacodesmus and Nesogeophilus are austral genera
and the species endemic; Schzzotaenza alacer is known from Chile, but the genus
is eminently austral-bicentric. For the same reason the endemic genus of Thysa-
nura merits to be noticed.

A very great number of insects have been reported from Juan Fernandez,
most of them endemic, also many of the genera. The majority has been described
only recently and very often nothing was said about their relations; where they were
stated they are, as a rule, to be found in South America. Isolated forms are
plentiful and bear witness of a long history. For the single termite an Antarctic
ancestry is postulated. Most of the butterflies collected have not yet been de-
scribed. Diptera are numerous and largely allied to American genera or species,
and the non-endemic forms mostly Chilean. I have not been able to get a proper
insight into the distribution of the many genera found elsewhere. Little can be
said about the beetles until Dr. KUSCHEL’s material has been described. Two
presumably austral-bicentric cases are noticed, Pycnomerodes and Eleuszs. As in
so many oceanic islands there is in Juan Fernandez a remarkable display of endemic
wingless Curculionids, living on the endemic plants of South American or Antarctic
parentage, and examples of strict specialization are known. Host and lodger look
back upon a long common history, but whether this implies a common original
ancestral home or adaptation in the islands I cannot tell. Of Hymenoptera, Hap/o-
gonatus, Prenolepis and; perhaps, J/etelza show Antarctic connections.

The antiquity of the endemic land shelis cannot be disputed. The Tornatellids
are an ancient Pacific group and their presence in Juan Fernandez as well as in
other isolated islands and archipelagoes has been considered a proof of former
land connections. Their display in Hawaii is unparalleled. GERMAIN summarized
his opinion on the evolution of the Hawaiian fauna in the following terms which,
mutatis mutandis, apply also to Juan Fernandez (z05. 995).

S’il est bien ainsi, cette famille primordiale [the ancestors of Achatinellidae, Amas-
tridae, Leptachatinidae and Tornatellinidae|} doit avoir une trés ancienne origine et
remonter au Paléozoique.... le peuplement malacologique de l’archipel des Hawaii
est fort ancien et doit remonter a des temps primaires. Il n’a put se faire, comme le
preuvent les développements précédents sur la répartition des genres et des espeéces,
que sur une aire contenue, ce qui exclut la possibilité de considérer les iles Hawaii
comme le resultat de l’activité des volcans sous-marins.

26 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

402 C. SKOTTSBERG

Finally, let us try to reconstruct the history of our islands, beginning with
the time when there existed a ‘Tierra de Juan Fernandez” in BRUGGEN’s sense.
It must have become isolated and reduced in size rather early. We do not know
if the fauna included vertebrates other than birds; if it did they did not survive
the long volcanic period. Unfortunately we know too little of their history in
Chile, when they first appeared in modern forms, and so forth. The absence of
all gymnosperms is difficult to explain. The Eocene flora of Chile contained several,
Araucaria, Libocedrus and Podocarpus, all still living there and accompanied by
Fitzroya, Saxegothaea, Pilgerodendron and Dacrydium, and even if no close land
connection existed, some of them ought not to have had much greater difficulties
to get transported across the water barrier than some of the angiosperms found
on the islands. If, on the other hand, a land bridge existed, I can see no obvious
reason why conifers did not use it or, if they did, why they didn’t take possession
of the new volcanic soil. Introduced araucarias, pines and cypresses do well on
Masatierra. The only possibility, remote perhaps, could be that they had not been
able to spread as far as to Juan Fernandez when the connection was cut. Their
absence gives us no clue to the time when this happened.

Another element in the Chilean flora would seem to come to our rescue, the
Nothofagus flora. In my sketch of 1925 (23z. 33) I expressed the idea that “the
connections between the islands and the main land were severed before the south
Chilean flora assumed its present composition, and also before the advancing
Nothofagus flora reached these latitudes”, and BRUGGEN is of the same opinion.
He states that tropical South America is the ancestral home of much of the
Chilean forest flora, and adds:

Pero, la actual flora de Chile central contiene, ademas, una mezcla con una flora
de clima mas fresco, que después de la separacion de la Tierra de Juan Fernandez
inmigr6é y que se caracteriza por los géneros WVothofagus, Araucaria, etc.

Araucaria was found in the Pichileufii beds which are supposed to be Eocene,
whereas the southern beeches appear in this latitude considerably later. In the
Magellanian region DUSEN, as we have seen, distinguished two horizons, a lower,
Oligocene, with Vofhofagus, and an upper, Lower Miocene, with Avaucaria. Accord-
ing to BRUGGEN the separation of Juan Fernandez from Chile took place during
the Oligocene Navidad transgression. We would think that, if the bridge lasted
longer, the Nothofagus flora ought to have invaded Juan Fernandez and to have
found suitable stations in the montane belt, and that the rising volcanic islands
offered acceptable habitats for a genus of such wide physiological amplitude.

Anyhow, during the final upheaval of the Andes when, in any case, most
of the land disappeared, leaving the submarine ridge with the rising volcanic
masses standing, Central Chile definitely ceased to be a source of the island flora.
This Chilean flora is, as REICHE pointed out, a product of the Andes with addition
of an Antarctic element.

Submergence resulted in the displacement of the vegetation belts and in much
loss of life when the volcanic eruptions began and lava flows and other ejected
material covered part of the country. Long before that, important changes had

DERIVATION OF THE FLORA AND FAUNA 493

taken place as a consequence of the cooling of the climate in post-Eocene time,
and with the widening space of sea between the islands and the mainland the
insular climate became more and more oceanic.

The rise of the Cordillera was an affair of great magnitude and long duration,
and millions of years passed before the last remnant of the Juan Fernandez—
Desventuradas ridge disappeared. Two eruption centres were formed, Masatierra +
Santa Clara, and Masafuera. The first two form an arc, suggesting a remnant of
a gigantic crater, but the place must be properly sounded in order to allow us to
prove or disprove this hypothesis. I cannot even guess where the eruption centre
of Masafuera should be looked for. Abrasion has, as could be expected, forced
the coast escarpment back more on the west than on the east side, reducing the
area of the island. A description of the topography of the islands is found in this
volume, pp. 89-168. There is no sign of recent activity anywhere, no parasitic
cones, no hot springs. When the map in “Der neue Brockhaus’’ (1938) marks
Juan Fernandez as a seat of active volcanoes, this applies, I suppose, to the
submarine eruption in 1835 near Masatierra and to the phenomena observed at
San Felix some years ago in connection with an earthquake on the mainland,
but otherwise nothing in the way of activity has been recorded after the tsunami
in 1751, when also an earthquake occurred, and other earthquakes were registered
in 1809, 1822 and 1835 (275).

As I said, I believe that Masatierra and Masafuera represent two separate
centres; the distance between them is 92 nautical miles and the sea is deep.
Whether they are of exactly the same age and became extinct at the same time
is difficult to tell; to judge by the topography, Masafuera makes an impression
of being much less eroded, and, as a consequence, younger, but the difference is,
I think, mainly due to differences in the basalt; the petrographical structure is
not quite the same. It is supposed that the eruptions began during Pliocene and
lasted a very long time and that part of the submarine ridge was still above water
when the main eruption centres became extinct. Otherwise the result would have
been two lifeless islands without a sign of the old endemic biota. The flora and
fauna were inherited from the sinking land. The process is easily observed in
many volcanic islands and nowhere to greater advantage than in the island of
Hawaii. As soon as the lava has cooled down, plants get established, microscopic
algae, modest tufts of mosses and particularly lichens of the genus Stereocaulon
(341), but a conditio sine qua non is that moisture is available, that erosion sets
in and soil is formed. Even under very favourable conditions and with the
sources for repopulation next door, it will take a long time before a closed vege-
tation cover gets established. With regard to Juan Fernandez, nothing much
could happen until the islands had been built up to a considerable altitude,
undoubtedly greater than now, when streams rushed down the mountain slopes
and started to excavate valleys, for no plant cover, not to speak of forest growth,
could get established until erosion and abrasion had done part of their work.
This means that a good deal of the fundament, now hundreds of metres below
the surface of the ocean, was still exposed and retained a portion of the original
flora and fauna, which became the principal source of the flora and fauna of the

404 C. SKOTTSBERG

volcanic soil. It goes without saying that chance played a dominant role and that
the fragmentary character of the island world is easy to understand. It is also
possible to understand why different species happened to become isolated on
Masatierra and Masafuera. According to JOHOW Masafuera was populated through
overseas transport from Masatierra, and this explained why the former was so
much poorer, but only 50% of the vascular plants found on Masafuera were known
to him. Other reasons for the dissimilarities between the islands are differences
in the topography, particularly in altitude, Masatierra is 915, Masafuera 1570 m
high. If we could lower Masafuera 650 metres, the entire highland region with
its special flora and fauna would disappear. The question of the origin of the
alpine flora is, as we have seen, difficult to answer. Did it exist in the islands
before the separation from Chile took place, and were there any habitats where
it could thrive? Many of the species are of Antarctic origin and immigrated to
the extreme south of America, with or perhaps after the Vothofagus flora, where
they abound in the bogs of the rainy zone and in the mountains above the
timberline. And if, as was explained above, the Vothofagus flora never had an
opportunity to spread to Juan Fernandez over land, nor were those Magellanian
plants able to come. There are several montane plants in Masafuera which un-
doubtedly date back to early times, but they are of different origin: the species
of Erigeron, Euphrasia formosissima, Megalachne masafuerana, Phoenticoserts regia,
Ranunculus caprarum and Robinsonia Masafuerae, all of them peculiar endemics,
and of these Ranunculus and Megalachne are found only along the high ridge
above 1300 m. Even if some of them are of Antarctic ancestry, their history is
another. When it comes to the Magellanian group, Oreobolus, Lagenophora,
Gleichenia, and so forth, it is difficult to exclude the possibility that they be
glacial or postglacial immigrants. Carer Banksit, Empetrum rubrum—only a single
plant seen—Galium masafueranum, Gnaphalium spiciforme, Myrteola nummularia
and Rubus geoides are, as was already told, not of Antarctic origin, but may have
extended far south after the recession of the ice and accompanied Oreodolus, etc.
And we cannot refuse to admit that various Chilean ferns, bryophytes and lichens,
belonging to the forest, were transported across from South Chile, for even if the
prevailing winds are westerly, storms from other directions occur, and there has
been plenty time. Still, the floristic difference between the two islands is a warning,
not to put too much faith in the efficiency of the natural agents, among which,
in this case, the wind stands foremost.

The biological differences between Juan Fernandez and the mainland increased
during the Ice Age. West Patagonia and Fuegia were covered by inland ice (z67);
if small, ice-free refuges occurred has not been definitely stated, but is not alto-
gether improbable; their part taken in the repopulation of the country was, however,
of minor importance. With the north-south trend of the Cordillera, the road toward
north lay open, and the big island of Chiloé was not ice-covered, allowing the
subantarctic flora and fauna to survive, perhaps also some of the hardy trees and
shrubs. The high crests of the Andes in Central Chile were covered by glaciers
descending into the valleys; squeezed between the mountains and the coast a
migration back and forth went on during the successive interglacial and glacial

DERIVATION OF THE FLORA AND FAUNA 405

periods. At times the subantarctic bog and heath must have occupied considerable
areas north of their present range, but during all this shifting to and fro many
species may have been lost, some of them surviving on Masafuera. The glaciers
did not come down to the coast of Central Chile, but the climate was cold and
numerous ancient stenotopic types, some of them surviving on Juan Fernandez,
perished.

The influence of the glacial periods cannot have been very destructive on
the islands. Some plant species reduced their range and became rare or extinct;
several, apparently with a very narrow physiological amplitude, are on the verge
of extinction today. In Masafuera, the timberline was lowered, I suppose, and the
forest patches in the valleys came down toward the sea. Masatierra has no climatic
upper timberline. In a depression on the summit of the highest peak, El Yunque,
Drimys, Fuania, Cuminia, Escallonta, Rhetinodendron, Dicksonia, etc. etc., luxuriate
more than anywhere else, thanks to the constant humidity.

Part 2.

EASTER ISLAND.

Chapter XIV.
Composition, distribution and relationships of the Flora.

I. Angiospermae.
Gramineae.

Paspalum \V.. More than 200; trop. to temp., most numerous in Amer.
forstertanum Fluegge. N. Caled.
scrobiculatum VL. var. orbiculare (Forst.) Domin. N. Guin., Austral., N. Caled.,
Polyn.
Axonopus Beauv. About 75, the majority in N. and S. Amer.
paschalis Pilger. Related, according to PILGER, to A. scoparius (Fluegge)
Pilger, a S. American species.
pa is
horridula Pilger. Related species in S. Amer. and Austral.
Sporobolus R. Br. Over 100, trop.subtrop., most numerous in Amer.
elongatus R. Br. |as indicus (L.) R. Br. in 2370]. S. As., Malays., Austral.,
Nes Zeal.
Calamagrostis Adans. About 150, mostly temp., north and south.
retrofracta (Willd.) Link. (Agrostis Willd., A. filiformis (Forst.) Spreng.) Aus-
tral., Tasm., N. Zeal., Polyn., Hawaii.
Dichelachne Endl. 2, Austral., N. Zeal.
sciurea (R. Br.) Hook. fil. Austral., N. Zeal.
Danthonia DC.
paschalis Pilger. Perhaps most nearly related to D. chzlensis Desv.
Eragrostis Host. Over 200, trop.—subtrop.
elongata Jacq. E. Ind. to Malays. and Polyn.

Cyperaceae.

Cyperus L.

eragrostis Lam. Very likely introduced, and this may be the case also with
the other species.

+ The area of genera and species also found in Juan Fernandez was indicated in Pt. 1 and
1s not repeated here.

DERIVATION OF THE FLORA AND FAUNA 407

polystachyus Rottb. Trop.—subtrop., wide-spread.

cylindrostachys Boeck. As the former.

brevifolius (Rottb.) Hassk. Pantrop.
Scirpus L.

riparius Presl. N. Amer., Calif. to S. Amer., south to Fueg., Falkland. I can-
not find that var. pfaschalis Kuekenth. deserves to be distinguished.

Juncaceae.

Funcus L.
plebeus R. Br. Colomb.—Urug., Austral., Tasm., N. Zeal.

Piperaceae.
Peperomia Ruiz et Pav.
reflexa Dietr. Pantrop., very widespread, north to Hawaii and south to
N. Zeal.; also on Rapa and possibly Pitcairn.

Chenopodiaceae.

Chenopodium ambiguum R. Br. Austral., Tasm., N. Zeal.

Polygonaceae.

Polygonum LL. About 150, world-wide.
acuminatum H.B.K. W. Ind., Centr. and S. Amer., Galdap. Is., trop. and
S. Afr., Orient.

Nyctaginiaceae.
Boerhaavia L. About 20. Pantrop.
diffusa 1. Widespread in the Pacific and a common weed in the Old and
New World; probably of aboriginal introduction.

Aizoaceae.
Tetragontia L.

expansa Murr.

Cruciferae.

Nasturtium R. Br. About 50. Widespread, mostly temp.
sarmentosum (Sol.) O. E. Sch. Austral—Polyn., Hawaii.

Leguminosae.
Caesalpinia L. At least 90-100; pantrop.
bonduc (L.) Roxb. Trop. As.—Polyn.
Sophora L.
toromiro (Phil.) Skottsb. Nearly related to the species from J. Fern.

Euphorbiaceae.
Euphorbia L. 1300-1600; world-wide.
hirta 1. Indomal.—Polyn., often adventitious.
serpens L. As the former, also common in trop. Amer.

408 C. SKOTTSBERG

Umbelliferae.
Apium LL.
prostratum Labill. Austral-circump., incl. A. australe Thouars.

Primulaceae.

Samolus L. 9 (127), 1 cosmop., 2 (5?) N. Amer., 3, southern S. Amer. iy oayamis
1 W. Austral., and the following.
repens (Forst.) Pers. Austral-circump.

Gentianaceae.

Erythraea Neck. (Centaurium Hill.) 30-40, subtrop.temp.
australus K. Br. Austral:, N. Caled., Fiji.

Convolvulaceae.
Calystegia R. Br. 7-8, temp.—subtrop.
sepium (L.) R. Br. forma. All continents, also reported from Australia, but
possibly introduced in the s. hemisph.
Tpomaea L.
pes caprae (L.) Roth. Pantrop.

Solanaceae.

Lyctum \.. About 100, most numerous in S. Amer.
carolintanum Walt. var. sandvicense (Gray) L. C. Hitchc. Rapa, Hawaii.

The flora is extremely poor, not much richer than in the low coral islands:
16 families, 26 genera and 31 species, and I am not at all sure that all of them
are indigenous and did exist here before man appeared on the scene; some may
have been accidentally or purposely introduced by the aborigines, by the American
whalers and in modern times. With regard to the “endemic” Solanum Insulae
Paschalis Bitter, see 249; it was used as medicine. Four species are endemic
(13%), but three of them belong to large grass genera needing monographic study.
FORSTER mentions (347), beside some cultigens, only Panicum filiforme Jacq.
(= Digitaria sanguinalis), a common weed, Sheffieldia (=Samolus) repens, Avena
filiformis (=Calamagrostis retrofracta) and Solanum nigrum; in his journal (348)
he refers to Apzwm, which he knew from New Zealand, to “J/émosa’’ (= Sophora),
and also to the former occurrence of savdalwood. In regard to the distribution
of Saztalum in Southeastern Polynesia, where S. zzsudare Bert., including varieties,
is known from Tahiti, Raiatea, Marquesas, and Austral Is., and S. hendersonense
F. B. R. Brown (very close to the former) is found on Henderson Island in the
extreme south-east of the island swarm, it did not seem incredible that sandal-
wood once occurred on Easter Island, perhaps introduced by the aborigines; not
very long ago S. yasz Seem. was introduced to Tonga and is well established there.
The matter is sufficiently interesting to be discussed here.

Tradition tells that when Hotu Matua, the legendary hero of the Easter
islanders, took possession of the island, he brought various useful trees and other
plants, a story first told by FORSTER (348. 583). METRAUX (zo. 15-17) enu-

DERIVATION OF THE FLORA AND FAUNA 409

merates mahute (Broussonetia papyrifera), makoi (Thespesia populnea), hauhau
(Triumfetta semitriloba), toromiro (Sophora), naunau (Santalum), and marikuru
(Sapindus saponaria). None are of American origin; all point towards Malaysia-
Polynesia.

In his description of a beautiful wooden hand presented to him by one of the
natives and now in the British Museum, FORSTER says that “the wood of which
it was made was the rare perfumed wood of Taheitee, with the chips of which
they communicate fragrance to their oils’’, undoubtedly sandalwood, with which
FORSTER was familiar. The natives were expert in the art of wood-carving, but
the hand is very unlike all other objects, which were made of Sephora wood,
called toromiro, while the name of the tree with the fragrant wood was xaunau.
COOKE (343. 722) translated this word “bastard sandalwood’’. This is the name
given to Myoporum sandvicense Gray in Hawaii, where the native name is azo
(naeo, naieo); Sazfalum is called z/zahz, in Marquesas puahz, on Rapa eahz, in
Tahiti and Tonga az. Whether or not the words xaunau and xazo have the
same base I cannot tell; on Raivavae, Rapa and New Zealand the name of
Myoporum is ngato, whereas true sandalwood, in this case Mida, is called
mazre, the Hawaiian name for the fragrant Alyaza oliviform7s Gaud., transferred
to Mida in New Zealand, where no A/yaza occurs.

Our main sources of Easter Island ethnology, ROGGEVEEN, LA PEROUSE,
Cook, Mrs. ROUTLEDGE, etc. do not mention sandalwood, while METRAUX pays
special attention to this subject, referring to FORSTER (7So. 17-18).

My informant gave “‘sandal’’ as the Spanish equivalent of the word nxaunau—
the correct Spanish word is sdindalo—and remarked that the tree had entirely disap-
peared since the time of SaLmon (between 1880 and 1890). The last one on the island
grew near Vai-mata, but died recently “‘because there were no more kings’’. From the
distribution of the sandalwood it seems likely that the sawnaw was the true sandal-
wood and not the bastard. Sandalwood “‘is found on the atolls of Elizabeth and Ducie,
the nearest islands to Easter Island’’.

Elizabeth I. is another name for Henderson, a rocky coral island; Ducie is
not quoted by Brown (35.1). To judge from the distribution, zazau could
just as well refer to Myoporum, a genus represented by endemic species in Rapa
—where, as we shall learn presently, aw means something quite different—
and Austral Is. but never reported from Easter Island. The wood, slightly fra-
grant, is said sometimes to have been used as a substitute for sandalwood in
Hawaii. METRAUX’s informant seems to have been convinced of the earlier ex-
istence of real Saxta/um on his island. If so, and if it had been brought from
Tahiti, it ought to have had the same name.

The story. does not end here. One of my correspondents, Mr. PAUL H. STEELE
of Sacramento, Cal., kindly called my attention to a book written by Padre
SEBASTIAN ENGLERT, who has lived more than 15 years on Easter Island (86),
a place in which Mr. STEELE is particularly interested. As I had no opportunity
to consult the book, Mr. STEELE copied and sent me the following remarkable
passage.

410 C. SKOTTSBERG

Nau o Naunau (Santalum): arbusto de la familia de las santalaceas, también lla-
mado max opata, porque crecia en los barrancos (opata) de la costa, entre rocas y
piedras. Ahora ha desaparecido. Los ultimos ejemplares que algunos de los nativos
actuales recuerdan haber visto todavia, se han secado hace unos 50 anos. El mau opata
daba, como frutos, nueces del tamafio de castafias, los “mako’i nau opata’’. Carl
Friedrich Behrens nombra nueces entre los frutos que los islefios les regalaron en gran
numero a él y a sus compafieros. Hotu Matua y su gente parecen haber traido gran
cantidad de estas nueces, porque de ellas se alimentaron en los primeros meses des-
pués de haber llegado a la isla. Al excavar la tierra en cuevas que estaban antigua-
mente habitadas se encuentran cdscaras de estas nueces. Estas cdscaras generalmente
no estén quebradas sino que han sido abiertas en forma de un pequeno circulo, para
ser usadas por los nifios en el juego del trompo. La madera del arbusto se utilizaba,
por su exquisito aroma, para confeccionar un perfume, como los veremos en otro capitulo.

This description does not at all fit either Saztealum or Myoporum. The fruit
of Santalum is an ellipsoid drupe with a thin fleshy mesocarp and a very hard
endocarp, and I have never seen or heard of a kind the size of a castafia (chest-
nut); in the largest I have measured (S. pyvularium Gray) the drupe was 16-18 mm
long and the stone 12-15 mm. According to HILLEBRAND (307. 390) the drupe
measures up to 24 mm in length, but I have not seen any as large as that. Nor
have I ever heard that the kernel is used as food; the idea that Hotu Matua’s
party could have maintained itself for months on nothing else is preposterous,
and I fail to see how the stone could be used as a whipping-top. The zawnau which
grew along the coast and produced the ‘‘cascaras’’ found in the caves cannot
have been a species of Santalum. Fortunately this could be proved. Mr. STEELE
had told me that Father ENGLERT had sent him two shells for his collection
of Easter Island curios and had promised him more, of which he intended to
send me samples. As time went by and no more came I asked him the
favour of sending me one of his precious specimens as loan, and he willingly
consented. It is a hard, brown and smooth, almost globular shell, 2.5 cm high,
3 cm wide, 2 mm thick, with a large irregular hole in the basal part. It has
nothing whatever to do with Saztalum. A passage in METRAUX’s book, p. 353,
put me on the track. He quotes a song which the children used to sing when
the tops were spinning, and it tells that the spinning-tops were made of makoz
—Thespesia populnea capsules! A comparison with herbarium material showed
that Mr. STEELE’s specimen is a typical capsule of 7hespesza, one of the “nuts”
found in the caves. They made very poor food but good spinning-tops. We
did not see 7hespesza on the island, but METRAUX observed it growing on the
cliffs at Poike (the eastern headland). Evidently the word xau or naunau has
been altogether misplaced by the Easter islanders, although both METRAUX
and ENGLERT were told that it was the name of Sanfalum. We find the same
word in Tahiti and the Tuamotu Islands for Lepidium bidentatuy: Montin, and
this is called xaupata, strikingly like ENGLERT’s xau opata, in Marquesas; in
Tahiti zaupata means Scaevola frutescens, which is called xaupaka in Hawaii and

ugaungau in Rarotonga, and on Rapa zau is used for Sonchus oleraceus. All
these plants were used as medicine.

DERIVATION OF THE FLORA AND FAUNA 4IiI
Geographical elements,

In 1934 I made an attempt to arrange the vascular plants according to their
distribution, with the following result (239. 278-279, translated from French).

1. Australian-Polynesian element (12).

a) Species found elsewhere: Paspalum (2), Sporobolus, Calamagrostis, Dichelachne,
Eragrostis, Juncus, Peperomia, Chenopodium, Nasturtium, Erythraea.

b) Endemic: Stipa horridula.

Palaeantarctic element (3).

a) Species found elsewhere: Apium, Samolus.

b) Endemic: Sophora toromiro.

American element (6).

a) Species found elsewhere: Cyperus vegetus, Scirpus, Polygonum, Lycium.

b) Endemic: Axonopus paschalis, Danthonia paschalis.

4. Wide-spread tropical element (10): Cyperus (3), Boerhaavia, Tetragonia, Caesalpinia,

Euphorbia (2), Calystegia, Ipomaea.

oP)

Going into more detail, I have tried to rearrange the angiosperms in the
following manner.

I. Palaeotropical element.—22 sp. (70 %).

1. Pantropical (6): Cyperus polystachyus, cylindrostachys and brevifolius,
Peperomia reflexa, Boerhaavia diffusa, Ipomaea pes caprae.

2. Malaysian-Polynesian (9): Paspalum forsterianum and scrobiculatum, Sporo-
bolus elongatus, Calamagrostis retrofracta, Eragrostis elongata, Nasturtium sarmen-
tosum, Caesalpinia bonduc, Euphorbia hirta and serpens.

3. Australian-Polynesian (7):

a. Endemic: Stipa horridula (?).

6. Not endemic: Dichelachne sciurea, Chenopodium ambiguum, Tetragonia
expansa, Erythraea australis, Calystegia sepium (?).

c. Also in tropical America: Juncus plebetus.

II. Austral-circumpolar element.—4 sp. (12.9 %).

a. Endemic: Danthonia paschalis, Sophora toromiro.
6. Not endemic: Apium prostratum, Samolus repens.

III. Neotropical element.—5 sp. (16.1 %).

a. Endemic: Axonopus paschalis.
6. Not endemic: Cyperus eragrostis, Scirpus riparius, Polygonum acumina-
tum, Lycium carolinianum var.

I am strongly inclined to regard Cyperus eragrostis as a late immigrant from
Chile. Lycitum carolintanum var. sandvicense is a puzzling case, a form of a
North American species but only found in Hawaii, Rapa and Easter Island. It
is a halophyte confined to the beach, but one might expect it also to occur on
the shores of California etc., from where it has not been reported. Beach plants

412 C. SKOTTSBERG

are included in all the groups, in 1 /pomaca, Caesalpimia, Chenopodium and Tetra-
gonia, in 11 Apium and Samolus, in UI Lycium, together they are 7, or 21% of
the angiosperms.

If we want to trace an ancient Antarctic source, just as we did in Juan
Fernandez, the Antarcto-tertiary group would consist of Danthonia, Funcus,
Sophora, Apium and Samolus.

To judge by the geographical position of Easter Island the dominance of a
Palaeotropical-Pacific element was expected. The island certainly is remote from
the large cluster of islands of South-eastern Polynesia, but much more distant
from South America. Consequently, the presence of a neotropical element is
surprising. Future researches will perhaps show that Sipa horridula should be
added, which makes little difference as long as the endemic Axonopus is
claimed to be of neotropical parentage, and Scrrpus riparius and Polygonum
acuminatum remain American. Their mode of occurrence and ecology oblige us to
regard them as truly indigenous, unless they have been intentionally introduced
in prehistoric time during one of the mythical cruises which, according to HEYER-
DAHL, put Easter Island in contact with Peru. A direct transport of seeds across
the ocean without man’s assistance is difficult to imagine, and it is futile to
speculate in land connections.

II. Pteridophyta (63, 279).

Polypodiaceae (in the old sense).
Asplenium L.
adiantoides (L.) C. Chr. var. squamulosum C. Chr. et Skottsb. The species
trop. As.—Austral., N. Zeal. and Polyn.; also E. Afr. and adjacent islands; the
variety endemic.
obliquum Forst. Austral-circump.; the only plant reported from Sala y Gomez.
Davallia Sm. About 40; wide-spread, mainly southern; As.—Polyn., S. Afr.,

Madag., a solitary species Ibero-Afr. and Macaron.
solida (Forst.) Sw. Indomal.—Polyn.—Austral.

Doodia R. Br. 12. “New Zealand and Juan Fernandez to Hawaii and Australia”
(COPELAND 69.158), but “Juan Fernandez” is a mistake for Easter I.

paschalis C. Chr. et Skottsb. Close to D. dlechnoides A. Cunn. (Austral.).
Dryopteris Adans.

Espinosai Hicken. Belongs to a neotropical group.

gougylodes \(Schk.) O. K. Pantrop.

dentata (Forsk.) C. Chr. Pantrop.
Elaphoglossum Schott.

Skottsbergii Krajina. Related to E. tahitense Brack.
Microlepia Presl. 45-50; a pantrop. genus ranging north to Japan and south to
Madag. and N. Zeal.

strigosa (Vhunb.) Presl. Indomal.—Polyn.

DERIVATION OF THE FLORA AND FAUNA 413

Polypodium U.

scolopendria Burm. Palaeotrop., wide-spread.

Polystichum Roth.
Fuentesii Espinosa. Belongs to the P. westitum assemblage.

Vittarta Sm. About 80, pantrop.—subtrop.
elongata Sw. Indomal.—Polyn.—Austral.

Ophioglossaceae.
Ophioglossum L.
lusitanitcum \.. subsp. corzaceum (Cunn.) Clausen. Boliv—Chile, Austral., Tasm.,
N. Caled., N. Zeal.
reticulatum L. S.E. As., Philipp., N. Guin., Melan—Polyn., east to Mangareva;
Mascarene Is.

Psilotaceae.

Psitlotum Sw. 2 wide-spread sp.
nudum (L..) Griseb. Pantrop.

Fifteen species are listed, 4 of them endemic (26.6%), the endemic variety
of Asplenitum adiantoides not counted.

Geographical elements.
In my earlier subdivision (279) the following groups were distinguished.

1. Australian-Polynesian element (6).
a) Species found elsewhere: Asplenium adiantoides, Elaphoglossum tahitense, Micro-
lepia strigosa, Vittaria elongata, Ophioglossum coriaceum.
b) Endemic: Doodia paschalis.
2. Palaeantarctic element (1).
Found elsewhere: Asplenium obtusatum.
. American element (1).
Endemic: Dryopteris Espinosai.
4. Wide-spread tropical element (4): Dryopteris gongylodes and parasitica (=dentata),
Polypodium phymatodes, Ophioglossum reticulatum.

The list contains I2 species; two have been reported later and some nomen-
clatural changes have been made. A new geographical arrangement follows here.

I. Palaeotropical element.—11 sp. (73.3%).

1. Pantropical (3): Dryopteris gongylodes and dentata, Psilotum nudum.
2. Palaeotropical (1): Asplenium adiantoides (endemic variety).
3. Malaysian-Polynestan (6).
a. Endemic: Elaphoglossum Skottsbergii.
6. Not endemic: Davallia solida, Microlepia strigosa, Polypodium scolo-
pendria, Vittaria elongata, Ophioglossum reticulatum (also Mascarene Is.).

4. Australian-Polynesian (1).
Endemic: Doodia paschalis.

414 C. SKOTTSBERG

Ul. Austral-circumpolar element.—3 sp. (20 %).

a. Endemic: Polystichum Fuentesil.
4. Not endemic: Asplenium obliquum, Ophioglossum lusitanicum subsp.

coriaceum.

III. Neotropical element.—r sp. (6.7 %).
Endemic: Dryopteris Espinosai.

We find the same dominance of a palaeotropical element as among the an-
giosperms. Ophioglossum lusitanicum is reported from Atlantic Europe, the Medi-
terranean and Macaronesia, whereas the subspecies is austral-bicentric, an example
of a remarkable discontinuous distribution. The American element is represented
by a single endemic species and there is no fern corresponding to Polygonum
acuminatum and Scirpus rviparius. Polystichum Fuentesti and Asplenium obliquum
are, perhaps, Antarctic, and the latter was classified as such above (p. 282), but it
is a seaside plant. If we follow COPELAND, Asplentum, Doodia and Davaliza are of

Antarctic origin.
Ill. Musci (33, 265).

I have to thank Dr. HERMAN PERSSON for kind assistance in finding out
about distribution.

Trematodon Michx (Dicranaceae). About 70, mainly trop., south to N. Zeal.
pascuanus Ther.

Campylopus Brid.
introfiexus (Hedw.) Mitt. See p. 227.
turficola Broth.
hygrophilus Broth.
dicranodontioides Broth.
saxicola Broth.

Fissidens Hedw.
pascuanus Broth.

Ptychomitrium (Bruch) Fuernr.
subcylindricum Ther.

Weisia Hedw. (Trichostomaceae). About 30, widely distributed (Eur., N. Amer.,
Ss, Se oe ey
flavipes Hook. fil. et Wils. Java, Ceylon, E. Austral., Tasm., N. Zeal.

Bryum Dill.
argenteum LL. var. lanatum (Palis.) Bryol. eur. Widely spread in warmer
countries, the species cosmop.
Philonotts Brid.
laxisstma (C. M.) Bryol. jav. E. Ind., Madag.
Papillaria C. M. (Meteoriaceae). About 70; a pantrop. genus.

DERIVATION OF THE FLORA AND FAUNA 415

pascuana Ther. ex Broth. Related to P. crocea (Hpe) Jaeg. Ind., Ceylon,
Java, Philipp., E. Austral., N. Zeal., Kermadec Is., Fiji).
Fabronia Raddi (Fabroniaceae). go-100, half of them Amer.; trop.—subtrop.

macroblepharoides Broth. Related to /*. macroblepharis Schwaegr. (Brazil);
other related sp. in Afr. and Austral.

Rhacopilum Palis.
cuspidigerum Schwaegr. Norfolk I., N. Caled., Samoa, Hawaii.

Besides, BROTHERUS mentions [Vezsza sp. and Macromitrium sp.

Fourteen species are reported, 9 of them endemic (64.3%), a high figure,
but there can be no doubt that only a part of the mosses has been collected,
and very likely the proportion of endemics will be reduced. Of greater interest
are the endemic aquatic species of Campylopus of the crater lakes, possibly relicts
from an era when the island was larger and higher than now and enjoyed a more
humid climate. Probably many mosses and other cryptogams disappeared with
the forest groves. Of the known species Campylopus introflexus, Wetsia flavipes
and Khacopilum cuspidigerum lave a southern distribution, PAzlonofis laxissima
is palaeotropical and Aryum argenteum cosmopolitan and perhaps anthropochorous;
the endemic species are supposed to have tropical relatives.

IV. Hepaticae (730).
Frrullania Raddi.
lagenifera Schwaegr. Known before from the type locality only, said to be
the Falkland Is., but this statement is subject to doubt; possibly Ins. Marianae
is meant instead of Maclovianae.

Conditions are not favourable to hepatics, but more species will be found.
A Lejeunea was collected, but the material seems to have been lost (l.c. 699).

V. Lichenes (297).

My thanks are due to Dr. A. H. MAGNUSSON and Dr. R. SANTESSON, who
helped with information on the distribution.

Arthonia Ach. About 500, the majority trop.—subtrop.
fuscescens Fée. Apparently only reported once before (trop. Amer.).
Opegrapha Humb. About 280, mostly warmer climates.
paschalis Zbr.
Graphis (Adans.) Ach.
lineola Ach. Trop.-subtrop.
Diploschistes Norm. About 30, cold to temp. regions, trop. mountains.
anactinus Zbr. Described from Japan.
scruposus (L.) Norm. Wide-spread N. and S. temp.
Hleppia Naeg. About 40, mostly warm and dry regions.
Guepini Nyl. N. Amer., Eur.

416 C. SKOTTSBERG

Lecidea (Ach.) Th. Fr.
paschalis Zbr.
Cladonia (Hill) Vainio.
pityrea (Flk.) Fr. Cosmop.
Acarospora Mass.
Skottsbergii Zbr.
Parmelia (Ach.) De Not.
reticulata Tayl. N. and S. Amer., W. and S. Eur., Afr., E. As., Austral.
conspersa (Ehrh.) Ach. var. /usitana (Nyl.). S. Eur.; the species cosmop.
Usnea Wigg.
subtorulosa (Zbr.) Motyka (344). Masafuera. Described as U. Stezneri var. by
ZAHLBRUCKNER, who also distinguished var. “#zcfa Zbr. and quoted it for Easter
Island: this is called U. tcta by MotTyKA, who records it for S. Amer. only.

Caloplaca Th. Fr:
Filing Abr. |. Pert.
lucens (Nyl.) Zbr. Patag., Falkl., S. Georgia.
Buellia De Not.
stellulata (Tayl.) Mudd. Cosmop.
Jernandeziana Zbr. J. Fern.
halophiloides Zbr. var. The typical sp. J. Fern.
paschalis Zbr.
glaztouana M. Arg. Brazil.
Rinodina (S. Gray) Mass. About 300; very widely distributed.
Perousii Zbr.
Pyxine (Fr.) Nyl.
enteroxantha Ny|l. forma. S.W. Eur., Japan.
Physcia (Schreb.) Vainio.
picta (Sw.) Nyl. Wide-spread trop.—subtrop.
Anaptychia Koerb.
speciosa (Wulf.) Mass. Widely distributed; in Amer. south to Fueg.

These 23 species, 5 regarded as endemic, represent, I am sure, only a minor
part of the lichen flora and do not lend themselves to geographical speculations.
There are several strange cases of disjunction serving, I daresay, to illustrate our
insufficient knowledge of the distribution of lichens.

V. Fungi.

Our collection contained a single species, Bovzstella pusilla Lioyd, known
before from Australia (zo2).

DERIVATION OF THE FLORA AND FAUNA 417

Chapter XV.
Composition, distribution and relationships of the Fauna.

Indigenous vertebrates, birds excepted, lacking. The principal occupation in
the island is farming, cattle and sheep are plentiful and roam over the island
which, with the exception of outlying rocks, has lost its primitiveness. As a con-
sequence of the changes in the plant cover, particularly the extermination of the
indigenous trees, also the fauna was impoverished, while through the introduction
of useful plants, numerous weeds and all kinds of goods many foreign insects
and other invertebrates made their appearance, as the lists below will show. As
little research work has been done hitherto, many more species will probably be
found, indigenous as well as introduced.

Aves (775).

Sterna lunata Peale. Molucc., Polyn., Fiji, Hawaii.

Anous stolidus (L.) unicolor Nordmann. Sala y Gomez. The typical species
trop.-subtrop., but not observed on the coast of America.

Procelsterna coerulea (Benn.) skottsberg? Loennb. Typical coerulea on Christmas
I., 4 other subspecies scattered over the Pacific.

Gygis alba (Sparrm.) royana Matthews. With the typical species wide-spread
trop.

Pterodroma heraldica Salvin paschae Loennb. The typical species S.W. Pacific.

Sula cyanops (Sundev.). Trop. seas throughout the world.

According to the natives some other sea birds occur, but there are no land
birds.

Oligochaeta (757),

Pheretina californica (Kbg). Introduced. Reported from Calif., Mex., Madeira
and Lower Egypt.

Araneida (22).

Scytodes lugubris TVhorell. Burma, N. Caled., perhaps all over Oceania.
Photcus phalangioides Fuessli. Eur., now spread over a large part of the globe.
Theridium tepidariorum C. Koch. Cosmop.

+ Tetragnata Paschae Berland. A large cosmop. genus.

Corinna cetrata Simon. N. Caled.

Hlasarius Adansoni Audouin. Cosmop.

Plexippus Paykulli Audouin. Cosmop.

Possibly all the spiders are adventitious (BERLAND l.c.). Two species were
determined as to genus only.

27 — 557857 The Nat. Hist. of Juan Fernandez and Easter Isl. Vol. I

418 C. SKOTTSBERG

Myriapoda (274).

Pachymerium ferrugineum Latz. Wide-spread in Eur., undoubtedly introduced,
probably from Chile.

Orthomorpha gracilis Koch, Latz. Apparently wide-spread; introduced.

A third species, belonging to Lamyctes, could not be named.

Collembola (276).

Entomobrya multifaciata (Tullb.). N. and S. Amer., Eur., N. As., N. Zeal,
J. Fern. Introduced.

Embioptera (222).

Oligotoma Vosseleri (Krauss). Ceylon, Sumatra, Java.

Insecta.
Odonata (225).
Pantala flavescens Fabr. Amer., Afr., As., Austral.

Orthoptera-Dermaptera (225, 207).

I am indebted to Dr. PRINCIS for information on the nomenclature and dis-
tribution.

Antsolabis Bormanst Scudd. Galap. Is., Masatierra.

Onychostylus notulatus (Stal; Allacta, 225.297). Formosa, Malays., N. Guin.,
N. Caled., Samoa, Tahiti, Marquesas, Hawaii. Introduced from Tahiti?

Periplaneta Australasiae (Fabr.). Probably originally African, now cosmop.;
the genus Afr.—Orient.

Diploptera punctata (Eschtz, D. dytiscoides, 225.297). Ind. Ceylon, Burma,
Malays., Austral., Samoa, Marquesas, Hawaii; the genus Oriental. Accidentally
introduced?

Melanozosteria philpotti (Shaw). N. Zeal. An Australian genus, represented on
some Pacific islands.

Blatella vaga Heb. N. Amer.; Asia? The genus probably Oriental. Probably

adventitious.

Orthoptera-Saltatoria (5°).

Gryllus oceanicus Le Guillou. Malaysia and Japan to Polyn. CHOPARD regards
its presence on Easter Island as a proof of the facility with which certain insects
are transported large distances; nothing will prevent animals, he says, to be carried

across the Pacific from Australia. In this case, however, I guess that man has
been the agent.

Thysanoptera (3).

Hlaplothrips usttatus Bagn. var. inermis Ahlb. The typical species in Hawaii.

DERIVATION OF THE FLORA AND FAUNA 419

Neuroptera (92).

Chrysopa lanata Banks. Wide-spread in S. Amer. and also found on Hawaii.
A wide-ranging genus of several hundred sp.
+ Chr. Skottsbergi sben-Peters.

Lepidoptera (73).

Agrotis ypsilon Rott. Cosmop.

Cirphis Loreyt Dup. Widely distributed.

Achaea melicerta Drury. On all islands in the Pacific and Indian oceans.

Phytometra chalcytes Esp. Eur., As., most islands of the Pacific and Indian
oceans.

It is not probable that any of these Noctuidae are indigenous.

Diptera (57).

Sarconesia chlorogaster (Wied.). Chile, J. Fern. Introduced.

++Lipsana insulae-paschalis Enderl.

Leptocera (Coprophila) ferruginata (Stenh.) var. zzsulae-paschalis Enderl. The
typical species, known from Eur., Ind. and S. Amer., lives in horse-dung and
was spread with the horse. The variety was picked from the carcass of a sheep
and has, ENDERLEIN remarks (l.c. 679), perhaps developed after the arrival in
Easter Island where, however, there are many horses.

Coleoptera.
Curculionidae (12).

Aramigus Fuller? Horn. A noxious beetle, probably of N. Amer. origin, in-
troduced into many countries and on some isolated islands.

+ Pentarthrum paschale Auriv.

Areocerus fasciculates Deg. Cosmop., introduced.

Dytiscidae (299).

+ Bidessus Skottsbergi Zimmerm. The occurrence of an endemic aquatic beetle
in the crater lake of Rano Kao among the endemic hygrophilous mosses is of
interest.

Elateridae (308).

Simodactylus Delfint Fleut. Chile? Austral., N. Guin., N. Brit., Solomon Is.,

Hawaii.
Staphylinidae (25).
Philonthus longicornis Steph. Cosmop.

Hymenoptera.
Formicidae (283).
Ponera trigona Mayr. var. opacior Forel. N. Amer., W. Ind., Chile. The
typical species in Brazil, a subspecies in Austral.
Cardtocondyla nuda Mayr. subsp. mznuta Forel. Hawaii. The typical species
Ind., Ceylon, Austral., N. Guin. One subsp. is Mediterranean.

27* — 557857

420 C. SKOTTSBERG

Tetramorium guineense Fabr. Probably of African origin, now pantrop.; in
hothouses in the temp. region.

T. simillimum ¥F. Smith. As the former.

Plagiolepis mactavishi Wheeler. Formosa, Hawaii, Society Is.

Prenolepis bourbonica Forel subsp. Skottsbergi Wheeler. The typical species
known from Chagos, Nicobar and Seychelle Is., and E. Afr., Pemba I. Other
subspecies in Ind., Comoro Is., E. As., Philipp. Is. and Hawaii.

Ants are easily carried about by man, but it seems likely that Easter Island
also has indigenous forms.

Vespidae (206).
Polistes hebraeus F. E. Afr., Madag., Ind., China; Tahiti?

Hemiptera (27).

Clerada apicicornis Sign. Wide-spread, introduced.
Reduviolus capsiformts Germ. As the former.

Mollusca (759).

Limax arborum Bruch-Chant. Cosmop., introduced.

Milax gagates Drap. As the former.

Melampus philippi Kuester. Pert.

+M. pascus Odhner. The genus distributed over the Pacific from Hawaii
towN. Caled. ‘S. Amer:

Tornatellinops impressa Mouss. (Syn. Pacificella variabilis Odhner l.c.). Dis-
tributed from Fiji to Easter Island. Perhaps introduced with living plant material.

The fauna, as known hitherto, presents the same picture of extreme poverty
as the flora, and even if future researches will double the number of species and
reveal the occurrence of groups not yet recorded, a considerable portion will con-
sist of late immigrants. The known endemics are very few and one or two of
them questionable, and our experience from the old list of Juan Fernandez Diptera
(84) bodes no good for the single endemic genus. Altogether half a dozen en-
demic species and some endemic forms of lower category have been described,
and of the species found elsewhere some are, perhaps, indigenous. An example
of remarkable discontinuous distribution is offered by MWelanozosteria philpotia,
New Zealand and Easter Island; possibly it will be discovered in intermediate
stations, but such stations are difficult to find in other cases where Easter Island
is the terminus: Axzsolabus Bormanst, Galapagos Islands + Juan Fernandez, Chry-
sopa lanata, South America and Hawaii, Ponera trigona, America, and Haplothrips
notatus, Hawaii. A direct overseas transport is not very probable, and I cannot
tell if these animals are likely to have been introduced with the traffic.

DERIVATION OF THE FLORA AND FAUNA 421

Chapter XVI.
The biogeographical history of Easter Island.

The composition of the present fauna and flora does not help us to throw
any light on the earlier history of Easter Island, and we do not know what they
were like before the arrival of aboriginal man many centuries ago. We know that
the island became densely populated, that the natural resources, evidently poor,
were exploited, the soil cultivated wherever this was possible and a number of
useful plants introduced from other parts of Polynesia; tradition tells that the first
colonists arrived from Rapa, but other opinions have also been expressed. ROGGE-
VEEN, the discoverer of the island in 1722, did not bring a naturalist, but to
judge by his narrative the island must have looked much the same as when
SPARRMAN and the FORSTERS, who came with COOK in 1774, made the first bio-
logical observations. FORSTER collected and cited a few species (776) and men-
tions, in his narrative (347), “Mimosa” (Sophora toromiro) and Apium, which he
had observed before in New Zealand. If there had been other indigenous trees,
they had disappeared; Broussonetia papyrifera, Thespesia populnea and very likely
also 7riumfetta semttriloba had been introduced but were scarce. For wood the
natives depended on Sofhora, and most of this was gone already. FORSTER
found the place very barren, but on p. 578 he speaks of a hillock covered with
toromiro, and later on another similar hill is mentioned (p. 592), but all the trees
were low, not over 9 or Io feet, the main trunk of the biggest as thick as a
man’s thigh. No wonder that the single canoe seen was a patchwork of pieces 2
or 3 feet long, and so was the paddle. The population did not exceed 700. When
THOMPSON and COOKE (343) visited the island in 1886, groups of trees were ob-
served in some places:

In other parts of the island may be seen, in places in considerable numbers, a
hardwood tree, more properly bush or brush, called by the natives toromiro. These
must have flourished well at one time, but are now all, or nearly all, dead and decaying
by reason of being stripped of their bark by the flocks of sheep which roam at will
all over the island. None of the trees are, perhaps, over 10 feet in height, nor their
trunks more than 2 or 3 inches in diameter (p. 705).

The last specimens of toromiro are restricted to the inside of the crater Rano
Kao. Easter Island was made a national park in order to protect the unique stone
monuments, and is a bird sanctuary, but otherwise nature is not preserved but the
land grazed over without restriction as far as I am aware.

Among the many isolated islands of the Pacific, Easter occupies a rather
unique position. Oceanic islands belong to two main categories, high volcanic and
low coralline; only the former are of greater biological interest and possess the
standard set of “‘peculiarities’’ described by HOOKER, WALLACE and others. Easter
Island seems to form a type by itself. It is volcanic and cannot be called low, for
the highest mountain is 530 m high and some of the craters reach an altitude

422 C. SKOTTSBERG

of 300-400 m, sufficient, one would think, to create a humid montane belt with
fairly luxuriant arboreous vegetation, but of this there is nothing, in any case
nothing left. Rains are frequent, and the amount of precipitation is not small,
but evaporation, favoured by high temperatures and the strong S.E. trade wind,
is great and most of the water rapidly disappears underground. The climax vege-
tation is an oceanic steppe-like meadow or grass heath, as some would prefer
to call it (372). The flora does not present many of the characteristics of oceanic
islands. There are no endemic genera, no peculiar endemic species, no prepon-
derance of woody plants; Sophora is the only tree and Lyczuwm the only shrub;
the flora is herbaceous, comprising few therophytes but many annual weeds. The
ratio, species : genus is 1.2: 1. With the exception of Gramineae, which dominate,
and Cyperaceae (some of these perhaps not indigenous) most large and world-wide
families are absent, even Compositae; there are no conifers, no orchids but a fair
number of ferns: in these respects the island conforms to typical oceanic islands.

On the other hand, Easter Island has little in common with the low islands,
atolls or other coralline structures with no rock foundation exposed, where en-
demics are, as a rule, absent and the fragmentary flora consists of species easily
transported by the natural agencies and by man.

The geographical position is unfavourable to immigration, the chances for
the arrival of seeds from America small, the distance being 3700 km, and 1850
km separate Easter from Pitcairn, the nearest basaltic islet. On the other hand,
the chances for establishment ought to have been good: new surroundings, though
perhaps not very varied, plenty of space, no competition, conditions furthering
the evolution of new species and genera as many biogeographers believe, but
nothing like that seems to have happened on Easter Island. The objection will
be raised that we do not know, for the bulk of the original fauna and flora may
have been destroyed by man and will remain unknown—but is it not surprising,
if this be true, that no peculiar, systematically isolated form was preserved to
our days? After all, perhaps not. Truly, in many high and well populated Poly-
nesian islands endemics are plentiful, growing on the elevated ridges, on the
precipitous mountain sides, in the deep recesses of the gorges where cascades
tumble down or even in less inaccessible, but uninhabited places, but the topo-
graphy of Easter Island is different and I can see no reason why not man and
his animals could have succeeded to exterminate practically everything of the
original nature except the lichens and mosses covering the rocks and a few herbs
and grasses.

Distance is not the only factor, time is another; the island is, somebody
will say, perhaps too young. It has a youthful appearance, the craters are well
preserved, but they are secondary and not responsible for the origin of the island,
and their well-preserved shape is no proof of youth. There is no sign of recent
activity——FORSTER’s idea that the decay during the 18th century of the old ab-
original culture was due to some volcanic catastrophe lacks foundation. As there
are no permanent streams, erosion must be slight. The various vazos and other

cones may be old enough and the foundation, on which they stand, very ancient.
As BRUGGEN says (33z. 290):

DERIVATION OF THE FLORA AND FAUNA 423

A pesar de que los volcanes ... tienen sus formas muy bien conservadas, no
existe en la tradicion de los nativos ningun recuerdo de una erupcién. La isla pro-
duce la impresion de tratarse del resto de un segmento de un enorme volcan central,
en cuya superficie inclinada se han formado los crateres actualmente visibles como
conos adventicios superiores.

Easter Island is small, but not very small, 117 sq. km, and GULICK (zz9)
called it “the wave-worn remnant of an island that could once have claimed about
twice that area’. Lack of time cannot have prevented the island to reach matu-
rity and to acquire some internal harmony of the flora and fauna. Still, they are
very disharmonious. The general situation of isolated islands is clearly set forth
by GULICK (l.c. 413-414):

It is evident that mature groups of islands will attain an internal harmony from
the standpoint of the systematist. But this harmony, instead of reflecting the pre-existent
harmony of some continental source (as is the case for continental islands or land-bridge
remnants), will be recognizably derivable by descent from a quite limited number of
original importations, at the start distinctly miscellaneous and “‘disharmonic’’, as was
observed to be the condition in Bermuda and St. Helena. Large series of related or
previously associated forms will be found from the beginning in continental islands,
but their counterpart must be brought into existence de novo if the group is truly
oceanic. But this distinction, obvious in theory, is in practice very difficult to recog-
nize, unless the oceanic condition is really extreme.

Ikaster Island complies with this condition and is often described as an ex-
ample of a truly oceanic island; this is the general opinion, but it was told above
(Chapter IV) that there is no lack of theories according to which Easter is a rem-
nant of a land mass of continental size, a mid-Pacific continent or a land bridge
uniting the Australian—New Zealandic area with South America and Hawaii; ARLDT,
GERMAIN, GUILLEMIN, J. W. GREGORY, MEYRICK, PILSBRY etc. were quoted. It
is not improbable that the island is the rest of a somewhat larger piece of land,
but this is all we can say. The bathymetrical conditions—see map—hardly tempt
us to construct bridges, even if not all signs of submergence are lacking. The
3000 m curve forms a large, almost closed crescent, on the north extremity of
which Easter Island rises. E.N.E. is Sala y Gomez, extending S.W.-N.E., 1200 m
long, 150 m broad and 30 m high, and this tiny islet is the only visible part of a
larger reef running in the direction N.E. 1/, N. and called Scott Reef, where the
smallest depth, 1950 m from the islet, is 35 m only; between this place and the
islet a series of soundings gave 55, 60, 49 and 46 m. The bank extends at least
a couple of km west of the islet with depths of 42 to 68 m, and nowhere within
this range a greater depth than 95 m was found (772. 74). The scale of my map is
too small to give these details. Proceeding east we find, in about the same dis-
tance from Sala y Gomez as this is from Easter Island another submarine ridge run-
ning N.-S., bounded by the 1000 m curve and with depths as modest as 862 and
308 m. Further east again, in 97 30’ w. |., approximately, is another ridge trending
W.-E., where the smallest figure is 497 m. None of these shallow areas were, I
believe, known to the bridge-builders. They must be welcome also to those who
look, if not for sunken continents, at least for submerged islands used as way
stations in the migrations. However, all this means a possible extension east,

424 C. SKOTTSBERG

towards South America but this is not where we ought to look for the vanished
world that could have inhabited a greater and higher Easter Island; rather, we
have better look in the opposite direction.

It is FORREST BROWN’s merit to have pointed to the large insular world
known as Southeastern Polynesia as as important floristic—and, I presume, faun-
istic—centre, and from his Flora (35) and the reports published by members of
other recent survey parties, sponsored by the Bishop Museum in Honolulu, the
following data were compiled. The Society Islands are excluded and only the
angiosperms considered. Included are the Marquesas, Tuamotu and Austral or
Tubuai Islands, and the more isolated Rapa (Oparu), Mangareva (Gambier), Pitcairn
and Henderson (Elizabeth) Island.

The Marquesas Islands are high, 800-1200 m, and belong, to judge from the
geology and topography, to the same generation of Tertiary islands as Tahiti,
Juan Fernandez, Macaronesia, etc. The Tuamotus proper are atolls. Of the Austral
Islands the high basaltic are considered, Rimatara, 95 m, Rurutu, 410 m, Tubuai,
400 m, and Raivavae (Vavitao), 440 m; further, the following outlying islands are
included: Rapa (Oparo), 640 m, Mangareva (Gambier), 400 m, Pitcairn, 350 m>
and Henderson (Elizabeth), an islet of raised coral said to be only 25 m high
(345) but nevertheless the home of an endemic Saztalum (compare Laysan of the
Leeward Hawaiian Islands with S. e//zpdicum var. laysanense). The Marquesas flora
is considered to be well known and the same may be true of the flora of the
other islands, even if no complete lists have been published; I suppose that all
the novelties have been described, but my figures for wide-spread species are,
perhaps, too low. There is a difference between my figures and those given by
BROWN, because varieties are counted by him as units equal to species, which
explains why his figures for the endemics are so high.

The largest families are Rubiaceae (36), Cyperaceae (25), Compositae (19),
Euphorbiaceae (16), Gramineae (13), Leguminosae (12), and Piperaceae (10). Other
large and important families, such as Araliaceae, Cruciferae, Ericaceae, Malvaceae,
Myrtaceae, Orchidaceae, Sapindaceae, etc., are represented by fewer species. We
have every reason to believe that the flora has suffered losses after man had
taken possession of the soil.

The total number of presumably indigenous species—many of aboriginal
introduction and not few later arrivals have become naturalized—is 282, of which
156 are endemic within the area. The genera are 145, of which only 3 are endemic
according to BROWN. The ratio species: genus is almost 2:1. No genus is very
large, the largest is Psychotria with 12 species, and 10 have from 5 to 10 species
each. The distribution of the species and the number of local endemics are indi-
cated in Table VIII. The figures do not pretend to be exact.

A large proportion of endemics and of woody, arboreous or fruticose species
—suffruticose excluded—are characteristic of oceanic floras of considerable anti-
quity. Of the 156 endemic species 113 (72.4%) are woody, of the 126 found
elsewhere 69 (54.8%). The herbaceous species are, with very few exceptions, peren-
nial. Systematically isolated types are few, and even the Marquesas Islands can-
not, in this respect, be compared with either Hawaii or Juan Fernandez.

DERIVATION OF THE FLORA AND FAUNA 425

Table VIII.

Distribution of angtosperms in Southeastern Polynesia.

Number of Number of % Woody species

species endemics endemics Number %
Marquesas <)<=- .- <i. I51 79 52.3 100 66.2
AOC at ous ion = 44 7.0 23 52.3
MUSLIM Prat vacate" 5° 5) 0 12 80 12 15.0 53 66.7
pe ere SS 89 44 49.4 62 69.6
Manpareval. =is2iftt Ss 28 3 10.7 17 60.9
EATLCAIERA <) accucehere © | 26 2 Th | 19 73.1
Henderson. .... . 21 3 14.3 15 Tha:
LCT | a ee a peo eee 282 1560 55-3 182 64.5

BROWN (35) regards southeastern Polynesia, with the Tuamotus in the centre,
as an old, submerged region:

Affinities point to the Tuamotuan region as one of the ancient mid-oceanic centers
of origin for a large part of the dicotyledonous flora of southeastern Polynesia (III. 6);

and, speaking of the distribution of F7zfchia, he writes (I.c. 364):

The grouping and affinity of these allied species strongly suggest the Tuamotuan
region as the center of origin at a rather remote period, possibly at the dawn of the
Tertiary or somewhat earlier, when it may be assumed that high (pre-Tuamotuan) islands
existed in place of the low (Tuamotuan) atolls of the present.

Within the Marquesas archipelago the sea is shallow:

Apparently, an emergence of roo meters would cause land to appear in six places;
an additional emergence of 300 meters would unite or bring into close contact all
land areas of the archipelago.... Botanical evidences, outlined in an earlier paper,
indicate that the islands were at one time 1000 to 2000 meters higher than at pre-
Sentes ..(Lc, 1.17).

The floristic affinities of this region is with Malaysia—Melanesia—Australia;
there is no neotropical element in spite of the prevailing direction of winds and
currents. BROWN, who could be expected also to look toward America, remarks:

The Cichorieae, to which /7¢chia belongs, are best represented in Europe and
America, pointing to a more remote American center of origin for the pre-Tuamotuan
ancestral stock.

This brings up the Dezdroseris- Thamnoseris problem. The four dendroseroid
genera form a very natural group, 7hamnoseris of Desventuradas Islands stands
apart, and so does F7fchza. If they are, at least distantly, related, and isolated
from all other Cichoriaceous genera, then the possibility of an Antarcto-tertiary
ancestry should be considered. I have, however, referred the Dendroseris assem-
blage to an ancient neotropical element, absent from the present continental flora,

1 Proceed. 2d Pan-Pacif. Sci. Congr., Vol. 2, 1923.

426 C. SKOTTSBERG

and Zhamnoscris finds no better place, but to derive /ztchia from an American
source seems little inviting. Another solution is, perhaps, in sight. Professor GUN-
NAR ERDTMAN kindly told me that, to judge by the pollen morphology, F7tchia
mav have to be removed from the Czchorzum subfamily where J. D. HOOKER
placed it next to Dexdroseris and where it has remained.

There is in the Pacific Ocean no island of the size, geology and altitude of
Easter Island with such an extremely poor flora and with a subtropical climate
favourable for plant growth, but nor is there an island as isolated as this, and
the conclusion will be that poverty is a result of isolation—even if man is re-
sponsible for the disappearance of part of the flora, it cannot have been rich;
the Marquesas Is., which have been inhabited longer, I believe, and formerly
had a large native population, still preserve a fairly rich and varied angiospermic
flora, half of which is endemic. The distances are too great to be overcome
except on very rare occasions. The nearest land is to the west, the small most
easterly islets of the Mangareva (Gambier) group, but winds (S.E. trade-wind) and
currents are unfavourable for transport from W., and Easter Island appears to lie
away from the cyclonic tracks. Beach drift is responsible for the arrival of several
species, /pomaea, Caesalpinia, Chenopodium, Tetragonia, Erythraea, Apium, Samo-
lus, Lyctum and perhaps some grasses and species of Cyperus, altogether about
1/, of the angiosperms. Storms bring light diaspores, but it is noteworthy that
Compositae are absent. I can find no special adaptations for bird carriage, but the
possibility of rare cases of epizoic transport cannot be excluded. However that
may be, Easter is a good example of an island peopled by “‘waifs and strays”.

Affinities are, as we have seen, with Malaysia—Australia or pantropical, whereas
the well-marked east Polynesian flora has contributed nothing, not even its leading
family Rubiaceae, rich in drupe-fruited forms. Sophora torom7ro is allied to S. “‘tetra-
ptera’ of Raivavae and Rapa; I cannot tell if this is the true ¢etrapiera, a native
of New Zealand, but I do not think it is, and as BROWN’s description (III. 120)
shows, it differs much from forvomiro, which comes very close to S. masafuerana.
Neither is of American ancestry: sect. Edwardsza is austral-circumpolar and gene-
rally regarded to be of Antarctic origin or, at least, history.

With the exception of Lycium carolinianum var. sandvicense, supposed to
belong to the beach drift, there is, if Fztchza is definitely excluded, no American
element in the flora of southeastern Polynesia, nor is it expected there. It is, as
we have seen, found in Easter Island. Of the 3 endemic grasses, S/pa was ten-
tatively brought to the palaeotropical element, Axoxopus to the neotropical. Dax-
thonia is an austral-circumpolar, tricentric genus. Three American, not endemic
species, Cyperus eragrostis, Scirpus riparius and Polygonum acuminatum, remain
to be accounted for.

If Easter Island once had a richer flora is an open question. According to
newspaper reports a palynological survey of the swamp in the crater of Rano
Kao was planned for HEYERDAHL's recent survey. The thickness of the loose,
water-soaked Campylopus peat was not measured by me; it is a somewhat dan-
gerous quagmire which cannot be bored with the usual methods, but samples may

DERIVATION OF THE FLORA AND FAUNA 427

be dug out from different depths, and if pollen of species not growing on the
island are found, some light will be thrown on the history of the flora.

The map accompanying this paper was prepared by the Oceanographical
Institute in Goéteborg. I am greatly indebted to the Director, Professor HANS
PETTERSSON, and to Dr. BORJE KULLENBERG for valuable assistance.

May, 1956.

Additions.

Ee 252.

Usnea Gaudichaudii Motyka, known from the “espinal” of Central and North
Chile, is quoted for Juan Fernandez (Masatierra) by MOTYKA 344.600 as found
by BERTERO, 1830. As its occurrence there seems little probable—BERTERO
collected also near Valparaiso, etc.—I have excluded the species from my list.

To Chapters IV and XVI.

A recent paper by R. FURON, “Importance paléogéographique des mouve-
ments de subsidence du Pacific Central’ (Rev. gén. des Sciences 62, 1955), should
be noted here. His object is expressed in the following terms:

Constatant combien les biogéographes manquent de documentation géologique, il. nous a
paru utile de regrouper les notions acquises au cours de ces derniéres années, notions qui
éclairent fort bien l’historie du Pacifique depuis le Crétacé (p. 307).

Whereas the Galdpagos Islands show, he says, a purely oceanic type of
rocks, andesitic basalt and tuff are found on Easter Island and andesite and
trachyte on Pitcairn. Contrary to what was told above, corals of Cretaceous age
were dredged in a depth of 2000 m on one of the Central Pacific guyots. The
deep borings through atolls, the latest on Eniwetok in 1953, have penetrated
through coral formations dating from Pleistocene to Eocene to the bedrock of
basalt, indicating a subsidence since the end of the Cretaceous of 2000 m. Perhaps
other parts of the Pacific would give still greater figures.

To Chapter V.

In the Proceedings of the Cotterwood Natur. Field Club 31, 1955, T. A.
SPRAGUE gives an account of the Drift Theory of Du Toir and finds that, from
a botanist’s viewpoint, this theory “offers the best explanation hitherto brought
forward of the major problems of biogeography”. With reference to the physical
side of the drift process he quotes HOLMES’ “Principles of Physical Geography’
(1944). In the case of Juan Fernandez, which certainly is one of the mzxor pro-
blems, I cannot see that the drift theory offers an acceptable solution.

To Chapter VII, p. 360.

An important paper by H. MoLHOLM HANSEN, “‘Life forms as age indica-
tors’, Ringkjobing 1956, confirms the opinions of SINNOTT & BAILEy and others.

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1939, IV. Berkeley 1940.

. —— Genera Filicum. Annal. Crypt. et Phytopath. 5. Waltham, Mass. 1947.

. Cortés, R. Tachynidae (Diptera). (Insect. J. Fern. 9.) Rey. Chil. de Entomol. 2, 1952.
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nificance to phytogeography. New Phytologist 45, 1916.

. DarBISHIRE, O. V. The Lichens. Wiss. Ergebn. schwed. Siidpolar-Exp. 1901-1903,

IV: 1. Stockholm 1912.

. Doncr, C. W. Lichens and Lichen Parasites. Reports Brit. Austral. N. Zeal. Antarct.

Research Exp. 1929-31. Ser. B. VII. Adelaide 1948.

. Domin, K. Monographische Ubersicht der Gattung Centella L. Bot. Jahrb. fiir System.

etc. ed. by A. ENGLER, 41, 1908.

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moose. Mém. Soc. Roy. de Sciences de Bohéme 1923. Prague 1924.

. Du Rirtz, G. E. Two new species of Euphrasia from the Philippines and their phyto-

geographical significance. Svensk bot. tidskr. 25, 1931.

. ——— Problems of bipolar plant distribution. Acta Phytogeogr. Suec. XIII, 1940.
. Dusen, P. Uber die tertiire Flora der Magellanslinder. Wiss. Ergebn. schwed. Exp.

1895-1897, I, 1907.

. —— Uber die tertiéire Flora der Seymour-Insel. Wiss. Ergebn. schwed. Sitidpolar-

Exp: 1901-1903, IM ro re.

. Du Torr, L. Our Wandering Continents. Edinburgh and London 1937.
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. Emerson, E. A. A new Termite from the Juan Fernandez Islands. Nat. Hist. Juan

Hern Lieeng24:

. Enperzein, G. Die Dipterenfauna der Juan Fernandez-Inseln und der Oster-Insel.

Ibid. 1938.

. Encier, A. Versuch einer Entwickelungsgeschichte der Pflanzenwelt seit der Tertiar-

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. ENGLERT, S. La Tierra de Hotu Matua. Valparaiso 1948.

. Epxinc, C. Synopsis of the South American Labiatae. Fedde, Repert. Beih. 85, 1935.
aaa lhe Labiataevof Chile,“Rey.- Universitaniar2 2) mgs 77,

. —-— The distribution of the American Labiatae. Proceed. 6th Pacif. Sci. Congr.

1939, LV. Berkeley 1940.

. —-— Distribucion geogrdfica y parentesco de las Labiadas de la América del Sur.

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REQ;
120.
Roi’.

122.

DERIVATION OF THE FLORA AND FAUNA 431

. Ere, A. Eine Reise nach der Robinson-Insel. Hamburg 1889.
. Espen-PETERSEN, P. More Neuroptera from Juan Fernandez and Easter Island. Nat.

Hist. Juan Fern. III, 1924.

. Evans, A. W. The Thallose Hepaticae of the Juan Fernandez Islands. Ibid. II, 1930.
. Fassert, N. C. Callitriche in the New World. Rhodora 53, 1951.
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significance. K. Sv. Vet.-akad. Handl. ser. 3, 19, 1940.

. Fruxe, C. L. Syrphidae. (Insect. J. Fern. 18.) Rev. Chil. de Entomol. 4, 1955.
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géogr. 29 année, no. 256, 1952.

. —— The American element in the Hawaiian Flora. Proceed. 7th Intern. Botan.

Congr. 1950. Stockholm 1953.

. Freeman, P. Mycetophilidae, Sciaridae, Cecidomyidae and Scatopsidae. (Insect. J.

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. Fries, R. E. Die Myxomyceten der Juan Fernandez-Inseln. Nat. Hist. Juan Fern. II,

1920.

. Fries, TH. yk. Die Gasteromyceten der Juan Fernandez- und Osterinseln. Ibid. 1922.
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7th Intern. Botan. Congr. 1950. Stockholm 1953.

. Gepren, H. Coleoptera-Tenebrionidae von Juan Fernandez. Nat. Hist. Juan Fern. III,

Ig21.

. Germaln, L. L’origine et l’évolution de la faune de Hawaii. Proceed. 3d Pacif. Sci.

Congr. 1926. Tokyo 1928.

. Gipgs, L. Notes on Phytogeography and Flora of the mountain summit plateau of

Tasmania. Journ. of Ecology 8, 1920.

. Gitiocty, L. R. Coleoptera-Nitidulidae. (Insect. J. Fern. 24.) Rev. Chil. de Entomol.

4, 1955:

. Goon, R. See Continental Drift.
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1951. Buenos Aires.

. GoopspeED, T. H. Plant Hunters in the Andes. New York 1941.
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New Zeal. Ass. Advancem. Sci. 27, 1949.

. Grant, ADELE L. A monograph of the genus Mimulus. Ann. Miss. Botan. Gard. XI,

1924.

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Tokyo 1928.

. Grecory, J. W. Theories of the origin of the Pacific. Proceed. Geol. Soc. London

86, 1930.

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1952.

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Gutick, A. Biological peculiarities of oceanic islands. Quart. Review Biol. 7, 1932.

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Guppy, H. B. Observations of a naturalist in the Pacific between 1896 and 1899. II.
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—w— The island and the continent. Journ. of Ecol. 7, 1919.

C. SKOTTSBERG

_ GuTENBERG, B. Geophysical and geological observations in the Pacific area and

tectonic hypotheses. Proceed. 7th Pacif. Sci. Congr. 1949, II. Wellington 1953.

24. GorrscH, W. Die Robinson-Insel Juan Fernandez und ihre biogeographischen Pro-

12
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125. HANDSCHIN, C. Neuroptera. (Insect. J. Fern. 15.) Rev. Chil. de Entomol. 4, 1955.

126. Harmston, F. C. Dolichopodidae. (Insect. J. Fern. 17.) Ibid. i

127. Hems.ey, W. B. (a) Report on the Present State of Knowledge of Various Insular
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Ser. Res» Ha M?.S.Challengers Bot. 18 35

128. Hennic, W. Phryneidae, Helomyzidae, Lonchaeidae, Piophilidae, Anthomyzidae,
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129. Herzoc, Tu. Geographie der Moose. Jena 1926.

30. —-— Die foliosen Lebermoose der Juan Fernandez-Inseln und der Oster-Insel. Nat.
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131. —-— Zur Bryophytenflora Chiles. Revue Bryol. et Lichenol. XXIII, 1954.

132. Hess, H. H. Drowned ancient islands of the Pacific. Amer. Journ. Sci. 244, 1946.

133. Hicken, C. M. Polypodiacearum argentinarum catalogus. Rev. Mus. de La Plata 15,
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134. Hit, A. W. Antarctica and problems of geographical distribution. Proceed. 5th Intern.
Bot. Congr. 1926. Ithaca 1929.

135. Hinton, H. E. See Continental Drift.

136. Hopcson, E. AMy & Sainspury, G. O. K. Bryophytes collected by G. E. Du Rietz
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137. HotrepauL, O. On the geology and physiography of some Antarctic and Sub-Ant-
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138. Hooker, J. D. Lecture on Insular Floras before the British Association at Notting-
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139. Hur, A.-M. Lichens. Deuxiéme Expéd. Antarct. Frangaise 1908-1910. Paris 1915.

140. Hutcuinson, J. The Families of Flowering Plants. I. 1926. II. 1934.

141. Hux ey, J. Species formation and geographical isolation. Proceed. Linnean Soc.
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142. Insecros. Los Insectos de las islas Juan Fernandez. Ed. by Dr. G. Kuschel. 1-12.
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143. IRMSCHER, EF. Pflanzenverbreitung und Entwickelung der Kontinente. I-II. Mitteil.
Hamburg. Inst. allgem. Botan. 5, 1922; 8, 1929.

144. JEFFREY, E. C. & Torrey, R. S. Transitional Herbaceous Dicotyledons. Annals of
Botany 35, 1921.

145. JEFFREYS, H. See Continental Drift.

146. JHERING, H. Das neotropische Florengebiet und seine Geschichte. Bot. Jahrb. fiir
System. etc. ed. by A. ENGLER. 17, 1893, Beibl. 42.

147. JoHANsson, L. Ein neuer Landblutegel aus den Juan Fernandez-Inseln. Nat. Hist.
Juan Fern. III, 1924.

148. Jounston, I. M. A revision of the South American Boraginoideae. Contrib. Gray
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149. —-— Papers on the Flora of Northern Chile. 1. The coastal Flora of the Depart-
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150. JoHow, F. Estudios sobre la Flora de las Islas de Juan Fernandez. Santiago 1896.

151. JorDAN, D. S. The origin of species through isolation. Science 22, 1905.

152. —-— Isolation with segregation as a factor in organic evolution. Annual Rep. Smith-
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153. JorpaNn, K. Coleoptera-Anthribidae from Juan Fernandez. Nat. Hist. Juan Fern. III,
EQGok.

154. Joyce, J. R. T. See Continental Drift.

Fess
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157:
158.
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HOW

162,
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DERIVATION OF THE FLORA AND FAUNA 433

Just, Tu. Geology and Plant Distribution. Ecol. Monogr. 17, 1947.

Kauset, E. Contribucioén al Estudio de las Mirtaceas Chilenas I-II. Rev. Argent. de
Agron. 9, 1942.

——- Notas Mirtoldgicas. Lilloa 13, 1947.

—— Revision del género Escallonia. Darwiniana 10, 1953.

KrtssLer, K. Ascomyceten, Fungi imperfecti und Uredineen von Juan Fernandez.
Nat. Hist. Juan Fern. II, 1928.

—— Nachtrag zur Pilzflora von Juan Fernandez. Ibid.

Kormitey, N. A. Lygaeidae (Hemiptera). (Insect. J. Fern. 1.) Rev. Chil. de Entomol.
25 OSes

Krause, K. Uber einen hapaxanthen Baum. Mitteil. deutsch. dendrol. Ges. 1921.

KraAuseEL, R. Pflanzenwanderungen im Tertidr. Proceed. 7th Intern. Botan. Congr.
1950. Stockholm 1953.

KUKENTHAL, G. Vorarbeiten zu einer Monographie der Rhynchosporoideae. VIII.
Fedde, Repert. 48, 1940. — XII. Ibid. 51, ro42.

Lawrence, G. H. M. Taxonomy of Vascular Plants. New York 1951.

Lesng, P. Coleoptera-Bostrychidae et Cioidae de Juan Fernandez. Nat. Hist. Juan
Ber: Tilts119.213:.

Linpsay, A. Biology and biogeography of the Antarctic and Subantarctic Pacific.
Proceed. 6th Pacif. Sci. Congr. 1939, II. Berkeley 1940.

Looser, G. Nueva localidad cerca de Antofagasta del Helecho Polypodium Masa-
fuerae Philippi. Rev. Universit. 28. Santiago 1943.

—— Los Blechnum (Filices) de Chile. Ibid. 32, 1947.

—— Comparacion de las Bromeliaceas de Chile con las de Argentina y Peru. Rev.
Chil. de Hist. y Geogr. n. 110 (1947), 1948.

—— El género Polypodium L. y sus representantes en Chile. Rev. Universit. 36.
Santiago 1951.

Lopez, J. E. Esploracion de las Islas esporddicas al occidente de la costa de Chile.
Anuario Hidrograf. de la Marina de Chile. Il. 1876.

Lourteic, Aticia. Ranunculaceas de Sudamerica templada. Darwiniana 9, 1952.

LONNBERG, E. The Birds of the Juan Fernandez Islands. Nat. Hist. Juan Fern. III,
1920.

—— Notes on Birds from Easter Island. Ibid. rg2r.

Mackenzig Lams, I. A monograph of the Lichen genus Placopsis Nyl. Lilloa 13,
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Masses, G. Lichenes. The Subantarctic Islands of New Zealand. Wellington 1909.

MatTFeELpD, J. Compositae. Nova Guinea 14: 4. Leiden 1932.

Mayr, E. The origin and history of the Bird Fauna of Polynesia. Proceed. 6th Pacif.
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Merravux, A. Ethnology of Easter Island. Bull. Bishop Mus. Honolulu 1940.

MricHaeLsEn, W. Oligochaeten von Juan Fernandez und der Oster-Insel. Nat. Hist.
Juan Fern. Ill, 1921.

Montane, C. Liquenes, in Gay, Hist. fis. y polit. de Chile. Botanica 8. Paris 1852.

Mumrorp, E. P. The present status of studies of faunal distribution with reference
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Mutter, J. Lichens. Miss. Scient. du Cap Horn 1882-83. V. Paris 1889.

NavAs, L. Neuroptéres des Iles Juan Fernandez et de l’Ile de Paques. Nat. Hist.
Juan Fern. III, 1921.

Nitsson, H. Syntetische Artbildung. Lund 1953.

Nixon, G. E. J. Hymenoptera-Braconidae. (Insect. J. Fern. 26.) Rev. Chil. de Ento-
mol. 4, 1955.

Nyseuin, O. Nesophilaemon n.g. fiir Philaemon skottsbergi L. Johansson. Zool.
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434 C. SKOTTSBERG
189. OpHNER, N. Hy. Mollusca from Juan Fernandez and Faster Island. Nat. Hist. Juan
Fern. III, 1922. Addenda, ibid. 1926.
1g0. OcLosin, A. Mymaridae (Hymenoptera). (Insect. J. Fern. 12.) Rev. Chil. de Entomol.
2; 9952.
191. —— Bethylidae y Dryinidae (Hymenoptera). (Insect. J. Fern. 14.) Ibid. 3, 1953.
192. OLiveR, W. R. B. The genus Coprosma. Bull. Bishop Mus. Honolulu 132, 1935.
193. Perkins, R. C. L. Introduction, being a review of the land-fauna of Hawaiia.
Fauna Hawaiiensis I. Cambridge 1913.
194. PreirFER, H. Oreobolus R. Br., eine merkwiirdige Cyperaceengattung. Fedde, Repert.
2a LOZ yc
195. Pic, M. Coleoptera-Anobiida de Juan Fernandez. Nat. Hist. Juan Fern. HI, 1924.
196. —-— Coleoptera-Clavicornia et autres de Juan Fernandez. Ibid.
197. Pircer, R. Uber einige Gramineae der Skottsbergschen Sammlung von Juan Fer-
nandez. Fedde, Repert. 16, 1920.
198. —— 7}. Das System der Gramineae unter Ausschluss der Bambusoideae. Bot.
Jahrb. ftir System. etc. ed. by A. ENGLER, 76, 1954.
199. Prats, L. Zur Kenntnis der Insel Juan Fernandez. Verhandl. Ges. fiir Erdkunde
3erlin XXIII, 1896.
200. Pope, R. D. Coleoptera-Colydiidae. (Insect. J. Fern. 25.) Rev. Chil. de Entomol. 4,
1955:
201. Princis, K. Uber einige neue bzw. wenig bekannte Blattarien. Arkiv f. Zool. Ar igeee
202. Rampo, B. Historia da flora do planalto riograndense. Anais Bot. Barbosa Rodriguez
V, 1953-
203. ReicHe, K. Grundziige der Pflanzenverbreitung in Chile. ENGLER & DRube, Vege-
tation der Erde 7. Leipzig 1907.
204. RicHarbs, O. W. Sphaeroceridae. (Insect. J. Fern. 21.) Rev. Chil. de Entomol. 4,
1955-
205. Riptey, H. N. The Dispersal of Plants throughout the World. London 3930.
206. Roman, A. Ichneumoniden von Juan Fernandez. Nat. Hist. Juan Fern. II, 1924.
207. RomeELL, L. Bacidiomycetes from Juan Fernandez. Ibid. II, 1928.
208. Saprosky, C. W. A new species of Ogcodes from the Juan Fernandez islands. Rev.
Chil. de Entomol. 1, 1951.
209. —-— Chloropidae. (Insect. J. Fern. 19.) Ibid. 4, 1955.
210. Sarnspury, G. O. K. A Handbook of the New Zealand Mosses. Bull. Roy. Soc.
of N. Zeal. no. 5, 1955.
211. SANTIS, L. pe. Hymenoptera-Eulophidae, Entedontidae (Chalcidoidea 1). (Insect. J.
Fern. 27.) Rey. Chil. de Entomol. 4, 1955.
212. SATCHELL, G. H. Psychodidae (Diptera). (Insect. J. Fern. 8.) Rev. Chil. de Entomol.
2, E52.
213. SCHENKLING, S. Coleoptera-Cleridae von Juan Fernandez. Nat. Hist. Juan Fern. IU,
1931.
214. ScHMID, F. Trichoptera. (Insect. J. Fern. 4.) Rev. Chil. de Entomol. 2, 1952.
215. Scumipt, A. Coleoptera-Scarabaeidae von Juan Fernandez. Nat. Hist. Juan Fern. Ill,
1931.
216. Scuorr, H. Collembola aus den Juan Fernandez-Inseln und der Osterinsel. Nat.
Hist. Juan Fern. III, rg21.
217. SETCHELL, W. A. Les migrations des oiseaux et la dissémination des plantes. Compte-
rendu sommaire Soc. de Biogéogr. 22, 1926.
218. —— Migration and endemism with reference to Pacific Insular floras. Proceed. 3d
Pacif. Sci. Congr. 1926. Tokyo 1928.
219. —— Pacific Insular floras and Pacific paleogeography. Amer. Naturalist 69, 1935.
220. Sewarp, A. C. A study in contrasts. The present and past distribution of certain

ferns. Linn. Soc. Journ. of Bot. 46, 1922.

DERIVATION OF THE FLORA AND FAUNA 435

. SewarD, A. C. & Conway, VERONA. A phytogeographical problem: fossil plants from

the Kerguelen Archipelago. Ann. of Bot. 48, 1934.

. Sttvestri, F. Thysanura and Embioptera. Nat. Hist. Juan Fern. III, 1924.
. Simpson, G. G. Antarctica as a faunal migration route. Proceed. 6th Pacif. Sci.

Congr. 1939. Berkeley 1940.

. Sinnott, E. W. & Battey, I. W. The origin and dispersal of Herbaceous Angio-

sperms. Ann. of Bot. 28, 1914.

. Sjostrept, Y. Odonata. Orthoptera. Nat. Hist. Juan Fern. III, 1924.
. SKorrsBeRG, C. Die Gefisspflanzen Siidgeorgiens. Wiss. Ergebn. Schwed. Siidpolar-

Exp. IV. Stockholm 1905.

. —— Studien iiber die Vegetation der Juan Fernandez-Inseln. K. Sy. Vet.-akad.

Handl. 51, 1914.

. —— Notes on the relations between the floras of Subantarctic America and New

Zealand. Plant World 18, 1915.

. —— The Phanerogams of the Juan Fernandez Islands. Nat. Hist. Juan Fern. II,

1922.

. —— The Phanerogams of Easter Island. Ibid.
. —-- Juan Fernandez and Hawaii. A phytogeographical discussion. Bull. Bishop

Mus. 16. Honolulu 1925.

. —— Einige Bemerkungen iiber die alpinen Gefasspflanzen von Masafuera. Ver6ff.

Geobot. Inst. Riibel 3. Ziirich 1925.

. —-— Einige Pflanzen von der Oster-Insel. Acta Horti Gotob. 3, 1927.
. —— Remarks on the relative independency of Pacific Floras. Proceed. 3d Pacif.

Sci. Congr. 1926. Tokyo 1928.

. —— Pollinationsbiologie und Samenverbreitung auf den Juan Fernandez-Inseln.

Nat. Hist. Juan Fern. II, 1928.

. —— The geographical distribution of the sandalwoods and its significance. Proceed.

4th Pacif. Sci. Congr. 1929. III. Batavia 1930.

. —— Notes on some recent collections made in the Island of Juan Fernandez. Acta

Horti Gotob. 4, 1929.

. —-— Marine algal Communities of the Juan Fernandez Islands. Nat. Hist. Juan

Fern. II, 1941.

. —— Le peuplement des iles pacifiques du Chili. Soc. de Biogéogr. IV, 1934.
. —— Greigia Berteroi Skottsb. and its systematic position. Acta Horti Gotob. 11,

1936.

. —— Die Flora der Desventuradas-Inseln. Géteb. K. Vet. 0. Vitterh. Samh. Hand.

5:e féljden, ser. B, bd 5:6, 1937.

. —— Geographical isolation as a factor in species formation. Proceed. Linn. Soc.

of Lond., Sess. 150, 1938.

. —— On Mr. C. Bock’s collection of plants from Masatierra (Juan Fernandez) with

remarks on the flower of Centaurodendron. Acta Horti Gotob. 12, 1938.

. —— Hawaiian Vascular Plants. Ibid. 15, 1944.
. —— Peperomia berteroana Miq. and P. tristanensis Christoph., an interesting case

of disjunction. Ibid. 16, 1947.

. —— Eine kleine Pflanzensammlung von San Ambrosio. Ibid. 17, 1947.
. —-— The genus Peperomia in Chile. Ibid.
. —— Biogeografiska Stillahavsproblem. Statens naturv. forskn.-rad, arsbok 4 (1949-

50). Stockholm 1951.

. —— A supplement to the Pteridophytes and Phanerogams of Juan Fernandez and

Easter Island. Nat. Hist. Juan Fern. II, 1951.

. —— On the supposed occurrence of Blechnum longicauda C. Chr. in Brazil. Svensk

bot. tidskr. 48, 1954.

. —— The Vegetation of the Juan Fernandez Islands. Nat. Hist. Juan Fern. Il, 1953:

C. SKOTTSBERG

_ Steumer, H. Revision der Gattung Pernettya. Notizbl. Botan. Gartens Berlin 12,

1935.

_ Smrru, A. C. Taxonomic notes on the old world species of Winteraceae. Journ

Arnold Arbor. 24, 1943.
— - The American species of Drimys. Ibid.

c. Souza Loprs, H. & ALBUQUERQUE, D. Calliphoridae, Sarcophagidae. (Insect. J. Fern.

22.) Rev. Chil. de Entomol. 4, 1955.

_ Sparre, B. Review of Croizat, Manual of Phytogeography. Rev. Universitaria 38,

1953-

. Sreppins, G. L. yr. Additional evidence for a holarctic dispersal of flowering plants

in the Mesozoic era. Proceed. 6th Pacif. Sci. Congr. 1939, 1V. Berkeley 1941.

_ Sreents, C. G. G. J. vAN. On the origin of the Malaysian mountain flora. Bull. Jard.

bot. Buitenzores ser. 35 13-14) LO35—30.

_ SrockweLt, P. A revision of the genus Stenactis. Contrib. Dudley Herbar. 1940.
_ Srraneo, S. L. & JEANNEL, R. Coleoptera-Carabidae. (Insect. J. Fern. 23.) Rev.

Chil. de Entomol. 4, 1955.

_ Svenson, H. Monographic studies in the genus Eleocharis V. Rhodora 41, 1939.
_ Swezey, O. H. Distribution of Lepidoptera in Pacific Island Groups. Proceed. 6th

Pacif. Sci. Congr. 1939, IV. Berkeley 1940.
TayLor, W. B. An example of long distance dispersal. Ecology 35, 1954.
Tueriot, J. Mousses récoltées dans l’ile Mas a Tierra (Juan Fernandez) en 1927,
par M. Gualterio Looser. Rev. Chil. de Hist. Nat. 31, 1927.

. —— Mousses de l'Ile de Paques. Revue bryol. et lichénol. Ser. II: 10, 1937.
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Tonnoir, A. L. Australian Mycetophilidae. Proceed. Linn. Soc. N. S. Wales 54,

1920.
TRAGARDH, I. Acarina from the Juan Fernandez Islands. Nat. Hist. Juan Fern. III,

13h.

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. Tuyama, ‘T. On Santalum boninense and the distribution of the species of Santalum.

Journ. Japan. Bot. 15, 1939.

. —— On genus Haloragis and Micronesian species. Ibid. 16, 1940.
. Umecrove, J. H. T. See Continental Drift.
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Sci. Congr. 1939, IV. Berkeley 1940.

. Vernorrr, K. Uber Myriapoden von Juan Fernandez und der Osterinsel. Nat. Hist.

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. Vipat Gormaz, F. Geografia nautica de la Republica de Chile. Anuario Hidrograf.

de la Marina de Chile 7, 1881.

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I1gtg.

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Jian Hern. Ti 922).
Wattace, A. R. Island Life. London 1880. 3d ed. 1911.

. Weeks, L. G. Paleogeography of South America. Bull. Geol. Soc. of Amer. 59, 1948.
. Wecener, A. Die Entstehung der Kontinente und Ozeane. Braunschweig. 3d ed.

LO22, co tamedea moo)

. Wemarck, H. Studies in Juncaceae with special reference to the species in Ethiopia

and the Cape. Svensk bot. tidskr. 40, 1946.

. Weise, J. Coleoptera-Chrysomelidae and Coccinellidae von Juan Fernandez. Nat.

Hist. Juan Fern. III, 1924.

. WHEELER, W. M. Formicidae from Easter Island and Juan Fernandez. Ibid.

DERIVATION OF THE FLORA AND FAUNA 437

. Wixuts, B. Geotectonics of the Pacific. Proceed. 3d Pacif. Sci. Congr. 1926. Tokyo
1928.
. Wits, J. C. Endemic genera of plants in their relation to others. Annals of Bot. 35,
1g2l.
. —— Age and Area. A Study in the Origin and Distribution of Plants. 1922.
. Winker, H. Geographie, in Manual of Pteridology, ed. by F. Verdoorn. The Hague
1935.
. Wirtu, W. W. Heleidae and. Tendipedidae. (Insect. J. Fern. 7.) Rev. Chil. de
Entomol. 2, 1952.
. —— Ephydridae. (Insect. J. Fern. 20.) Rev. Chil. de Entomol. 4, 1955.
. Woo.tpripGE. See Continental Drift.
. Wutrr, E. V. An Introduction to Historical Plant Geography. Waltham, Mass. 1943.
. Wycopzinsky, P. Contribuci6n al conocimiento del género J/etapterus Costa de las
Americas y de Juan Fernandez. Rev. Chil. de Entomol. 1, 1951.
. ——— Thysanura from Juan Fernandez Island. Ibid.
. —— Simuliidae (Diptera). (Insect. J. Fern. 6.) Rev. Chil. de Entomol. 2, 1952.
. ZAHLBRUCKNER, A. Die Flechten. Bot. Ergebn. schwed. Exped. nach Patagonien
und dem Feuerlande. VI. K. Sv. Vet.-akad. Handl. 57:6, 1917.
. —— Die Flechten der Juan Fernandez-Inseln. Nat. Hist. Juan Fern. II, 1924.
. —— Die Flechten der Osterinsel, nebst einem Nachtrag zur Flechtenflora von
Juan Fernandez. Ibid. 1926.
. ZIMMERMAN, E. C. Insects of Hawau. I. Introduction. Honolulu 1948.
. ZIMMERMANN, A. Coleoptera-Dytiscidae von Juan Fernandez und der Osterinsel. Nat.
Hast. wan’ Fern: Tl 1924.
. Bryan, W. A. in “El Mercurio’”’, Valparaiso, Apr. 23, 1920.

. Gourtay, W. B. in “The South Pacific Mail’, Valparaiso, Feb. 2, 1928.
. QuenseL, P. Die Geologie der Juan Fernandez-Inseln. Bull. Geol. Inst. Upsala 11,

IQ13.

. Brancut, E. C. La Isla de Robinson. Valparaiso 1922.

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. Drupe, O. Handbuch der Pflanzengeographie. Stuttgart 1890.

. Merriti, E. D. The Botany of Cook’s Voyages. Chronica Botanica 14: 5/6. Waltham,

Mass. 1954.

. HILLeEBRAND, W. The Flora of the Hawaiian Islands. Heidelberg 1888.
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Hist. Juan Fern. II, 1924.

. SCHEDL, K. E. Chilenische Borkenkifer. Rev. Chil. de Entomol. 4, 1955.
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Chil. de Entomol. 2, 1952.

. Arwipsson, TH. Einige parasitische Pilze aus Juan Fernandez und der Osterinsel.

Svensk bot. tidskr. 34, 1940.

2. Hate, T. G. A fossil fertile Zygodiwm from the Tertiary of South Chile. Ibid.
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WW W
DW Ww Ww

oo |

©} N we

C. SKOTTSBERG

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337. ——— La Flora fosil de la region del alto Rio Chalia en Santa Cruz (Argentina).
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Svensk bot. tidskr. 34, 1940.

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342. —-— The Vegetation of Easter Island. Nat. Hist. Juan Fern. II, 1927.

343. Cooker, G.H. Te pito te henua, known as Rapa Nui. Rep. Smithson. Inst. 1897.
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345. Bricnam, W. T. An index to the islands of the Pacific ocean. Mem. Bishop Museum
Vol. I: 2. Honolulu rgoo.

346. Forsvrer, G. Florulae insularum australium Prodromus. Goettingen 1776.

347. —— A Voyage round the world ...commanded by Capt. James Cook 1772, 3, 4
and s. Vol. I. London 1777.

348. —-— De plantis esculentis insularum oceani australis Commentatio botanica. Berlin

1786.

Contents.

Part 1. The Juan Fernandez Islands.

Chapter I. Composition, distribution and relationships of the Flora ...... 193

Chapter Il. Sources of the island flora as judged by the total distribution of the
I . . . - J = ‘s . —
geographical elements, with special reference to the composition of the Chilean

We ef 3 tae Uva ML ot, ca teas Ga een ee A oe. eS et eee A
Chapter III. Composition, distribution and relationships of the Fauna .... . 292
Mech. Continental and Oceanic islands.<5 5°) 20). ea ke ALF
pee ¥. fhe Pacific Ocean and. Continental Drift-. .:9.. 52... .. : 325
Peery to bransoceanic: Inigration fi 0529S os, Sosy wele 8 ee Se ere 31
“hapter VII. Biological characteristics of isolated islands ......°..... 351
Peer viil. Evolution im: Oceanic islands! Je0 x25. (Fe Gh eS) eye ee - 363
Paer 1X. Juan Fernandez—oceanic or contimental? . . - 2 2.2042 es se. 372
Chapter X. The Chilean coast line and the history of the Andes ....... 380
Saager XI. Fhe Tertiary floras of Chile and Patagonia... ...-....-... 386
Chapter XII. Antarctica as a source of the present circumpolar floras .... . 389
Chapter XIII. The history of Juan Fernandez—a tentative sketch .-..... - 394

Part 2. Easter Island.

Chapter XIV. Composition, distribution and relationships of the Flora... . . 406
Chapter XV. Composition, distribution and relationships of the Fauna... . . 417
Chapter XVI. The biogeographical history of Easter Island .......... 421
enn STEAL Te oP! Se Ga cee Pe Pte Mae, the ee Et ode ie is. ad vo be gw eel 428

Printed October 26th, 1956.

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