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Full text of "The natural history of Juan Fernandez and Easter Island"

THE 



Natural History 



Juan Fernandez 



AND 



Easter Island 



EDITED BY 
CARL SKOTTSBERG 




THE NATURAL HISTORY 
OF JUAN FERNANDEZ 
AND EASTER ISLAND 

EDITED BY DR. CARLSKOTTSBERG 



VOL. I 
GEOGRAPHY, GEOLOGY, 
ORIGIN OF ISLAND LIFE 



WITH 14 PLATES 



AWE 




UPPSALA 1920-1956 

ALMQVIST&WIKSELLS BOKTRYCKERI AB 






707782 



Table of Contents. 

1. Skottsberg, Carl. Notes on a visit to Easter Island 3 

2. Hagerman, T. H. Beitrage zur Geologic der Juan Fernandez-Inseln .... 21 

3. QuENSEL, P. Additional Comments on the Geology of the Juan Fernandez 

Islands 37 

4. Skottsberg, C. A Geographical Sketch of the Juan Fernandez Islands ... 89 
5. Derivation of the Flora and Fauna of Juan Fernandez and Easter Island 193 



The Nat. Hist, of Juan Fernandez and Easter Isl. Vol. I. 



"\iica4a "Ho Orwo 




EASTER ISLAND 
after Ihe map of fhe Qrilean H\'dr. OHice 

-t:aooooo 

Ueiifhi 1/1 miners abox-e sea leve? 
. Obsrrvatiart spaf 



310 



/^i&oVo^uH 



I. Notes on a visit to Easter Island. 

 

■■. ' By 
CARL SKOTTSBERG. 

With 14 plates, i 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 191 7. A short and very popular account of our visit 
appears in my book »Till Robinson-on 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 (ew 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 Perouse 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 
Perouse on account of adverse winds; finally we resolved to cross the island 



, t ARL SKOTTSBERO 



(,n liorseback, and arrived at Matavcri, the scat of tlie farm house, on the 19th. 
hi the meantime u e liad made some excursions in the northeastern part, where 
Mt Katiki was ascended. At Matavcri we were cordially received by Mr. 
ri:k( V i:i)MlNhS, the mana-er, and were invited to to take up our quarters in 
his house. ( juite naturally, the natives were in a state of great excitement 
ovc-r the arrival of the vessel with their much beloved bishop, the missionaries 
(two Capuchin l^rethrcn) and the many useful articles reported to be onboard; 
and conse(iuentl\- the\- wore 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 })ait of our scientific baggage, which had been left on the beach 
at La Perouse, transported to Mataveri. From our headquarters the district 
round Hanga Koa and Ilanga Piko etc. was visited and several trips under- 
taken to Kano Kao and Orongo, the famous stone village. Further, our work 
was exlendend to Kano Aroi and Mt. Terevaka, the highest mountain, and 
also to tlie 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 ca{)acit\' for hard work had become reduced on account of illness. 
The >Ha«iucdano» left ICaster Island on July 1st. 



(IKNKK.M. NOTF.S ON TIIK GEOGKArilV OF EASTER ISLAND 

Tlic topographical features of the island are fairly well illustrated on the 
accom[)anying map, the result of Chilean Navy surveys of later years. The 
position of the observation spot in Ilanga Roa is given as Lat. 27° 08' 06" S., 
Long. 10/ 25' 54" W. Mrs. Roitledge's map is based on U. S. Hydro- 
graphic ()ft'ice chart no. i 1 19, 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 tlic 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. art- properly s|)elt and righth' placed. 

1 lie island is known to be wholly volcanic. There are no signs of recent 
action, save for a couple of tcj)id sj^rings below high water mark reported to, 
but never seen l)\ us. It is rather curious that both THOMSON (Smiths. Inst. 
Ann. kcp. iSS.^. Washington 1S91) and Agassiz (Mem. Mus. Comp. Zool. 
( ambridgc 33, i<i(>9) should discuss volcanic eruptions and great earthquakes 
as a possible reason for the destruction of the megalithic monuments and for 
the (lisappearani e 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; someiinies thev 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. 1 lie tufas and ashes of the numerous cones present a 
great variation ot colour contributing to lessen the monotony of the scenery. 
The attention of the visitor is especially drawn to the three great A"^;/^. 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 i 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 buUrush, an 
endemic variety of the widespread Scirpus riparius, called paschalis by Dr. 
KUKENTHAL. 

The country NE of Rano Kao is hilly, one of the cones being known as 
Punapau (Plate i), 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 Perouse 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- 



TAkL SKOTISUKKG 



health. The slope of Katiki (another name not found on Mrs. RouTLEDGE's 
map, but frecpienlly used) is comparatively gentle. The top was found to be 
412 m. higli (300 m. on the Chilean ma[) 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 tlie action of the sea. The one nearest Katiki, Vaintu 
Rova, is of a light yellowish colour; the liight is 310 m. On the south slope 
wc 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 
|)erfectly fresh, being considerably above the level of the sea; but it was dirty, 
(Mving to the filthiness or cleanliness (call it what you will) of the natives, who 
never go to drink v.ithout washing themselves all over as soon as they have 
ilone . . . (the edition in Iweryman'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 llanga 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, i 218 
mm., the average of 8 years' observations. This is, indeed, no small amount, 
surpassing that of Juan h\'rnandez, where erosion has modelled the entire island 
into a system of deej) valleys and sharp ridges, l^ut 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 
(juitc astonishing. The climate is warmer in Easter Island than in Juan Fer- 
nandez, the evaporation undoubtedly much greater, the winds at least equally 
tre<iucnt. Anybody will note the rapid disappearance of the water; after a 
licavy raiti, 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 j)orosity of the soil. Occasionally, water is 
encountered by digging deep holes; but to dig through the hard rocks must 
have l)ccn too difficult a matter for the natives. Subterraneous streams are 
reported, and arc, of course, to be expected; and running water has played an 
important })att in the formation of the numerous caves round the coast. 

1 iic 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, cxci-pt for a few specimens in the crater of Rano Kao, where the last 
stunted liwarfs of the famous toromiro (SopJiora foroiiiiro) still linger, in com- 
pany with mahute ' Ihvnssnurt/a ptipyrifcni), hau-hau (or jau-jau, Spanish j), 
called //////;/A/A/.- l,v l-t I MIS I believe that it is T. scmitriloba\ and \:\ [Cor- 



NOTES ON A VISIT TO EASTER ISLAND 7 

dyline iennhialis 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 formo (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 n6t 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. 

^ In Anuario Hidrogr. de la Mar, de Chile (30) 1916, p. 55 C. De la Maza mentions a 
small indigenous tree which he calls »tumahiti». »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, 
I 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 Melia, but this is called miotaijl (vide 
F. FuENTES, Resefia botanica de la Isla de Pascua. Inst. Centr. Meteor, y geofi's. de Chile. 
No. 4. Santiago 191 3). Tumahiti sounds like a corruption for te mahute, the paper mulberry, 
which certainly cannot supply materials for the implements mentioned. 



S CARI. SKOIISBEKG 

The dwarf trees now cxistiiiL; are, as has already been stated, almost 
wholly confined to the crater of Rano Kao and are on the verge of extinction. 
They ^row not far from the lake, where the steep slope is covered with very 
larj^e 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; 
unfortunatls', 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, widespread species, 
either Polynesian or introduced from the old world via Chile, Tahiti or other 
places. Occasionally ferns arc found, also outside \.\\c x^no, Microlcpia strigosa 
being the most common. We discovered two species of Ophioglossum, to which 
the natives attribute medicinal (jualities. 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. 



ARCIIAKOI.OGICAL REMAINS: THE AHU 

Tliese structures, the burial-places — but not the only ones — of the is- 
landers, liave been called » terraces* by most authors. Such a word tells us 
very little, while the word ahu is a proper technical term, strongly and justly 
recommended by Mrs. RoUTLEDCH-: 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 u{)set as a result of internal warfare. During our staty in La 
Tcrouse Hay we devoted some time to the inspection of the ahu. One not far 
from the landing-{)lace, 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. 'Ihis ahu was measured and described (see Plates 3-4 and text fig. l). 
The central part is 23,; 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 1,35 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, o,y high, and 0,45 broad, another 2,45 long and 
I in high. They are closely (itted. The space behind this wall was filled 
with bouUkrs, the surface paved with larger, flat stones, making a level 
platform. ^ 

I he wings are larger than the centre, the cast 31,3, the west 32 m., giving 
as total length of the ahu 86,7 m. Their front wall is as high as or even 
higher (about 2 m) than that f)f 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. Ihe aim sl()i)cs gently inwards. This .slope, which could be 



NOTES ON A VJSIT TO EASTER ISLAND 



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 1,9 X 0,7 m. with a depth of i 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. 






I ' '  ' (K M 1/ 1(1 I U '!'<(/ ■. 






Fig. I. Diagrammatic sketch of an ahu west of the landing-place in La Perouse Bay. 
Scale I : 750. s fallen image, /i hat, 7/1 stone- wall, ^ vault. 

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



Total length 10,27 ni. 
Length of body 6,55 m. 

» » head 2,52 m. 

» » neck 1,2 m. 

Width of body at base 2,y m. 
Thickness of body at base 1,6 m. 



Width across shoulders 3,2 m. 

» » head 2,6 m. 

» » neck 1,9 m. 

Circumference round shoulders 7,9 m. 

» » neck 5,2 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 Faro, 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 



CAKL SKOTTSHKRG 



that It is the hir^cst one ever found on an ahu and the last one to be upset. 
Are there two exactly siniihir ahu close to each other, each with an unusually 
lari^e imager I think not. The measurements were taken by my wife and 
myself with a tape 25 m. loni; and the figures committed to paper on the spot. 

It couUl po>sibly also be the same ahu as no. 34 Punahoa of THOMSON 
1. c, p. 505. lie also <^ives the total length of the single moai as 32 feet. 
Ilmvever, the entire structure is said to have a length of 175 feet and a width 
«)f S feet, which figures must be entirely incorrect if ahu Pare is meant. 

\W what kind of apparatus or devices the statues were transported from 
Kano Raraku to the coast, in some cases to rather inaccessible places, remains 
a m\ster\'. 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-bodietl men» (p. 49S). 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 fioor in order to serve 
the puri)ose, the images being rather fragile. Mrs. R. has traced the few high- 
wa\-s leading from Rano Raraku to the coast; but if really the images were 
draggetl 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 
re(|nired could have been brought together. It is astonishing that no tradition 
on the means of transport survives. According to Mrs. R. the natives in- 
variabl\- offered one exi)lanation: that the images were transported by the aid 
of supernatural forces. 

< )n p. 4X6 Thonlsox discusses the possibilit)^ of a transport by sea. Near 
a grouj) of ahu he discovered a fine landing-place made by art, »admirably 
adapted to the landing of heavy weights*. hVom old drawings we know what 
the aboriginal canoes were like — not a single one, as far as I know, has been 
preserved to our <la\s but they were not strong enough to support any very 
heavy weight ( )tie might suggest that large rafts were built but, on the other 
hand, there are several ahu which are unaccessible from the shore. 

.Still, there is another nietod to be reckoned with, although further specu- 
lation on this matter may appear j)retty useless. Some sort of a sledge-like 
apparatus could h.i\c been cntistructed without the need of timber of any con- 
sitleral)le si/.e. .\ sledge would slide (luite well over the grass, provided that 
the road was cleared from stones. A great number of people could be simul- 
taneously engaged in j)n!liiig, while, if rollers were used, the image must have 
been more diffK-ult t(» handle. ( )nce 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 aj^plied when raising the 



NOTES ON A VISIT TO EASTER ISLAND II 

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 9 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 

^ Geiseler (resp. Weisser) describes a statue of 23 m. length (Die Oster-Insel. Berlin 
1883, p. 9). 



12 CAKI, SK.OTTSEF.RG 

neither the iinaLTcs stamlini; on the mountain nor those found scattered over 
the ishuui were ever intended for the ahu, but that the latter lined the roads 
leadin<; from the mountain to the coast. I cannot add anything to the ex- 
phmation of the standing statues. If they were put up to celebrate »bird- 
nien». it seems cjuaint that not tiic 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. 

I'he rutle stone implements (toki) used by the sculptors are often found. 
In Ilanga Koa we came across a large and very well wrought stone adze of a 
ratiier international type, but not found in any of the accounts on the island. 
It had been picked up on tlie seashore at low water and is quite incrustated 
witii the siiells of animals. It measures 20 cm. (Plate 14, fig. i). A few stone 
chisels were also obtained; two are figured on Plate 14, fig. 2, 3. 



KKM.MNS OV llorSK.S ANP PLANTATIONS 

I''oundations of old houses are seen in many places, and several were 
noteil on our excursion to Alt. 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. The\- 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 pef)ple do not seem to be sure as to their former use. A fine 
tower at the landing-place in Hanga Ho Orno (La Perouse) 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,.» m. and is 6,; 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 j^reat ahu. Mrs. R., on the authority of some resident, explains 
these structures as look-out towers whence watchers on land communicated the 
wlicreabouts of the fish to those at sea; these contained a small chamber 
below which was us(-d as a .sleeping aj)artmcnt» (p. 218). 

It sounds strange that these solid towers should have had no other pur- 
pose; tlian to serve as lookout 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 
»tish are not picntifuh — an observer must seek an eminence dominating a 
considerable space of water. TiioMsox tells us that from the towers the move- 
ments of the turtles wer(> watched. The ob.server must keep outside the 
t(nver. or on the top of it, not a very comfortable place. Really, the tower 
itselt would have been little more than a refuge in bad weather and during 
the night, but for such a jnupose 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. 





2. Diagrams of old plantations; a seen from above; b three types of shelter, in section. 
I = bananas, 2 = Melia azedarach, 3 = Andropogon halepensis. Scale i : 230. 



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 Pi^ROUSE's 
Voyage, reproduced by Stolfe 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 



CAF^L SKOTTSBERG 



crepancics bctw ccn tlic 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 PIiROUSE's 
and I'lNAKl's narratives designs of ahu which do not at all correspond to 
modern descriptions or photographs. 

In the vicinit)- ot Ilanga IIo Orno we saw many remains of native planta- 
tions. 'ihe\- are of several types. One, seen in fig. 2 a, is probably of a 
later date, as the material has been taken from an ahu, the front wall of 




l-ur V n. Tsv.) .iMrd-mcii)> on rock at Oron^o (hciolu ol the rock 1,6—1,7111.) b. Incised marks 

on door poost at Orongo. 



which fornix the back wall of the garden. Circular miniature gardens are 
represented in fig. 2 /'. The need of shelter and moisture is well unterstood. 
Mclia was said to be u.oun for the sake of the timber. Probably it is of recent 



intii)(iuction. 



•>K'»N(;o AND WW. lURI) cui/r 



Mrs. koriiKixiK (i,-votcd nuich time to the survev of the Orongo village, 
an(l as a detailed plan was made and every hou.se measured and described, I 
shall content mysell with a few short remarks. 

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



NOTES ON A VISIT TO EASTER ISLAND 



15 



vide fig. 3 b. 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 
»ariki-mau», vide 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. VlVES, 
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 i? 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 before 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 ofi"ered 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 EDWARDS. 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 



,6 CARL sKonsr^KKG 

the earliest liiscoverers liad seen Mude carthen-\vare» on the island, a statement 
due to some misinterpretation. Of stone implements, besides the toki, and the 
stone adze and chisels, we t;x>t one lishiiook, 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 tlie si)hcrical 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 stonc». llioM.soN has described and figured many such stones, but 
none of them i)resent any likeness to this one. 

The object on Plate 14, Fig. 6 is not, as might be suspected, a broken 
spear-liead or ))iataa, 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 
ct)mmonl\- found in the soil and also manufactured to satisfy the demand of 
visitors. Two, of an ordinar\' type and apparently old, are seen on Plate 14, 
h'igs. 7, S. According to THOMSON there were at least nine kinds, all with 
dilVerent names, a statement well needing the corroboration of Mrs. R. 



ORKIIN OF THE PEOPLE 

The histor\- of Faster 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 imj)ortant conclusions from the legendary traditions still alive. A tale 
of two ditTerent races and two successive colonizations runs through the old 
legends. Tlie anthropological evidence seems to be in favour of a double origin, 
Melancsian and I'oKnesian. The comparative studies of the Bird Cult in the 
.Solomon Islands and I^aster Island (by PI. Balfolr, vide Mrs. R.) seem nothing 
less than convincing. The bird rei)resented in the numerous carvings, paintings 
etc. of ICaster Island is not the holy bird of this place, but the frigate bird, 
u<»rshi|)|)cil in the Solomon Islands. 'Phe bird figures were called penguins by 
Lkhm.wn (l-.ssai dune bibliograi)hie, Anthropos, 1907), which undoubtedly 
must be a mistake, especiall)' as penguins hardly ever visit these waters. 

it wi- sum up the results obtained, there is evidence that the Easter Is- 
landers ate ot a twofold origin and that, after the Melanesian immigration, a 
P(»l\-nesian immigration followed. l"he population now tends to assume a 
multicoloured aspect; there has been a late influence from Tahiti (so we were 
told) and various white men ha\'e contributed towards the »amelioration» of the 
race. The xouul; girl figured on Plates 12-13 ^^•'■'^ ^^^^ to be of » pure Easter 
Island race , but whether rejjrcsenting a IVlelanesian or Polynesian type, I am 
unai)le to trli. 

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 viaunga 
(mountain), inaJiul, (paper mulberry), // (Cord^line), ciimara (sweet potato) a. o. 
A large vocai)ular\-, collected by Padre Roi SSEI,, was publi.shed in Santiago, 



NOTES ON A VISIT TO EASTER ISLAND 



17 



191 7, but it miist be used with much criticism, as it contains many Tahitian 
words and also corrupted EngHsh, French or Spanish; good examples are anio 
(agneau) and rnutone = sheep, himene (hymn) = to sing,' teperanate = serpent, 
tokini = stockings, tiaporo (diablo) = devil, viretute = 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 reHgious 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 (1. 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 RoG- 
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. 
191 1): »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 ubas? Most likely the same word 
as the Malesian ubi (uwi, huvvi), yams (Dioscorea alata), now called ufi 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 gehort 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 [Ipomaea batatas or Batatas 
edulis) from New Zealand through Polynesia to Easter Island. According to 
Cheeseman (Manual of the New Zealand Plora) 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. 



I 8 CARL SKOT'ISBI-RG 

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 legion 
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 etimolojico. Santiago 1910), the word cumara is found in the 
Quichua language; it is not indicated as the principal name of the sweet potato, 
which is apichii, 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 ICuropeans. 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 
H.XLI.IKR (Uber friihere LandbriJcken, Pflanzen- und Volkerwanderungen zwi- 
schen Australasien und Amerika. Mededeel.'s Rijks Herb. Leiden 13, 191 2) 
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 
Ro<;gevkkn as existing 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 barahii mentioned by F. ViDAL GoRMAZ, Jeografia nautica, 
p. 177 (Anuario Midrogr. 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 1 9 

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 Edw.ards 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 ho 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 

^ 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. 



2 CARL SKOTTSBEKG 

re^^ardini,^ the cxistino^ monuments. Last year a small book appeared, entitled 
»lJet sunkne kontinent (Atlantis)», b\- a Xoruegian, C. SuND (Copenhagen I919), 
where also the supposed Pacific continent is spoken of. As might be expected, 
I'Lastcr Island forms an important item. With my permission, two of my photo- 
<,'raphs 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 lie should not happen to draw from a single reliable source, not even 
from m\- popular dcscrif)ti()n, which was known to him. Mr. SUND tells us of 
the Iv^^vptian influence in Easter Island, of the enormous foundation walls and 
ruins of temples; almost ever\- mountain had sculptured designs of goods, fishes 
and p\'ramids, the cave paintings w:ere in the l^oltec 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 Egj^ptiari 
one, even with the same kind of cement. All materials, bricks, glass, porcelain, 
ever>thing was known in I£aster Island; religion, symbols and habits were the 
same as in I\gypt, 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. SUNI) 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 
bv such fantastic inventions. 



iMually, 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. I'KK( V I-:i)MiN])s, Mr. I. ViVES and Baron Erland NoRDENSKlOLD. 



Explanation of Plate 14. 

1. StiMic ;u1/.:, no. U). i. 307; iu)t quite V-'- 

2. .Stone ciiiscl, no. 10. 1. 320; alino.st ^i. 

V  >^ no. K). I. :;:i ; ,^ a,. 

4. I'lsh hook of stone, no. iq. i :,2y, almost "/;. 

V Stone 1^11. no. 10. I. 500, not qu;?e nat. si/e. 

(\ Knite or scrape- no. 19. i. 31^, not ciuite nat si/e. 
7. Speai-Iie.id, no. ly. i. 31 -,. •' „ 

■'^- " . UO. lU. I. :;i J . -j.:. 

'I'lie orii,'inaK in the .Museum. Ciothenbiii". 



Nat. Hist. Juan Fernandez and Easter hi. Vol. I. 



Plate i 




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Nat. Hist. Juan Fernandez and Easter IsL Vol. I. 



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Nat, Hist. Juan Fernandez and Easter Isl. Vol. I. 



Plate 5. 




.\(//. llist. '"fUiVi l-t))iiuhii:: auJ l.astiy I si. \'ol. I. 



Plate 6. 



/ 




m 



Nat. Hist. Jtian Fcrnartdez and Easter Isl. Vol. I. 



Plate 7, 




\<i/. Hist. '7ua)i l\))iiviiii:: a)id luistcr Isl. ]'ol. I. 



Plate 8. 




Nat Hist. Jtian Fernandez and Easter hi. Vol. I. 



Plate 9, 




3 o 

0) 



5 rt 



la 



\(!/. Hist, 'lua)! l-\>)uvuii:: a)id f.astir I si. I'o/. I. 



Plate io. 




Nat. Hist. Juan Fernandez and Easter Is/. ]^o/, I. 



Plate ii, 







Xd/. Hist. Juan Foiitvidcz ajid liastcr Is/. J 'of. I. 



Plate 12. 





Nat. Hist. Juan Fcj'nandez and Easter Isl. Vol. I. 



Plate 13. 




Photo by K. B'dckstrbm 



Same girl as in Plate 12. 



.\(i/. I list. '7iui// J-\)iiii)!(/i-: ivid Juis/ir Is/. I'ol. /. 



Pl.ATK 14. 




2. Beitrage zur Geologic der Juan Fernandez-Inseln. 

Von 

TOR H. HAGERMAN. 

Mit 12 Textfiguren. 

Die Schwedische Pazifik-Expedition 1916 — 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 iibergeben 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 Verfiigung. 

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 diirften, 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 Canon-Landschaft gebildet. Die wilden Terrainformen sind deutlich 
aus den zahlreichen Photographien zu erkennen, von denen viele in Skottsberg's 
Reisebeschreibung^, wie auch in Vol. I und II dieses Werkes veroffentlicht 
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 konnen als verschiedene Erstarrungsformen ein 



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

2 L. c. p. 256. 

3 C. Skottsberg, Till Robinsonon och varldens ande. Stockholm 1918. 

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



2 2 TOR H. HAGERMAN 

unci (lesselbcn Magmas betrachtet werden; spaterhin sind allenthalben stellen- 
wcisc sckundare Veranderungen infolge hydrotliermaler Prozesse entstanden, die 
abweichende Ausbildunoen hervorgerufen haben. 

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

Charakteristisch fur alle diese verschicdenen 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 Aciisen. 13och ist zu bemerken, dass derselbe 
nicht zu den extremsten Typen gehort. 

Audi die Felds{nite weisen eine konstante Zusammensetzung auf. Sie sind 
fast aile zwillingsgebildet nach dem Albit- oder Karlsbader-Gesetz. In Schnitten 
senkrecht zu M zeigen die Albitlamellen eine max.-Auslosung von 32 — 33°, in 
einigen iMnzclfallen diese W'erte mit hochst 2° variierend. Der Feldspat ist also 
ein Labrador von eincr Durchschnittzusammensetzung Ab42An58. 

Da die Hasalte also mineralogiscli einander nahe vervvandt sind, wurden sie 
liauptsaclilich nach der Struktur in folgende Typen eingeteilt: 

basaltische Laven, (teils dichte, teils grobkornig doleritische, teils schlackige) 
Tuftc, hydrothermale Um\vandlungS[)rodukte. 

Basaltische Laven. 

Diese (iesteine bilden, wie friihere Verfasser bereits betont haben, den Haupt- 
bestandteil der Insel. Auf Grund des vorliegenden Materials konnte man den 
X'erlauf der verschicdenen Lavastrome und deren Neigungsverhaltnisse nicht be- 
stinimen und so auch keine Klarheit iiber die P>uptionsstellen erlangen. 

Die Fundstiitten der dem Verfasser zur Untersuchung vorliegenden Hand- 
stucke sind ziemlich gleichmassig iiber die Insel verteilt. Von den zu den Laven 
gchorenden (iesteinsproben sind ungefahr ^/^ sehr porose und schlackige Typen, 
wahrend ^3 (6 Stck.) dichte, dabei gleichmassigere und feinkornige Gesteine 
darstellen. 

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

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

Die I'Linsprenglinge sind Olivine und ihre Umwandlungsprodukte sowie 
i'eldspat. Die Augitkorner konnen sich zuweilen der Grosse der Einsprenglinge 
naiiern. 

Die Olivineinsprenglinge erreichen ihre grosste durchschnittliche Ausdehnung, 
ca. 2 mm, in Proben, die von dem nordlichen Ufer der Padrebucht stammen. 
Gerade diese ICinsprenglinge zeigen meistens in einer scharf begrenzten Zone 
die von Oll.NSF.i.' fruher beschriebene Iddingsitumvvandlung. Aus Dunnschlififen 
der genannten (icsteine ist klar ersichtlich, wie die Umwandlung von den Ran- 

' L. c. p. 260. 



BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ- INSELN 23 

dern nach der Mitte zu ausgegangen ist und sich 0,02 — 0,03 mm in den Olivin 
hineinerstreckt hat. An manchen Stellen ist die Umwandlung langs der Spalt- 
risse vor sich gegangen, wahrend anderweitig die Durchgange merkwiirdigerweise 
vollkommen unveranderte Teile der OHvinkerne durchqueren. (Vergl. Fig. i.) 
In der Gesamterscheinung des obengenannten Praparates mochte der Verfasser 
die Iddingsitbildung als das Resultat einer von aussen kommenden chemischen 
Beeinflussung ansehen. WASHINGTON^ verweist die Iddingsitbildung bis auf mag- 
matischen Ursprung zurijck. Man braucht vielleicht nicht so weit zu gehen, da 
dieselbe ebensogut einer hydrothermalen Umwandlungsperiode zugeschrieben wer- 
den kann. 




Fig. I. An den Randern iddingsitumwandelte Olivine. Olivinbasalt von der Padrebucht. — 

Vergr. 56 x . Photo Hj. Olsson. 

In einem anderen Gestein von der Padrebucht, das etvvas feinkorniger ent- 
wickelt ist, treten auch Feldspateinsprenghnge von 0,5 — i mm Lange auf, neben 
Resten von Olivinen. Der Olivinumwandlungsprozess hat hier ein anderes Pro- 
dukt hervorgebracht, namUch gewohnHchen Serpentin. Dies ist auch bei den der 
Grundmasse angehorenden OHvinkornern der Fall. Dieselben sind vollkommen 
als Serpentin ausgeflossen. Auch der Augit scheint an den Randern etwas ange- 
grififen 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 ii. M.) SW. von Tres Puntas, W. von der Villagra- 
Bucht, herstammend. Iddingsit- und Serpentinumwandlungen der Olivine sind, 
wie ich besonders hervorheben mochte, 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. 
1st man erst einmal auf diese spharolitahnlichen Bildungen aufmerksam gewor- 
den, so findet man sie haufig hauptsachlich in den feinkornigsten Proben dieser 
Gesteine ausgebildet. Besonders schone Beispiele hiervon zeigt ein Handstiick, 
das der »ryramides einem Ikrggipfel ungefahr in der Mitte der Insel unvveit 
des SKIA'IKK-Denkmals, entnommen wurde. (Fig. 2.) 

lune sehr ahnliche Krscheinung ist von Reiter^ beschrieben worden. Er 
sclimolz 45% Albit, 45% Augit und 10% Magnetit zusammen. »Der Schliff 



V /r 




Pyroxen (-^^ ">PU<tioklAS ^ M Ag Weht 



Fig. 2. Phigioklas-AugitSpharoIit. I5asalt von der '>Pyramide». — Vergr. 250 x 

Zeichnung vom Vcrf. 



eincr durcli 7 Stundcn abgckiihlten Schmelze zeigt eine sphariodale Anordnung 
der Kristalle, in dem niagnetitreiche Kerne von Glaspartien mit einzelnen aus- 
gcschiedenen Augit- und Plagioklasleisten und Kristalliten umgeben sind. Die 
W'iederhokmg des X'crsuchcs bei 30-stundiger Abkiihlung ergab eine Schmelze 
mit kornig-pori)hyrisclicr Struktur. Zweifelsohne ist dies audi in dem vorliegenden 
l^'alle zutrcffeiid, indcm die s{)harolitrulirenden Laven einer raschen und ungestorten 
Abkuhlung ausgcset/.t gewesen sein durften. 

In Anschluss an dicse Spharolite seien hier die in Pig. 3 abgebildeten kreuz- 
formig licgenden Olivinkristalle erwahnt. Der DiinnschlifT entstammt einem etwas 
grobkornigen Gestein von Bahia Cumberland. Wie aus der Figur deutlich her- 

^ U. H. Rkii F.R, Kxporimentellc Studien an Silikatschmelzen. Neues Jahrbuch. Beil. Bd. 22 
1906 , p. 197. 



BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ-INSELN 



25 



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



Im Vaqueriatal tritt, wie aus der untenstehenden Photographic (Fig. 4) er- 
sichtlich, ein fast horizontal liegendes Gestein auf. Nach ihrer grobkristallinischen 
Struktur zu urteilen, sind diese Basalte 
moglicherweise als intrusiv aufzufassen> 
Das nur an zwei Seiten zugeschlagene 
Handstiick ist vorziiglich durch Schrump- 
fung unter rechtem Winkel zerkluftet. 
(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- 
selbe wie jener der Vaqueriaprobe nur 
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 iibrigen dem vorgenannten Gestein vollig gleich, ist 
der bei »Tres Puntas» genommene Basalt. Die Handstiicke bestehen aus langen, 
schmalen, dreiseitigen Prismen. 




Fig. 3 a. 



Zentrisch angeordnete Olivine. 
Vergr. 56 x . Verf. phot. 



Unter den schlackigen Laven weisen einige eine auffallende Analogic 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. 

2 L. c. p. 263. 



26 



TOR H. HAGKRMAN 



Tuffe. 

Unter dem mitgebracliten Material befinden sich zwei Proben von ausge- 
sprochenen Tufien. Der eine, ein poroses, dichtes Gestein, stammt von El Puente, 
dem Istmus zwischen der Padrebucht und Carbajal und ist ein Palagonittuff mit 
einigen sporadischen Augit- und Magnetitkornern. Der andere Tuff stammt von 
dem nordlichen Ufer der Padrebucht, von wo einige umgevvandelte Olivinba- 
salte (s. S. 23) herriihren. Ausser Augit und Magnetit enthalt derselbe einige 
grossere vollkommen reine Olivinkorner in einem teilvveise kryptokristallinisch aus- 
sehenden Glase. Stellt man diese verschiedenen Bildungen aus der Nahe der 




Fi^. 3 b. Detail von Fig. 3 a.— Verg^r. 170 x. Verf. phot. 



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

Hydrothermale Bildungen. 

K\n aragonithaltiges (iestein vom Ufer gleich siidlich vom Yunquegipfel diirfte 
als liydrothermal umgewandelter Basalt angesehen vverden. (Fig. 6.) Das Hand- 
stuck ist ein von weissen Streifen durchzogenes scharfgriines Gestein, das u. d. M. 

1 L. c. p. 266. 



BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ-INSELN 



27 



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 eigentumliche Bildung dargestellt, wie sie auf dem offenen 




Fig. 4. Das Vaqueriatal. Wasserfall iiber den saulenformig 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 iiber 
eine Vegetationsdecke geflossen sind, wobei Wurzeln etc. mit einer Kruste von 
oben angegebener Zusammensetzung iiberzogen 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- 



1 L. c. p. 265. 



28 



TOR H. HAGERMAN 




hangt. Sowohl in Diinnschlifif wie in Handstiick 
zeigt dieses Gestein, das als fast vertikale Gange 
auftritt, eine voUige tjbereinstimmung mit einem 
der dichten Basalte von Masatierra vom Gipfel des 
Cerro Negro SO von Yunque, 190 m ii. d. M. Ur- 
spriingliche 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 natiirlich vollig in Serpentin 
umgewandelt. Das ganze Praparat ist von Ilme- 
nitskeletten durchwachsen. 

Masafuera. 



I'ig. 5. Handstiick voin Vaqueria 
K'ang. Verf. phot. 



Etwa 180 km vvestlich von Masatierra erhebt 
sich die Insel Masafuera. Abweichend von Masa- 
tierra in Bezug auf die einheitliche Mineralzusammensetzung der Gesteine Uefert 
Masafuera Beispiele petrographisch weit verschiedener Typen. Basalte mit den 
dazugehorenden Gangformen von ungefahr gleichem Mineralbestand wie die auf 
Masatierra vorkommenden gibt es zvvar 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 angefiihrten Natrontrachyte. 



Natrontrachyt. 

Leider ist das einzige mitgebrachte Handstiick dieses Gesteins von einem 
losen Block am Fuss der Steilwand von Tierras Blancas abgeschlagen. Skotts- 
HER(; hat indessen miindlich berichtet, dass zahlreiche Blocke desselben Gesteines 
in den Talusbildungen von Tierras Blancas vorkommen, und dass, soweit er ver- 
stehcn koniitc, dasselbe hellgraue Gestein den ganzen oberen Teil der Steilwand 
bildct; seiner Kartenskizze nach zu urteilen tritt dasselbe bereits 400 m ii. d. M. 
auf. Dies ist von Bedeutung fiir das Feststellen der Eruptionsfolge, die spater kurz 
erwalint wcrden soil. 

Das (iestein besteht aus gleichmassigen Kornern und ist sehr reich an Feld- 
spat. Die I^'eldsj)atleisten erreichen eine Lange von 0,3 — 0,4 mm. Sie sind gut 
parallclorientiert und verleihen dem Gestein eine trachytoidale Struktur. Da Albit- 
zwillinge nicht vorhanden sind und der Feldspat durch die Anlagerung der diinnen 
Individucn unscharfe Bcgrenzungen zeigt, konnte eine genaue Bestimmung des- 
selben nicht ausgefuhrt vverden. Die Lichtbrechung halt sich im allgemeinen etwas 
liber Kollolith (n 1,535), stellenweise ist das Relief jedoch ganz verschwunden. 
Urn eine nalierc Kcnntnis von den Feldspaten zu bekommen, ist eine Alkalibestim- 
mung des Gesteins ausgefuhrt worden. Diese ergab 3,45 % KgO und 7,34 % NagO. 
Dies wiirdc einem Gehalt von 20,44 % Ortoklas und 62,27 % Albit im Gestein ent- 
sprechen. Auf (irund der Lichtbrechungsverhaltnisse diirfte jedenfalls neben einem 
Kali-Xatronfeidspat auch cin 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 gegeniiber zeigt. 




..— Aragonitband, von 
einem Augitindivi- 
duum iiberquert 



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



J 






9 


m 


i^fe' 


^■^M 








---^r 


'^:f^ 


y • « 


 . ' z , . 


. . . * 





Fig. 7. Sinterbildung von Puente. Massstab in Cm. Verf. phot. 



Der von QuENSEL^ beschriebene Natrontrachyt ist ungefahr 1200 m ii. d, M. 
gefunden worden und zeigt bei einem Vergleich einige Abweichungen. Das Hand- 
stuck 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. 

1 L. c. p. 283. 



30 



TOR H. HAGERMAN 



Andesit. 



Von (lem Berggipfel »Las Torres >, 1 370 m ii. d. M., und in losen Blocken 
von dem Ik'rgrucken XO von der genannten Stelle 1 200 m ii. d. M. sind Hand- 
stiicke von einem feldspatreichcn aschgrauen Gestein mitgebracht worden. Obwohl 
etwas olivinreicher, stelien dieselben den von Olensel^ beschriebenen, einem 
1 100 m ii. d. M. liegenden Niveau entnommenen Trachytandesiten sehr nahe. Als 
Kinsprenglinge koniinen Feldspat und Olivin vor, u. d. M. zeigt der erstgenannte 
eine Zusammensetzung von AbggAnga 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 0,05 mm. Die Magnetite sind vollig idio- 
morph. Die Klassifizicrung hiehergehorender Gesteine ist etwas unsicher. Die 
trachytoidale Struktur konnte auf einen gevvissen, nicht wahrnehmbaren Alkali- 
gelialt in der (irundmasse deuten. Geniigende Griinde, sie als Trachytandesite zu 
bezeiclinen, liegen jedoch nicht vor. 

Sehr iiiteressant ist ein bei Correspondencia (1420 m ii. d. M.) genommenes 
(iestein. Mikroskopisch zeigt dieses Handstiick ein aschgraues, porphyrisches 
Aussehcn, doch sind einige Partien bedeutend dunkler. Als Kinsprenglinge kom- 
men 1^'eldspat und Olivin vor. U. d. M. zeigt der erstgenannte eine Zusammen- 
setzung Ab42An58 und erreicht eine Korngrosse von durchschnittlich 2 mm bei 
einer niax.-Lange von 5 mm. 

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

In den dunkleren Schlieren finden sich nun Olivine mit Magnetiteinschliissen 
vollgesteckt. Iksonders an den Randern ist der Magnetit so reichlich vorhanden, 
dass die Olivinkorner vollig opak sind. Sowohl aus diesem Grunde als auch in- 
folge der abgerundeten Form der Mineralkcirner scheint es, als ob diese Olivin- 
korner einer kraftigen Resorption ausgesetzt gewesen waren. Naheliegend ist nun, 
dass diese dunkleren Schlieren mit ihrem grosseren Kisengehalt Riickstande auf- 
geloster Hruchstucke sind, welche urspriinglich zu dem Typus gehorten, die einem 
schlackigen l^asalt vom (jipfel des Inocentas entsprechen, der spater beschrieben 
werden soil. 

In einer luitfernung von kaum i^/g mm von einem der erwahnten Olivin- 
korner treten Individuen des anderen Typus auf. Diese sind ganz einschlussfrei, 
vollig idiomorph mit scharfen Hegrenzungsflachen, erreichen einen Durchschnitt bis 
zu 4 mm und ents{)rechen vermutlich der intratellurischen Olivingeneration des 
Ilauptgesteins. 

Der Feldspat in der Grundmasse der helleren Schlieren tritt in Stengein von 
ca. 0,14 mm Lange auf und verleiht durch seine Parallelorientierung dem Gestein 
eine Fluidalstruktur. Die Zusammensetzung desselben ist Ab46An54. Im iibrigen 
enthalt die Grundmasse Magnetit und Pyroxen. 

Die (irundmasse der dunkleren Partien unterscheidet sich von der obenge- 

^ L. c. p. 282. 



BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ-INSELN 3 1 

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 gelbgruner Farbe und weist parallele Ausloschung auf. Wegen der 
kleinen Dimensionen der Korner konnte eine sichere Bestimmung derselben nicht 
ausgefuhrt werden. Mit grosster Wahrscheinlichkeit liegt hier nur eine feinblattrige 
Ausbildung von Ilmenit vor. 

Basaltische Laven. 

Auf dem Uferplateau an der Ostseite der Insel bei dem Casastal steht ein 
feinkorniger 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 Ab4oAn6o 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 Grosse von 0,3 — 0,4 mm und sind etwas iddingsitumgewandelt. Von 
den Mineralien in der Grundmasse werden die Plagioklase am grossten, 0,08 mm. 
Der Pyroxen ist dagegen so klein, dass eine nahere Bestimmung sich nicht aus- 
fiihren Hess. 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 Vertikalzerkliiftung 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 FcgOg 
verursacht wird. Untenstehende Photographic (Fig. 9) stellt einen dieser iibrigens 
sehr sparlich vorkommenden Einsprenglinge dar. 

Die Grundmasse ist der des vorstehend erwahnten Olivinbasalts voUig gleich, 
nur etwas grober. 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 iibrigen findet 
sich Magnetit und der farblose Pyroxen. 



1 S. p. 276. 

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



32 



TOR H. HAGERMAN 



Der hochste Berg auf Masafuera ist der Inocentes. Von dem Gipfel, ca. 1 500 
m ii. d. M., wurde ein rotlicher, 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 Ciestein von Correspondencia. Dunklere Partien oben und unten 
sichtbar. — Vergr. 10 x. Photo. E. Dahlstrom. 



die hohen Interferenzfarben des OHvins, dazu weisen die Korner auch im iibrigen 
den Habitus des Oiivins auf. Bei Beobachtung in konvergentem polarisiertem 
Licht bei einem Achsenaustritt wurde keine Krummung des Achsenbalkens beob- 
achtet, vveshalb der Achsenvvinkel nahezu 90° sein durfte. Dies deutet auf cine 
fur liasahe normale Zusammensetzung von OHvin mit verhaltnismassig niedrigem 
Eisengehalt. 

Kand. S. L.\M)E1<(;kkn fuhrte eine Eisen- und Titanbestimmung der Ge- 
steinsarten aus, vvobci folgende Werte erhalten wurden: 18,82% FeaOg und 3,59 % 
TiOg. Dies entspricht einem Gehalt von 6,81% Ilmenit und 1 5,44 % Magnetit. 
Diese Zahlen sind natiirlich zu hoch, da Fe auch in den geringen Mengen Olivin, 
Pyroxen und Glasbasis enlhalten ist, die sich in dem Gestein befinden. Es ist 



BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ-INSELN 



33 



jedoch besonders interessant, dass, obgleich das Magma eisenubersattigt war, das 
Eisen die Konstitution des Olivins nicht nennenswert beeinflusste. 

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






6V\ 



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



Olivinkrystall einen breiten, vollig opaken, schwarzen Saum von Magneteismen 
besitzt». Das Phanomen stimmt mit dem vorliegenden voUkommen uberein, nur 
dass bier der entgegengesetzte Fall vorliegt, indem das ganze Korn voUstandig 
opak sein kann, auch in sehr diinnen Praparaten. 

Reiter2 hat sich mit der Zusammenschmelzung von Olivin und Magnetit 
beschaftigt und sagt: »Bei Abkuhlung 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-TuluI 
in Syrien. T. M. P. M. Bd. 7 (1886), p. 483—484. 

2 L. c. p. 232. 



34 



lOR H. HAGKRMAN 



Wir erhalten hiemit also Magnetite in zonarer Anordnung in den nachher 
entstehentlen Olivinen ein^esclilossen; ...» 

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

Unter den I^insprenglingen komnit, wenn auch selten, hier und da ein Feld- 
si)atindividiiiim \ or. dessen Zusamniensetzung ungefahr Ab4oAn6o ist; es handelt 
sich also um einen Labrador, unbedeutend abweichend von dem Feldspate der 
ubrigen (iesteinsarten. 

Die basischen Tlagioklase der Grundmasse zeigten zum Teil unscharfe Be- 
grenzungen. durcli die Anlagerung der diinneren Individuen verursacht. 

Die Pyroxene sind bedeutend kleiner und erreichen einen Durchschnitt von 
ca. 0,02—0.03 mm, wenn auch einzelne stengelige Individuen etwas grosser werden 












-0*^3^ 





r/iA 



Fi 



•'K- 



9. Kotj)igmentierte Olivineinsprenglinge. Basalt von El Ovalo. — Vergr. 56 
Photo. Hi. Olsson. 



konncn. \'on diesen kommen zvvei verschiedene Ausbildungen vor. Der eine (nor- 
male) ist mit den in den beschriebenen basaltischen Gesteinen auftretenden Py- 
roxen indentiscli und vermutlich als Diopsid zu betrachten, da er beinahe farblos 
und kaiun pleochroitisch bei einer Ausloschung von ca. 45° ist. Der andere Typus 
ist braimgelb und tritt besonders uin die Locher herum, niemals zusammen mit 
dem erstgenannten Tyjius atif. stellt aber wahrscheinlich nur eine Pigmentierung 
dcsselbeu dar. Auch hierin zeigt das Gestein eine grosse Analogic mit den vor- 
erwahnten. von Dos.s beschriebenen I^asalten. \\x sagt auf Seite 481: »Hier be- 
sitzen die in der Xahe der von Kalkspat ausgefiillten Hohlraume gelegenen Augite 
eine goldgelbc I'\-irbc; . . . . 

Im iibrigcn bcsteht die (irimdmasse aus kleinen, idiomorphen Magnetitkornern, 
Ilmenit luul Glasbasis. Der ganze (iebirgskamm soil aus diesem ausgesprochen 
basischen Gestein l;cstehen. 

I^ereits mikroskopisch zeigt ein anderes Handstuck, das vom Strandabhange 
nahe des Casatales abgeschlagen ist, eine auffallende Ahnlichkeit mit dem Ino- 



BEITRAGE ZUR GEOLOGIE DER JUAN FERNANDEZ-INSELN 35 

centesgestein. (Laut Angabe von Skottsberg soil das ganze ostliche Ufer aus 
diesem Gesteinstypus bestehen, abwechselnd mit den obenerwahnten Feldspat- 
basalten.) 

Auch hier treten magnetitfuhrende 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 Abkiihlung ausgesetzt gewesen ist als 
das vorgenannte, kann man dasselbe diesem gleichstellen und moglicherweise 
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 iiber die Veranderlichkeit 
der efifusiven 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 diirfte mit Riicksicht 
auf den geringen Umfang der Insel die umstehende Tabelle ein gutes Bild der 
Eruptionsfolge gevvahren, wobei die altesten Gesteine beim Meeresniveau, also 
links beginnen. 

Vor allem verstosst der stark basische Basalt aus der Hohe von 1500 m ii. 
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 konnen. 



36 



TOR H. HAGERMAN 



1 Natrontrachvte . . . 

! 




i ! o 

! i 1 














1 

X 










i 
1 





1 


1 






1 1 


O 


O 




1 




Basanit 
















1 

X 










j^^^^l^^FeUispat ^q i '\ 1 






















Ohvin- i oo 


O 


1 i 






















Er/iibersatti^ae Ha 
salte 





i 1 




















O 



13 14 1500 

m ii. d. M. 



Die \'erteilung der Effusivgesteine der InseJ Masafuera. 
O bezeichnet Das Material Skottsberg's. 
X » » , QUENSEL's. 



Ks ist selbstverstandlich denkbar, dass die in den tieferen Teilen des Zufluss- 
kanales versunkcnen schweren Magmas zuletzt herausgestossen wurden, doch spricht 
gcgcn diese Annahme das bereits in einer Hohe von 400 m ii. d. M. auftretende 
alkalisciie Ciestein. 

Scliliesslich ist aus der Tabelle zu ersehen, dass aus einer Hohe von 500 — 
1000 111 Liberhaupt kein Material untersucht ist. Die Masafuera Gesteine sind 
jcdenfalls in Minblick aiif den Mineralbestand so verschiedenartig, dass eine nahere 
L'ntcrsuchung der noch unbekannten Hohenlagen wichtige Aufschliisse ergeben 
durfte. 

/'»'j-/ ^7-intcd February 2^th, ig2^. 
Reprinted ivithout change April 13th, igjj4. 



3- Additional Comments on the Geology of the 
Juan Fernandez Islands. 



By 

PERCY OUENSEL. 



Contents. 

Page 

Introduction ^^ 

Main Geological Features 40 

Petrology of the Volcanic Formations 44 

Masatierra 4^ 

Masafuera 56 

Regional Relations. 

Tectonic Connections -74 

Petrographic Connections 77 

General Conclusions 79 

Acknowledgements 82 

Tables of Analyses 83 

Bibliography 86 



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 5 
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 CocilCRANE 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 Cmsoniana: "Lord 

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



3S IMKlV (JIKNSKL 

Cochcrane brought from the siiniiiiit (1.500 feet) a piece of black porous lava; 
and uiuler it lie foiiiul some dark iiardened clay full of cells, the inside of which 
appear sli^litly \ itritk-d. The ishuul seems chiefly composed of this porous lava; 
the strata of which, beini; crossed at ri^ht an^^les by a very compact black lava, 
dij) on tile eastern side of the island about 22' and on the west side l6°, pointing 
to the centre of tiie island as an apex" (l, j). 198). 

In 1S30 C\ Hi:kii:r<» j^ublished some observations under the title 'Notice 
sur rilistoire naturelle de file juan hV-rnandez'. With regard to geological ques- 
tions he sa\s: "je pense ciu'un geologue n'y trouverait cjue du basalte dans les 
etats, lueiue dans cehii de la j)lus parfaite decomposition; plusieurs blocs sont 
parsemes dune cristallisation j)articulicre, a laquelle on donne, je croix, le nom 
d()li\ine ... 11 n'\- a pas de trace de volcan ; les pierres qu'on prend pour 
de la hue, et dont (iuel(|ues-unes ressemblent assez aux scories on de la pierre 
j)once. ne sont, a mon avis, (jue du basalte decompose; on trouve aussi cette 
rociie sous forme sj)hericiue. et composee de couches concentriques ..." (2, 
p. 34;; compare in the latter respect h^ig. 10 on page 52 of this paper). 

.\. (Ai.ix i.Kic;!!, who accomj)anied Captain P. Parker Kixc; on the sur- 
\f\ing \()\ages of II. M.S. Adventure and Beagle on their first expedition 
1S26 1S30, 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 N'ear 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 carbonatee metastaticpie. It is further mentioned that the basalt in parts 
is almost colunuiar, and in others has a [)eaked 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 i)uhlishe(l in the Phil. Mag. and Annals of Philosophy, 
\'ol. l.\, I S3 I, }). 220). 

Captain King recaj)itulates Caldcleugh's narrative, as given above, with the 
addition: "In captain llAi.i.s interesting journal, there is a list of geological and 
mmeralogical specimens, of which one from Alasafuera is named vesicular lava" 
I4. p. 304). '1 he uhimate destiny of these specimens is unknown. 

Members of tlie Dumont dl'rville expedition, when visiting Masatierra in 
1S3S. collected and specified several different saiu})les of the lavas from the island 
'-^- P- ■'4'- 1 he material tor the new analyses of basalts from Masatierra, which 
L\(K<iix recentl\- caused to be made and which will be referred to later on, are 
evidentlx- trom this collection, as Lacroix says they were made from specimens 
collected by the Dumont dCrMlle exj)edition. G. (iUANCiK records some observa- 
tions as tollows: "Toutes les roclies aj)j)artiennent a diverses varictes de trapp 
et de diorite basaltique amorphes et vesiculaires, on ne trouve dans ces roches 
volcanicjues aucun autres mineraux (jue de I'olivine et de la chaux metastatique. 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 39 

La roche basaltique s'y presente le plus souvent en couches superposees, quel- 
quefois en escarpements interrompus et fractionncs et sont traverses par des dikes 
d'ejections plus modernes. Le basalt forme des pics eleves dans I'lle 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 iiber- 
all mit senkrechten Wanden, deren Hohe zwischen ico 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 Ki.iste, so dass man in diesen Stellen ohne Miihe 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 PLruptionen wiederholten sich haufig, und so floss eine Lavaschicht iiber 
die andere, um spater zum Teil iiber 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 hochsten 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 



I'KRrV (UF.NSKL 



the microscope tlie fiiiulaineiUal mass, in wiiich the plagioclase crystals are embedded^ 
has a dolerite structure. Tlie felspar crwstals show large extinction angles (38° — 41°) 
which ma\- be compared with tiiose of b\townite. The sections of the mineral 
are cracked and pervaded with zeolite matter, which forms an irregular network. 
This matter which looks slightlx' gre\-, when seen by ordinary light, remains 
obscured l)etween crossed nicols . . . The olivine, of which large sections are seen, 
is uniform!)- changed into a red hematite; these sections, however, still show 
extinctions like those of the unaltered olivine" (8, [). 176). The characteristic 
change of olivine to iddingsite in the basalts, which will be treated below in 
some detail, is here evidentl\- noted b)- Renard, though by him named hematite. 
I"'or the rest his description mostly refers to the colour and texture of differ- 
ent samples of tlolerite and basalt. 

In 1886 L. l).\K.\i'SK\ published a short report on detached rounded lumps 
of magnesite from Hahia del Padre in Masatierra, locally known as "piedras de 
campana' but named "(dockenstein" by the author (9). R. PoilLMANN 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 K. I'( »lll.M.\NN, participants in Fedkrico Joiiow's expedition to the islands 
in 1891, has been published by the latter as an introduction to Johow's monograph: 
'Ivstudios sobre la Flora de las islas de Juan Fernandez' (11, p. i). 

As a member of the Swedish Magellanian Expedition of 1907 — 1909 I had 
the opportunity, together with Professor Skott.SBERG, of visiting the islands in 
1908 and later described the rock specimens then collected in a paper: 'Die 
(ieologie der Juan I^'ernandezinseln' (12). In 1916 — 17 Skottsberg again visited the 
islands and collected specimens from some new localities. These were subsequently 
described by '1\ H.\(;krmax under the title 'Beitrage zur Geologie der Juan 
I'ernande/. Inseln' ( I 3). 

Since Skottsberg's visit in 1917, no further exploration in the field has been 
|)ublishe(l concerning the geology of the islands in question. And, with the exception 
ot tour recent analyses of lavas from Masatierra from collections made by members 
of Dumont d Frvillcs A'oyage au I'ole Sud et dans rOceanie' in 1838, commented 
on 1)\- L.\(k(»i\ (14, p. 64), no further observations have, as far as known, been 
j)ublishcd on the gecjlogy or petrography of the Juan Fernandez Islands. On the 
other hand (|uestions relating to their lithological connection with other intra- 
|)acific islands and their geophysical position in relation to eastern-pacific volcanic 
centres has been tlie subject of repeated discussions during later years. 

Main Geological Features. 

1 he two islands Masatierra and Masafuera j)resent very different aspects with 
regard to their bulk conti-uration. Masatierra exhibits a rugged appearance of 
isolated jagged cliffs lI"io. 21. The highest peak, VA Vunque, is 927 m. On the 
clear-cut sliore boundc-d escarpments one can distinguish many hundred lava 
beds overlying one another. In thickness they can vary from .some few metres 
up to 20 m or more. 



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




Fig. I. 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, 
sharply 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 i ooo m. 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 i 500 m. 

The sketch in Fig. i 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. BrCggen (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 precis est indetermine, 
mais il est certainement tertiaire, pleistocene ou meme, dans certains cas, recent" 

(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 F:xpedition 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 place 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 occurr-ed four times, causing much destruction, uproot- 
ing trees and drowning cattle. Shortly after the explosion, a large column, some- 



42 



I'F.KCN (JUKNSEL 




I■"iJ,^ 2. 'I'lu' south sIoijc of the high ridge along the western lialf of Masatierra as seen from 

Portozuelo. Photo C. Skottsberg. 



what rcscinblin<; a water-spotit, was seen ascending from the sea off point l^acalab, 
which proved to be smoke, but at 7 |).m. volcanic flames were visible through 
the smoke, which lasted till 2 a.m. on the 2ist. The depth of the water on the 
spot, where the eruj)ti()n took j)lace, was from 50 to 80 fathoms; no alteration 
in the dej)th was detected after the eruption had subsided" (17, p. 818). Sutclifife 
has published an account of the 'earthquake' in a separate publication (18) and 
reproduced a sketch of the sidj-marine eruj)tion (19, p. 387). 

It is obvious that these narratives must refer to a sub-marine volcanic explo- 
sion. Cii ARMS 1) \i<\\ IN also mentions the phenomenon in his 'Geological Observa- 
tions on the X'olcanic Ishmds' (20, p. 149). Renard gives the f)osition of the 
e.\|)losion as i I-jioHsh mile from the island and remarks "that the close prox- 
imit>- of a \olcanic centre seems therefore to be implied" (8, p. 176). 

1 he sid) marine eruption nuist be taken as conclusive evidence that the 
iuHuediate nei^hl)f)iMhood of the Juan I'Y-rnandez Islands has been the seat of 
volcamc action within the last 11; years. A point of further interest is that the 
e.xplosion was simultaneous with violent earthcjuakes on the Chilean coast, as 
Darwin already obserxcd (see j). 75). 

Hruggen refers to some finther observations of sub-marine eruptions in 
the vicinity oi' the Juan l-'ernandez Islands, recorded by Fk. Goll in his paper 
'J)ie lu-dbeben Chiles' (Munchener Geogr. Studien 1904, Nr. 14). The following 
denotements by GoM, are taken from Hriiggen (16, p. 332): 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 43 




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



44 PERrv (jrHNSKi, 

"'Se«4un CioW se prodiijo", cl 12 de febrero de 1839 una erupcion submarina 
\- marenioto a unas 120 kni al este de la Isla de ^las a Tierra. rLl mismo autor 
cita la obscrvacion si<;iiicnte hecha en un punto iin poco mas austral: 'l^^n octubre 
de 1867, se sintio un temblor submarino en 34 55' S y 7/° 38' W (unas lOO 
millas SIC de Juan I-'ernande/.V despues el buque navego durante dos boras por 
agua tie color bianco lechoso. habiendo mucho pescado muerto en la superficie,' 
Se trata probablemente del mismo fencHiieno que describe JoSK M. Po.MAR en 
la forma sigiiiente, aun(]ue dice (jue el [)unto se halla a 100 millas al S\V en 
\ ez de Si'', de Juan I-'ernantlez: 'ICn 1867, el capitan SlMPSON de la barca britanica 
C'oronella navegaba en el I'acitico con mucha calma y vientos contrarios, con 
e.\cej)ci()n de un hierte \ iento acompaiiado de siete temblores que se produjeron 
como a 100 millas al SW de la Isla de Juan Fernandez; durante dos horas navego 
|)or agua tan blanca como leche; sondeo, pero no toco fondo en 100 pies de 
|)rorim(latl, \ io muclios pescados muertos y una gran cantidad de pajaros por 
todas j)artes. Agregaba el capitan Simpson que si hubiera estado 10 millas mas 
adelante, el cluxiiie hubiera sido peor y hubiera causado averias al buque.'" 

Concerning the earthquake of Vallenar in the province of Atacama on the 
loth of Nov. 1922 Ikuggen cites I^AII.EV WlLUS' observations on the contem- 
poraneous volcanic activity on San Felix. 

I^ruggen concludes his opinion on the submarine explosions, given above, 
as follows: "A la teoria del origin de los tsunamis por erupciones submarinas podria 
objetarse (jue serian erupciones muy excepcionales, ya que consisterian en una sola 
o nuiy pocas explosiones que causan las pocas olas sismicas, apagandose luego 
la actividad. V.n 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 
suhciente para causar el tsunami. Tambien 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 submarines 
parecen estar en relacicm mas estrecha con los focos sismicos de los grandes 
terremotos chilenos" (16, p. 332). 

Petrology of the Volcanic Formations. 

The following descrij)tion of the rocks of Masatierra and Masafuera is based 
on specimens collected by myself in 1908 and by Skottsbkrc; in 191 7. 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. 

1 he specimens from Masatierra in both collections give conclusive evidence 
that this inland 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 ot oJiMiie. The specimens from Masafuera on the other hand indicate 
more obx lou-, dissimilarities in the com[)osition of the rocks at different levels. 

In the following the principal petrological features of the two islands will 
first be treated. I'lHlcr 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° 37i' 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 {c(. 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 1 50 m above sea 



' The name oceanite was first proposed by Lacroix in 1923 (in Mineralogie de Madagascar, 

Vol. Ill, p. 49). The name is given "k cause de leur abondance dans les iles du Pac-hque". 

^ Holmes, Q. J. Geol. Soc, 172, 1916, p, 231; Washington, Am. J. of Sc. V, 1923. p. 471- 



rKKCV (llKN'SEl, 








ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 47 




Fig. 5. Basaltic lava beds. Western head of Bahia Cumberland, Masatierra. Photo C. Skottsberg. 



level. The two speciinens 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, 191 1, 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 an}^ signs of even 



48 



PF.RCV QU F.N SEL 






^ 



M^^m^'- 



^;^ 



v!i>;?r: 



> 



Fij,''. 6. Ficritc basalt oceanite). x 9. Puerto Frances, Masatierra. 




ig. 7- I ><'l('riti< basalt, x ^y. Bahia Cumberland, Masatierra. 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEIZ 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, llmenite 
in tabular form or in skeleton cry.stals 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. llmenite 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 500 m). 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 (i5' — 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 in a 
varying content of olivine. 



so 



rKKcV (UENSKL 




I'"ij(. H a. X'esirular aphyric feldspar basalt cokminar structure . 

(see Fig-. 9 . 



30. Portezuelo, Masatierra 




Fig. <S 1). Same spe( inien as Fig. 8 a, niagiiified x 65. Vesicle in upper right corner filled with 

o})al. Portezuelo, Masatierra, 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 5 I 




Fig 9 Northern wall of the Portezuelo Pass, showing columnar structure of feldspar basalt 

CFig. 8 a). Photo C. Skottsberg. 



PERCY QUKNSEL 




li-. 



uial weathering of basalt. Bahia Cumberland, Masaticrra. Photo C. Skotisbci^;. 



Oiiantities of pyroclastic material are found interbedded between the lava 
Hows all over the island. Pohlmann has described such products from Bahia 
del Padre as follows: "Uberall sieht man Decken von basaltischem, meist saulen- 
formi^ abj^esondertcm P'.ruptiv^estein mit Schichten von vulkanischem Auswurfs- 
niaterial wie Asche, Lapilli, l^omben u. s. w. wechsellagernd. 

Mine solchc aus Lapilli und l^omben zusammengesetzte Schicht von mehreren 
Mctern Maclitij^keit an der rechten Seite der Bucht in der Nahe der sog. Kapelle 
<;elc^fen, liefcrt das Material der sog. 'Glockensteine', Der Vorgang ihrer l^ildung 
ist, kurz gcsagt, folgendcr: zuniichst entstehen traubige und nierformige Con- 
crctioncn zwischcn den losen zusammengefiigten vulkanischen Massen; dieseweissen 
Knollen (JMg. i 2) gelangen beini Abstiirzen der Schichten an den Strand und erhalten 
als Kollstcinc (lurch die Thiitigkeit des Wassers ihre gerundete P'orm" (lO, p. 321). 

I ha\e prc\i<)us]y noted that the lava beds of the eastern parts of the island 
are also often interbedded with agglomeratic layers (12, p. 257). 

Some stra>' occurrences of superficial tuffs seem still to have evaded destruction 
by erosion. Wlu-n Kcnard says that "amongst the specimens collected at Juan 
I'"cnian(iez (Masaticrra) !)>• the Ciiallenger l^x|)edition in 1 875, we have not found 
any specimen which iniglit belong to any recent eruption, no tuffs, no volcanic 
ash are to be found and e\cr\thing seems to prove that they have been washed 
away by the waves and the atmosj)heric agencies" (8, j). 176), this last conclusion 
seems (juestionable. 1 lagerman refers to two specimens in Skottsberg's collections 



ADDITIONAL COMMENTS ON THE OEOLOGY OF THE JUAN FERNANDEZ ISLANDS 53 

as representing recent tuffacious material. The one he describes as "ein poroses, 
dichtes Gestein von El Puente, dem Istmus zwischen der Padrebucht und Carbajal 
und ist ein Palagonittuff mit einigen sporadischen Augit- und Magnetitkornern. 
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 tuffaccous origin. The 
description was given as follows: "In den zentralen Teilen von Cumberland Hay 
liegen noch ziemlich machtige, meist lebhaft rot gefarbte Tufflager, die sehr stark 
umgewandelt sind. Bruchstucke von Olivinkristallen, lapilliartige Lavabruchstucke, 
P>zkorner und Glas liegen in einer Grundmasse, die aus einer weichen, mit Mes.ser 
schneidbaren, roten lateritahnlichen Lehmsubstanz besteht, die durch Verwitterung 
aus dem ursprlinglichen Tuffmaterial hervorgegangen sein durfte. 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 jeolojica 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. P^n ningun otro punto, ni en Masatierra, ni en Masafuera 
se ha observado una configuracion jeolojica analoga a la mencionada" (11, p. 4). 

The rocks referred to by Pohlmann certainly show a divergent aspect, l^ut 
I do not think they can be taken as represehting 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 

' Ur. Schulze died before publishing his observations. 

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



54 



\'\\<C\ (UKNSKI. 




I \'<.|. 



llumciatc. I'.aliia CumixTland, Masalicrra. Photo C. Skottsbcrg. 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 55 




Fig. 12. Lumps of magnesite ;the so-called Glockenstein . Nat. size. Rahia del Padre, Masatierra. 
Reproduced from P. Quensel, 12, p. 269. 




Fig. 13. Scoriaceous lava with newtonite and bole. Nat. size. Bahia del Padre, Masatierra. 
Reproduced from P. Quensel, 12, p. 269. 



will in this connection only summarize the facts. The rocks in question are very 
fine-grained and of a greenish colour. The abundant cleavage plains are coated 
with epidote and calcite. Pyrite is finely dispersed throughout the rock. Under 
the microscope one can see that original phenocrysts of both olivine and augite 
are more or less completely changed to serpentine and chlorite. The matrix con- 
sists of olivine, augite, plagioclase, magnetite and ilmenite — i.e., the usual 
mineral assemblage of the basaltic lavas. The greater part of the silicates are. 



I'KRiV (JlKNSKL 



li()uc\cr, \cr\- nuich altered to serpentine, chlorite and prehnite. Secondary albite 
and some scapolite is also j)resent toj^ether with abnndant pyrite. Small cavities 
are tilled with epidote ami calcite. in the slaj^i^y lavas in the immediate vicinity 
lai\L;er ca\ ities are tilled with a j)iirel\- white cla>--like substance which was found 
to be newtonite. In intimate association with this mineral is nearly always found 
a vellowi^ii waw substance which shows ever\' resemblance to what mineralogi- 
call\- ma\- be si^nitied as bole (I''i<;. 13). It is very brittle with a conchoidal 
fraclme. in water it readil\' disintegrates into small angular fragments. Tliis bole 
mineral is also t'ound tilling cracks, or occurs in smaller masses in the cavities 
of the basalts around the baw 

in t!ie agglomeritic lavas of the vicinit)- large cavities are filled with hard 
compact magnesite. As earl\- as 1886 Darapsky described this mineral by the 
r.ame "( ilockenstein" and gave an analysis thereof, which shows it to be an 
exceptionall)- pure magnesite (9, p. I 13). Without doubt the magnesite is primarily 
derued tVom olivine, the decomposition being caused by the same processes as 
ha\e changed the basaltic lavas nearby. 

The whole asj^ect of the rocks from l^ahia del Padre, with magnesite, calcite, 
serpentine, chlorite, sca{)olite and pyrite as secondary minerals, seems without 
doubt to indicate that the lava beds in question 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 Pohlmann. 

In a s[)ecimen collected by Skottsberg from the shore south of El Yunque, 
Ilagerman also tound evidence of a far-reaching decomposition. Under the heading 
"! Ixclrothermale I^ildungen" he gives the following description: "Das Handstiick 
ist ein von weissen .Streifen durchzogenes scharfgriines Gestein, das u. d. M. 
grosse Augitkristalle in einer vollig zeri^ossenen Serpentinmasse zeigt. Das Pra- 
))arat ist von Aragonitbandern durchzogen. Dieses Gestein muss als ein stark 
umgebildetes ( )livinfels bezeichnct werdcn" (13, p. 27). The large olivine crystals of 
the duniti'. tound as inclusions in the picrite basalt at Puerto Prances, show, on the 
contrarx-, no signs of secondar\- alteration (see p. 47). There is therefore no doubt 
that the higiily decomposed '( )hvinfels' described by Hagerman has succumbed 
to a later decomposition of much the same nature as has been active around 
I)ahia (K-1 I'adre. 



Masafuera. 

I. at. ^y 52' S.. Lon^. <So'' 54' W. : 

1 he la\as ot Masafuera piesent a more varying aspect than those of Masa- 
tierra and contam .sc\cral t>pes of petrographic interest. They have, however, 
hitherto only been sinumanl\- described in the j)a|)ers by myself (l 2, p. 274) and 
b\' I lagerman 1 1 3. p. j8i. 

I he rocks which predominate at lower levels are mostly vesicular to slaggy 
basalts. The\- are well represented around the Ouebrada 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 57 



QwebradxvNegrcb 
Q Saitdahto 
Q SandaJLo 

QuiebraSa. PcUo 



Chx>%<ta 
QiLeibrajdcLiiano 




Ra3a de-la, CoLonicL 
Casae o Banuedxxnjo 
Ctuca. 

Q.BUndaSo 



Q Cdbreros 



P*«-Negra. 



Cables 10 

(§065 f*')  



3 SeaMiles 



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



S8 



)r F.N SKI. 





Fig. 15. \- 



.t !li 



nicrior of Quel^rada do la Loberia vieja towards the shore. 
Photo C. SkottsberL--. 



of Masatierra, olivine is more scarce in the dominant "basalts of Masafuera, but 
is sin(^tilarly rich in the basaltic dikes which in hundredfold vertically traverse 
the island. 

The rocks |)re\ailin<^ at heights up to c. i ooo m may for the greater part 
be classihed 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 (V\i^. 16). 

In many sj)ecimcns of the vesicular lavas the vesicles are filled or lined with 
zeolite minerals. In some samples the \esicles present, from the outward rim, 
fust a coating ot glass, followed by chalcedon and chabasite and a central 
replenishment of well develoj)ed natrolite spherolites. 

At an elevatif)n of about i ooo m a lava bed of a \ery ditTerent aspect is 
met witii. In contrast to the dark basalts of lower horizons, the rock now in 
(jiiestion is light gre\- in colour. Large j)henocrysts of olivine (up to 5 mm in 
diameter) and labrad'jrite are tmiformly distributed in a very fine-grained ground- 
mass, consistmg 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 asj:)ect. The surface 
teels rough and grain\- which, together with the light grey colour, at first gives 
the impression that the rock would ha\e a trachytic or trachy-andesitic compo- 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 59 




Fig. 16. Vesicular feldspar basalt, x 8. Normal development under i 000 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 exprimee; elle est restee a I'etat potential dans la verre. 
La poudre des ces roches, traitee par HCl s'attaque plus au moins facilement" 
(C. R. 169, 1919, p. 402). Bartii 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; 
22, 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 basanilc, the 
calculated norm shows no nepheline and an excess of 1.99 % of aluminium. Ihis, 
together with a content of 3.36 % H2O, gives the impression tiiat the rock is de- 
composed. On this account Lacroix, in reproducing my anal jos ■:: ilie Juan 
Fernandez rocks, omitted this analysis, "car les resultats ne curresi- nclcr.t pas 



6o 



rF.RCV ()L-F.NSKL 




17. ( )li\ inc h; 



lit. Nat. size. Elevation c. i 000 in. Masafuera. The olivine plienocrysts more 
or less completeh- altered to deuteric iddingsite. 



:i la composition mineralooique clccrite; le calcul fait voir qu'il s'a^^it d'une roche 
altcrce" ( 14, |). 65). 

A reiiewetl scriitin}- of the rock has offered new aspects regardin<^ its petro- 
looic and j^etro^raphic position. The essential point has been that the large olivine 
phenocrwsts are altered to such a great extent to iddingsite. We have every inter- 
mediate phase frotii a more or less broad rim of iddingsite around a kernel of per- 
fectly fresh ()li\ ine to c()mj)lete pseudomorphs of iddingsite, retaining the crystal 
habit of the olivine. The circimistancc 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 
la\a in which it occiu's. 

l)iM'ing later }'ears sexeral papers have been published, in which the origin 
ol iddingsite has been discussed in detail. The conceptions, there j:)ut forth, to 
all intent conform with the characteristic features of the lava from iMasafuera. 
As the petrogcnesis of the rock in (juestion is of a certain interest, some signi- 
ficant (|uotatioiis may be given, relating to the formation of iddingsite from 
ditk-rrnt localities. 

I\i»ss and .Shannon simimarize their conclusions as follows: "Iddingsite is 
not confmed to weathered surfaces; its development shows no proximity to joint 
cracks and e\ idences 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, tiiat is to say the result of metasomatic {)rocesses associated 
with the later stages (jf a cooling magma." They also emphasize that the 
magma must ha\e come to rest before iddingsite formed, for though it is a 
very brittle mineral, it is ne\er fractured or distorted by flow (C S. National 
Museum, l*roc. 6-, 1925, Art. 7, p. i<S). 

Al Korssi.Ai comes to the conclusion that "iddingsite is the result of oxida- 
tion [)rocesses that acted rapidly on the olivine during the liberation of copious 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 6 1 

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

Finally Edwards in his paper 'The formation of iddin^site' 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, j). 281). 

The formula for iddingsite is given by Ross and Shannon as MgO • FegOa • 
3 Si02-4 H2O with magnesia replaced in part by CaO, approximately in the ratio 
I : 4. The calculated composition according to this formula includes 7.90 % HaO"*" 
and 7.90 % HaO" (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 PI. 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% H2O, we may conclude that an appreciable amount of the 3.46% H2O, 
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 Otto 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 ir the iddingsite 



VFAiCX ()rKNSKI. 




l•'i;^^ i8. hklinj^site forming sharp outer borders around unaltered oli\inc. < loo. From specimen 
Fig. 17. Elevation i 000 m. Masafuera. 



aV, ^*,.. -< 




fy— 



r 



^^f^ JSr^^. 



y 



Fij^. 19. I'henocryst of labradorite, traversed In' a zeolitic matter, x 170. From specimen 

I-"i<^. 17. Masafuera. 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 65 

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 w^eathering or 
later thermal activity. 

In consequence of the changes in chemical composition which must have 
taken place in connection wTth 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 ooo and i lOO 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 higluM' elevations 
were reached. New specimens in Skottsberg's later collection di^::o\ e my earlier 
supposition in two essential respects. In the first place, rock.-, of mi ch the same 



^'4 



I'KIUN ()rKNSEL 




Fig. 20. Sotla trachyte. :■: 50. Elevation c. I 050 m. Masafuera. 




Iiytc boulder. • 30. Xic. +. Elevation about 400 m. 
Ticrras lUancas, Masafuera. 

character as the .soihi trachyte have now been found at lower levels. Secondly, 
Skottsbet-o reached the .siniiniit of the i.sland and was there able to establish that 
basaltic lavas ()cciij)y tlie whole highest i)art 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 Ticrras Blancas. According to Skotts- 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 65 




F\g. 22. Vesicular phyric feldspar basalt with stray phenocrysts of olivine, 
cimen from i 420 m elevation. Masafuera. 



30, Analysed spe- 




ij. Aphyric feldspar basalt. 

El Ovalo, Masafuera. 



66 \'V\iC\ (JUKNSKL 

berg's intorniation, boulders of tliis character were ver}' numerous in the talus 
under a steep escar|)nient and can here hardlx' ha\e originated from higher eleva- 
tions tlian about 400 ni. 

In outwarti a[)|)earance this rock shows an aspect different to the compact 
soda tiachxte i)et\\een I 000 and i loom, which on account of its dense texture 
at first sii;ht i^ives the impression of a (piart/itic rock. The new specimen has 
on the other hand a roui^h. loose grained structure. Minute inter^jjranular cavities 
in the rock are seen under the microscope to be tilled with limonitic matter. But 
allowing for tiiis structural dissimilarity the rocks from the talus below Tierras 
Hlancas seem to l)e closel}' related to the soda-traclu'te as described above (Fig. 21). 
Taking into consideration the seemingl}- horizontal j)()sition of the lava beds of 
-Masafuera. the conclusion must be that la\as of soda-trachytic composition have 
been emittetl at different times and that they have alternated with lavas of more 
normal basaltic comj)()siti()n (see j). 71). 

.S|)ecimens of the rocks from between i 100 and i 400 m j)resent again a 
different asj^ect. I'^ive sam|)les from I lOO, I 200, I 300 and I 420 m all have 
\er\- much the same a})pearance. The}' are all phyric lavas of an ash-grey colour, 
i-'eldspar and, more rarel}-, olivine occur as ])henocrysts 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 feldsjjar circuiting the phenocrysts in a more or less well-defined 
manner (I'^ig. 22). 

In m\- previous paj)er I named the rocks of this nature trachyandesite. 
Ilagerman sa\-s, referring to much the same specimens in Skottsberg's collection: 
"Die Klassifi/ierung hiehergehdriger Gesteine ist etwas unsicher. Geniigende 
Grimde sie als Trach\-andesite zu bezeichnen, liegen jedoch nicht vor" (13, p. 30). 
I iagerman names the rock andesite in his j)aper. 

Tliese rocks evidently have a rather wide distribution on ^lasafuera, j^robably 
lorming the whole comi)lex of la\a beds between i 100 and i 420 m. To certify 
their j)etrographic ])osition an analysis has now been made of a typical specimen 
( Tabk- I, No. 5). The calculated norm and Xiggli values definitely classify the 
rock as a tel(lsj)ar basalt, in many respects of similar composition as some of 
tile analxsed basalts from Masatierra (Table I, \o. i — 4). A difference of interest 
is the still lower content of KoO; the Xiggli value X' is now 0.08 against O.I 5 — 0.20 
in the basalts fiom Masatierra. 

I hese phyric feldspar basalts from ^lasafuera in general aspect also very 
much resemble the apii\ric fel(lsj)ar basalts from between 400 — 600 m elevation 
on Masatierra. ( )ccasionall\- at lower levels they also on Masafuera may be aphyric 
in texture il'ig. 23). dhey seem also in several respects to correspond to certain 
ph\ric and aph\ric basalts from the I lawaiian Islands, as described by Washinotox 
(Petrology of the I lawaiian Islands. I. Kohala and Mauna Kea. Am. J. of Sc. V, 
'9-3' P- 4''^"*- ^ '> this (juestion I will return under a concluding heading dealing 
with the chemical and petrological connections between the rock of the Juan 
I-'ernandez Islands and other volcanic islands of the eastern Pacific. 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 67 




Fig. 



•4. Olivine basalt, supersaturated with iron oxide, x 35. Vesicular flow-breccia. Los Inocentes 
^elevation c. i 500 m), Masafuera. 




Fig. 25. Olivine phenocryst from specimen Fig. 24. x no. Specks and 
olivine as rests in a pseudomorph after olivine. Los Inocentes. 



cak^ <:i tUKiiLcred 



6S 



PKKCV ()1-i:nski. 




I"ij4. 26. Disintegrated phenocryst of olivine, caused by high temperature oxidation. The rim 

around the olivine crystal is hematite. Within the olivine the light grains are mineral components, 

formed by exsolution. < 150. Los Inocentes, Masafuera. Photo P. Ramdohr. 




Pig. 27. I'art of disintegrated olivine |)heno( ryst, enlarged x 600. Against the dark background 

of olivine the exsolution proiUn t is seen in the form of small composite grains consisting of 

hematite light and a darker undefined component. Larger lij^ht grains see PI. II, P^ig. I. 

Photo P. Ramdohr. 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 69 




Fig. 28. High-temperature exsolution of spinel, enclosed in olivine phenocryst (PI. I, Fig. 2). Light 
lamellae, according to Ramdohr probably magnetite in groundmass of excess spinel, x 600. 

Photo P. Ramdohr. 




Fig. 29. Magnetite in groundmass of the lava bed as seen in PI. I, Fig. i pa:t-a!iy c!):inge(l to 
hematite (martite). x 300. Photo P. Ramdohr, 

5 — 516795 The Nat. Hist, of Juan Fernandez and Easter Isl. Vol. I. 



■JO I'KKCN (JIKN'SKI. 

An observation of importance with rej^ard to the distribution of different 
lava beds on Masafuera is tlie find of a deviatini^ type of olivine basalt from Los 
Inocentes. Accordin*; to Skottsber^'s observations such la\as probably occupy 
the hii^diest part of the island, representing^ elevations above I 420 m. The few 
samples brouj^ht back are hi<4hl\' scoriaceous flow breccias containing a high 
content of iron oxides (h'ig. 24). Hagerman has in the preceding publication 
of this series given a descrij)tion of these lavas which he characterizes as slaggy 
ohvine basalts with large olivine phenocrysts, supersaturated with magnetite 

(•3. V- ^S'^- 

'l"he phenocrysts of olivine are under the microscope found to be almost 
opacjiie, 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 (l^'ig. 25). \o 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. Gavklin and Dr Uv'i"KNlM)GAARl)'r kindly undertook to examine some 
polished sections of the rock. Professor P. Ramdoiir (Heidelberg) contempora- 
neously supervised a section for the same reason. It thereby became apparent 
that the seemingly opacpie constituent was not magnetite and that the mineral 
assemblage of the pseudomorphs was of a complicated nature. Professor Ramdohr 
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 i(liom()r|)hic 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. Repeated attempts have been made to determine the second com- 
ponent both in polished sections and with X-ray powder photographs. No con- 
clusive evidence 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 saiue processes as have controlled the formation of the deuteric 
iddingsite, though in the samj)lcs at hand this mineral is not extant. P:dwards 
seems to have described a very similar formation in the iddingsite-bearing basalts 
from two X'ictorian localities in Australia. After concluding that the iddingsite 
must have been formed before the ultimate consolidation of the lava flow, Fdwards 
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. Fventually all the original 
olivine vanishes, and the iddingsite, which had formed a rim about it, is completely 
rej)laced 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 7 I 

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 X 100 



o I 2 3 4 5 6 7 8 91011121314 1500 



Olivine basalt supersaturated 
with magnetite 

Light grey phyric feldspar 
basalt 

Soda-trachyte 

Iddingsite-bearing phyric oli- 
vine basalt 

Dark phyric and aphyric 
feldspar- and olivine basalt 

5*— 516795 



























X 


X 


X 










X 












X 


X 








X 


X 


v 


X 

























I'F.KfV (JUFNSF.L 




Fi^^ 30. Picrite basalt (masafuerite;. Dike rock. Nat. size. Loberia vieja, Masafuera. 

} Ia<^erman 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 sotne slight corrections on the base of renewed examination. 

My former supposition that the basaltic lavas only occur up to an elevation 
of about I 000 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 VV^est-East 
direction, are worth special notice as representing rocks exceptionally rich in 
olivine. In this res})ect they exceed the most olivine-rich picrite basalts from 
Puerto I'Vances on Masatierra. l^OWKN has commented on these rocks as follows: 
"( )ne otiier rock may be mentioned in this connection. It is a picrite basalt from 
Juan PY'rnandc/., 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 olivmc 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 73 




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




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



74 



PERCY (UKNSEL 



fact does not prove that they were not in solution in that material at an earlier 
time. Hut if one finds the condition siiown 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, thc\' always contain either all of their olivine or all in excess 
of a (juite small amount (apparently some 12 — 15 %) as relatively large pheno- 
crysts. rhe\-, therefore, force acceptance of the stated conclusion" (23, p. 163). 
The (juotation above refers to purely theoretical questions but indicates the extreme 
j)osition these rocks hold in petrographic classification with regard to the abnor- 
mally high content of olivine in basaltic lavas. JoilANNSEN 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 h'rances, 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 peridot; dans I'echantillon analyse, tons les grains 
de ce mineral se touchent, ils sont reunis par une petite quantite de plagues de 
labrador, englobant des microlites d'augite et des lames d'ilmenite" (14, p. 44). 
Johann.sen 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 
upj)er 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, subse(juently consf)lidated 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 ciiannels 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 crystalHsation in accordance with l^owen's conception of these 
dikes as (jucnched 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 75 

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- 
pagos" (20, p. 144). 

In his paper 'Constitution lithologique des lies volcaniques de la Polynesie 
Australe' Lacroix gives expression to much the same trend of thought when 
he writes: "Particulierement interessantes sont les iles volcaniques qui se trouvent 
a une plus ou moins distance de I'Amerique du Sud, les iles Juan Fernandez, 
San Felix et San Ambrosio, et enfin Galapagos, puis au large de I'Amerique 
centrale, I'ile Clipperton. Bien que la connaissance de la lithologie de ces iles 
soit loin d'etre completement eclaircie, on peut a present assurer que leur laves 
different de celles des Cordilleres des Andes, c'est-a-dire de la serie circumpacifique 
et montrer qu'elles se rattachent a la serie 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 signale, au large 
de Valparaiso 5.651 metres. A I'ouest de ces fosses sont situees les deux iles 
volcaniques anciennes de San Felix et San Ambrosio; au Sud de ces iles le 
croiseur Chilien Presidente Pinto a trouve, sur une etendue de 760 km, des 
profondeurs si faibles qu'il est probable qu'une crete sous-marine, orientee a 
peu pres N — S, s'allonge dans la direction de I'ile Juan Fernandez" (25, III, 

P-I359)- 

In his 'Description and Geology' of San Felix and San Ambrosio, Bailev 
Willis writes: "San Felix and San Ambrosio are volcanic islands in the South 
Pacific Ocean, San Felix being situated in latitude 26°: 5' south and longitude 80°;' 
west of Greenwich and San Ambrosio lying about 16 km to the east-south-east. 
They are about 500 miles west of Chanaral 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 ol 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 



PEKCN OlENSEI. 



can be made. The deptli of the ocean in this region, which lies west of the 
Ricliartls Deej), varies from 4.000 to 5.000 meters. The islands, therefore, repre- 
sent tlie 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 paf)er on the geology of Galapagos, Cocos and Easter 
Island, L. J. CliriUi has j)ublished some noteworthy remarks on the regional 
relations of the \olcanic islands of the eastern central Pacific. He writes as 
follows: "L'nder 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. Xo 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 Martjuesas 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, Piaster, Sala-y-Gomez 
and the Juan I^'ernandez 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 to a 
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. Vd'ix 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. 
\'olcanoes 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 j)roved by the history of their coral reefs. 
I here is not sufficient evidence, however, to determine whether the folds which 
l)r()bably underlie Cocos, St. Felix and St. Ambrose islands have suffered a 
similar movement ' (27, |). 43). 

Ji .\\ Hiu(;(;k\ has recently in his book 'Fundamentos de la Geologia de 
Chile' discussed the geotectonic position of the Juan P'ernandez Islands. He 
writes: "klsta zona (la regicui situada al este de Llico, en Arauco) de dislocaciones 
tan extranas a la structura de la Cordillera de los Andes, coincide con la region 
donde una ancha loma submarina se desprende del continente. F:ncima de la 
loma se levantan las islas Juan I^'ernandez y mas al norte las de San Ambrosio 
y San Felix. 

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



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 77 

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 termino de las capas de la Quiriquina en el sur de Arauco y tambien 
el hecho que sedimentos marinos de Eoceno no se conozcan mas a sur. 

Pero en el Oligoceno, cuando la costa del piso de Navidad se extendio 
hasta la region de Ypiin (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 immigro la flora del Eoceno. Cuando mas tarde se hundio tambien 
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 (occanites 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, therelore. 



7 8 I'KRCV (jrKNSRl. 

possible tluit earlier, lowermost flows are less sodic and more typically basaltic 
than the upper, which were the ones examined. " Washington continues: "In 
this {:)redominantly highly sodic character of the lavas San Felix appears to 
differ widel\' from other Pacific islands. At IMasafuera, it is true, both soda 
trach\-te (and nephcline basanite)', closely like those of San Felix, occur, but 
these are accompanied b\' basalt and [)icrite basalt, whereas at the neighbouring 
juan b'ernandez (Masatierra) the lavas appear to be, to judge from Quensel's 
description, only olivine basalt with neither trachyte nor basanite. Trachyte, also 
highl\' 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- 
canitjues de la Polynesie Australe' Lacroix compares the petrographic character 
ot 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 I'analogie la plus grande avec les basaltes de Masatierra et avec 
ceux des iles (iambier, c'est-a-dire avec les plus calciques des basaltes du 
I'acificjue et les plus pauvres en potasse" (14, p. 68). And further on in the same 
paper: "Les iles Juan P'ernandez se groupent au voisinage des iles Marquises et 
1 on a vu (ju'il faut comparer ce que Ton sait des roches des iles Galapagos aux 
donnces concernant ceux des iles Gambier" (14, p. jy]. 

'1 hese 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 
ot the Gambier Islands, however, goes a step further, as Lacroix describes from 
tiiere, 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 reterence to chemical similarities with other volcanic islands of the Pacific is 
Conrad Hi kki. Under the title 'Chemismus und provinziale Verhaltnisse der 
jungcruptivcn (iesteine 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 I-'ernandez, San P'elix 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 etudiees presentent la commune caracteristique d'etre 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 k is 
always under 0.4 and for the most typical regions of volcanic rocks of the intra- 
Pacific Ocean under 0.25 (28, p. 173). 

The value k 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 K2O (Niggli value k 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 
Gambler 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 ']']\ 

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. 



8o PKRCV QUENSEL 

soda-trachyte from Maui and Analuilu, on the first island in connection with a 
'picritic basalt' [V. S. (ieol. Survey, Prof. Paper 88, 191 5, p. 26 — 28). This corre- 
sponds exactly to the rock assenibhif^e of Masafuera. 

Petro(^ra{)hic 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 [)henocr)'sts 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" (1. c, p. 485). This description might as well refer to the light 
grey basalts of Masafuera at elevations between i lOO and i 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 I'ernandez, 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 com[)ly with Lacroix when he says: "Les roches de cette ile 
(Hawaii) constitueraient une division speciale, ayant une originalite propre" 
(14, p. 76). 

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 
(kila|)agos Archii)elago display the closest similarities. According to the analyses, 
j)ublished by Richardson, both the basalts and the soda-trachyte are in chemical 
composition very similar to equivalent rocks of Masafuera. Also soda-trachyte is 
f)f 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 T'ernandez 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 ' 1 27, j). 64). The low content of potassium is in common for the 
basalts from both island grouj)s. 

On the other hand we must evidently exclude any petrographical relationship 
between Juan I^'ernandez and San P'elix — San Ambrosio where the lavas have, as 
far as is known, a more j)ronounced alkaline composition, classified by Washington 
as soda-trachytes and nc))hcline basanites (1. c, p. 382). Richardson says: "The 
Juan I<ernandcz are i)etrologically closer to the Galapagos than are any other 
islands . . . Hoth are situated comparatively near the American coast of the Pacific, 
but their similarity is not shared by San VeWx 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 8 1 

to be the highly alkahne rocks of Tahiti in the mid-Pacific (14, p. ']']). 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 P^rnandez 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 Gambler 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. ']6) 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. 0° 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 serie nephelinique" 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-bearirig types and the presence 
of virtual free silica in many. Lacroix unites them as belonging to "la serie 
sans nepheline" (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 vulcanic 
nature. On San Felix volcanic gases were issuing from a crevice on the southern 



82 I'KRCv qiens?:l 

rim in May 1923. l^aile\' Willis sa\'s: "In this sense and to this extent we 
may consider San VcWx 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, {)()urin^ over the 1 00 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 eartlujuakes 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 
petrolog\- 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, Petr()gra{)hy of the Pacific Islands. Bull. Geol. Soc. of America, 27, 1916, 
P- 331)- 

Acknowledgements . 

The writer is indebted to Professor P. Ramdoiir, Heidelberg, and to Pro- 
fessor S. (kWKI.ix and Dr W. Uytenp.ogaardt 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 (). Mki.i.is has helpfully co-operated in taking most of the other photo- 
micrographs. The landsca[)e j)hotographs have kindly been put at my disposal by 
Professor C". Skoi 1 si!KR(;. They are taken by him on his visits to the Juan 
i'ernandez Islands in 1908 and 1917. 

A grant from the foundation Lars Mikrtas Mixnk is gratefully acknowledged. 

.Mineralogical Department, University of Stockholm, 
March 1952. 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 83 

Table I. 
Analyses of the rocks from the Juan Fernandez Islands. 



SiOa . . . 
TiOj ... 
Al,03 . . . 
Fe^O, .. 
FeO ... 
MnO . . . 
CaO . . . 
BaO.... 
MgO . . . 
Na^O . . . 
K^O . . . . 
P.O. ... 
Cr,03 . . . 

F 

CI 

S 





Masatierra (I— IV) 




Masafuera 


(V— VIII) 


I 


II 


III 


IV 


V 


VI 


VII 


VIII 


46.08 


46.10 


46.50 


47.62 


46.51 


43-37 


43-47 


63.43 


348 


3.80 


3.04 


3-42 


3-25 


1.03 


2.68 


0.28 


15.98 


16.54 


13-34 


16.24 


16.68 


8.48 


17.30 


18.64 


2.75 


5.48 


5-34. 


474 


4.30 


2.91 


6.87 


2.78 


8.83 


7.25 


6.71 


7-14 


8.17 


11.00 


7.09 


1.02 


0.23 


0.24 


O.II 


0.18 


0.18 


0.13 


0.07 


0.09 


10.54 


10.64 


10.04 


10.60 


8.85 

O.OI 

6.40 


5-03 


6.09 


1.68 


6.53 


4.58 


8.89 


4.41 


25-93 


8.60 


1.38 


3-61 


2.94 


2.32 


2.60 


2.80 


1-33 


2-53 


6.77 


1.36 


0.88 


0.63 


0.68 


0.36 


0.58 


0.74 


3.82 


0.32 


0.30 


0.29 


0.36 


0.46 


0.19 


0.27 
0.13 


0.18 










0.15 
0.08 


0.08 


0.18 


0.04 


— 


— 


— 


— 


- 


trace 


0.12 


0.01 


0.34 


0.96 


1.63 


1.24 


1.09 


— 


— 


— 


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 i 420 m. Masafuera. A.-M. Bystrom anal. 
VI. Picrite basalt (Masafuerite, Oceanite) dike rock. Masafuera. N. Sahlbom anal. 
VII. Iddingsite-basalt. Elevation i 000 m. Masafuera. N. Sahlbom anal. 
VIII. Soda trachyte. Elevation i 200 m. Masafuera. N. Sahlbom anal. 

Analyses I — IV have been made for A. Lacroix from specimens collected at Bahia Cumberland 
by members of Dumont d'Urville's expedition in 1854, These analyses were first published in 
Lacroix' paper: Constitution lithologique des lies volcaniques de la Polyne'sie 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 — VlII are produced from my earlier paper on the geology of the Juan Fernandez 
Islands (No. 12). 



84 



PER IV ()UEXSEL 



Analyses calculated as water free. 



■ii(\, . 

VVJ >3 

FrO , 
Mn( > . 
CaO , 
I'.aO. . 
M.^O 
Nii,,( ). 
K,() . 
IV>. . 

I' . . . . 

a .. . 



I 


11 


III 


1\- 


y 


VI 


VII 


VIII 


4r..2i 


4().()H 


47-S3 


48.60 


47.36 


43-34 


45-27 


63.36 


349 


3.83 


312 


3-49 


3-31 


103 


2.79 


0.28 


If). 03 


i^'.75 


1372 


16.57 


16.99 


8.48 


18.02 


18.62 


27^> 


5-55 


5-49 


4.84 


4.38 


2.91 


7-15 


2.78 


8..S6 


7-34 


6.90 


7.29 


8.32 


10.99 


7.38 


1.02 


0.23 


024 


0.1 1 


0.18 


0.18 


0.13 


0.07 


0.09 


10.53 


10.78 


10.33 


10.82 


9.01 
0.01 
6.62 


5-03 


6.34 


1.68 


0.55 


4M 


9.15 


4.50 


25.91 


8.96 


1.38 


3.62 


2.98 


2.39 


2.65 


2.85 


1.33 


2.64 


6.76 


1.36 


0.89 


0.65 


0.69 


0-37 


0.58 


0.77 


3-81 


0,32 


0.30 


0.30 


0.37 


0.47 


0.19 


0.28 


0.18 


— 


1 




— 


 — 


— 


0.14 


— 


— 


1 







0.15 
0.08 


0.08 


0.19 


0.04 


1 100.00 


1 100.00 


100.00 


100.00 


100.00 


1 00.00 


100.00 


100.00 



Molecular proportions (x 100) 



Sio, 

!•>.< \ 
!-•<•( ) 
.MnO 
(■;!( ). 

r.ao 
.Mk<> 

Na/) 

IV \ 
CrV'a 
I" ... 
CI . . 



Ill 



76.94 
4-36 

15.72 
i-73 

12.33 
0.32 

18.84 

16.24 
5.84 
1.44 
0.22 



[V 



77.72 
4.81 

16.43 
3-47 

10.22 

0.34 
19.22 

11.51 
4.81 
0.94 
0.21 



79.64 

3-9' 
13.72 

3.44 

9.60 

0.15 

18.42 

22.69 

3.85 
0.69 
0.21 



80.92 

4.36 

16.25 

3.03 
10.15 

0.25 
19.29 

1 1. 16 
4.27 
0.73 
0.26 



VI 



VII 



78.85 
4.13 

16.67 
2.74 

11.58 
0.25 

16.06 
0.01 

16.17 
4.60 
0.39 
0.33 

0.79 
0.23 



72.16 
1.29 
8.32 
1.82 

15.30 
0.18 

8.97 

64.26 
2.14 
0.62 
0.13 



0.23 



75-37 
3-48 

17.68 
4.48 

10.27 
o.io 

11.30 

22.22 
4.26 
0.82 
0.28 
0.09 

0.54 



105.49 

0-35 
18.27 

1-74 
1.42 

0.13 
2.99 

3.42 

10.90 

4.04 

0.13 



0.1 1 



ADDITIONAL COMMENTS ON THE GEOLOGY OF THE JUAN FERNANDEZ ISLANDS 85 

Norms. 



Q 

C . 

or 

ab 

an 

ne 

di 

hy 

ol 

ap 

il . 

ml 

hm 

cm 

sal. 

fern 



II 



8.08 
18.14 
23.48 

6.76 
21.78 

10.46 
0.74 
6.62 
4.01 



56.39 
4361 



0.14 

5-23 
25.22 
29.71 

17-39 
6.28 

0.71 
7.30 
8.03 



60.30 
3971 



III 



0.40 

3.84 
20.19 
24.81 

19.32 
16.84 

0.71 
5-93 
7-97 



49.24 

50.77 



IV 



V 



VI 



VII 



4.25 

4.06 

22.39 
31.29 

16.00 

7.50 

0.87 
6.62 

7.02 



61.99 
38.01 



0.66 
2.17 

24.12 

32.49 

7-43 
19.20 

I.I I 
6.27 
6.34 



59-44 
40.35 



3-45 
1 1.22 

15-47 



6.65 
3-51 
53-07 
0.44 
1.96 
421 



30.14 
69.84 



1-99 
4.56 

22.33 
29.62 



23.40 
1.58 
0.67 
5.28 

10.37 

0.20 
58.50 
41.50 



Quantitative system. 



I. 


Ill 




3 


4 


Camptonose 


or 


:ab 


an 


II. 


III 




4 


4 


dosodic 


or 


ab 


an 


III. 


III 




4 


4 


dosodic 


or 


ab 


an 


IV. 


III 




4 


4 


dosodic 


or 


ab 


an 


V. 


III 




4 


5 


persodic 


or 


ab 


an 


VI. 


IV 




I 


4 


doferrous 


or 


ab 


an 


VII. 


III 




4 


4 


dosodic 


or 


ab 


an 


VIII. 


I 




2 


4 


Laurvikose 


or 


ab 


an 



16.14:36.55 : 47.31 
8.69:41.92:49.39 
7.86 : 41.34: 50.80 
7.03:38.78:54.19 
3.69:41.03: 55.28 

11.44:37.23: 51.33 
8.07:39.51 : 52.42 

25.91 : 65.88: 8.21 



Niggli values. 



4-49 

0.79 

22.48 

57.15 
7.12 



3-43 

0.45 

0.53 
2.78 
0.86 

91-93 
8.05 



si . . 
qz . 
al .. 
f m . 
c , . 
alk 
c/fm 
ti . . 

P •- 
k . 
mg. 
o. . . 
w . 



103.7 

-35-5 
21.2 
43-6 
25.4 
9.8 
0.58 
5.88 
0.30 
0.20 
0.50 

O.II 

0.21 



1 10.4 

-22.4 

23-3 
41.2 

27.3 
8.2 
0.66 
6.83 
0.30 
0.16 
0.40 
0.24 
0.40 



III 



105. 1 

-18.9 

17.8 

51-9 

24.3 

6.0 

0.47 
5.16 
0.28 
0.15 
0.58 
0.17 
0.41 



IV 



118. 7 
-10.5 

23.9 
40.5 
28.3 
l-7> 
0.70 
6.40 
0.38 
0.15 
0.40 
0.22 
0.37 



110.7 

23-4 
47.0 
22.6 
7.0 
0.48 
5.80 
0.66 
0.08 
0.48 
0.16 
0.32 



VI 



VII 



69.8 

41.0 

8.0 
80.6 

8.7 

2.7 

O.II 

1.25 
0.13 
0.22 
0.77 
0.04 
0.19 



99-7 
27.1 
23-4 
550 
14.9 
6.7 
0.27 
4.60 
0.26 
0.16 

0.53 
0.22 
0.46 



VIII 



236.3 
+ 2.3 
40.9 
18.9 
6.7 
33-5 
0.35 
0.78 
0.29 
0.27 
0.40 
0.41 
0.69 



86 PKRCV QU KNSEL 



Bibliography. 

1. 'V\i. Sr 1(1 iFKK. Criisoiiiana. Manchester 1843. 

2. C. Hkrtkko. Notice siir I'Histoire naturelle de Tile Juan Fernandez. Annales des 

Sciences Naturelles, Paris ICS30, I'ome XXI, p. 345. 

3. A. Cai.dc i.FiuJH. On the (leology of the Island of Juan Fernandez. Geol. Soc. of 

London, Proceedings, Vol. i, 1826 — 1833, p. 256. (Also published in Phil. 
.Mag. and Annals of Philosophy, Vol. IX, 1831, p. 220.) 

4. P. Pakkkr Kin(;. Narrative of the surveying voyages of H. M. S. Adventure and 

Beagle. Vol. i. Proceedings of the first ex])edition 1826 — 1830, p. 304. 

5. J. DiMoNi dTrviiik. \'oyage au Pole Sud et dans I'Oceanie. Histoire du Voyage, 

III, 1842. }) . 114. 

6. I. (1ran(.k. (leologie, Mineralogie et (leographie physique du Voyage. Dumont 

d'Urville, Voyage au Pole Sud et dans I'Oceanie, 2^ Partie, 1854, p. 39. 

7. L. Pi A IK. Zur Kenntnis der Insel Juan Fernandez. Verh. der Gesellschaft fiir 

Frdkunde /.u Berlin. Band XXIII, 1896, p. 221. 
<S. A. RiNARD. l\ej)ort 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 Fxpedition, Vol. II, 1889, No. 15. (Also published in 
I'rench under the title 'Notice sur les roches de Pile de Juan Fernandez'. 
Bull. Acad. Belgique, tome 10, 1885, p. 569.) 
9. L. Daraj'skv. Uber den C.lockenstein von Juan Fernandez. Verh. des deutschen 

wissenschaftlichen Vereins zu Santiago, 1886, Heft 3, p. 113. 
o. R. PoHi.MANN. Das Vorkommen und Bildung des sog. Glockensteins (Magnesit). 
Ibid. Band II. 1893, Heft 5—6. 

I. Noticias preliniinares sobre las condiciones jeograficas i jeolojicas del Archi- 

pielago. Publ. in V. Johow: Fstudios sobre la Flora de las islas de Juan 
I-'ernande/. Santiago de Chile 1896, p. i. 
2. P. (^)rKNSKi.. Die (ieologie der Juan Peinandezinseln. Bull. Ceol. Inst, of Upsala, 
\ ol. XI, I 9 I 2, J). 252. 
r. Ha(;i:r.man. lieitriige zur Geologic der Juan Fernandezinseln. The Natural 
History of Juan P'ernandez and Faster Island. Vol. I, 1924. 
14. A. La( Roix. La constitution lithologique des iles volcaniques de la Polynesie 

Australe. Mem. Acad, des Sciences. Paris. Tome 59, 1927. 
I 5. !■. \()N Wo IK. X'ulkanisnnis. Stuttgart. 'Peil I, 19 i 3, p. 290; 'Peil II, 1929, p. 77 i, 805. 
i<>- ]■ Bkm(;(;kn. Inindainentos de la (ieologia de Chile. Santiago de Chile 1950, p. 59, 

17. 'P. II. Ti/Akii, H. N. MosKiKv, J. G. Puchanan and J. Murray. Narrative of the 

cruise of II. M.S. Challenger. Vol. I, 1885, p". 818. 
>8. T. SiK i.iKiK. The eartluiuake of Juan Fernandez as it occurred in the year 
I S3 5. London i.S3(). 

Sixteen years in Chile and Peru from 1822 to 1839. London i84i,p. 387 
(with a sketch ot the sub-marine explosion). 
i'w. Darwin. ( ;colo,L;i( al observations on the volcanic islands of South America, 

visited during the voyage of H. M. S. P)eagle. London 1876, ]). 144. 
I'. Bar in. Pa( ificite, an anemousite basalt. Journ. Washington Acad, of Sc. Vol. 
-X X, I ()3o, p. 60. 
- - Mineral()gi( al Petrf)^ra])hy of the Pacific lavas. Am. J. of Sc. Vol. XXI, 
1 93 I , ]). 3 So, 510. 
L. BowKN. The evolution of igneous rocks. Princeton 1928, p. 163. 



ADDITIONAL COMMENTS ON THE GEOLOGY OF IHE JUAN FERNANDEZ ISLANDS 87 

24. A. JoHANNSEN. Petrography, 111, j). 334. 

25. E. DE Margerie. La face de la 'Jerre. Ill, ]). 1359. 

26. Bailey Willis and H. S. Washington. San Felix and San Ambrosio, their geo- 

logy and petrography. Bull. Geol. Soc. of America. Vol. 35, 1924, p. 365. 

27. L. Chubb and C. Richardson. Geology of Galaprgos, Cocos and Kaster Islands. 

Bernice P. Bishop Museum, Honolulu. Bull. 100, 1933, p. 43, 47, 64. 

28. C. BuRRi. Chemismus und provinciale Verhiiltnisse der jungeruptiven (iesteine 

des pacifischen Oceans und scine.'r I'mrandung. Schw. Min.-petrogr. Mitt. 
Band 6, 1926, p. 177. 

Unfortunately an interesting paper by Gorden A. Macdonald on the 'Hawaiian 
Petrographic Province', published in the Bull. (ieol. Soc. of America (60:2, 1949, 
p. 1588), in which comparisons with Juan Fernandez and other Central Pacific Islands 
are discussed, has evaded my attention until this paj)er was already in print. 



4- A Geographical Sketch of the Juan Fernandez Islands. 

By 

C. SKOTTSBERG. 

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'i5" S. and 78°49'5o" W., and 
of the summit of Masafuera, 33^46' S. and 8o°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 Sutclifife to have occurred in 1835, and another E of 
this island by Goll in 1839 (Bruggen pp. 326, 332). Sutclifife (i, 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 i8th 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 ¥., 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. i. 



90 



C. SKOTTSBERG 



features; of the mountains only Mt. Vunque appears. The names on the map are 
Monkey Key, luist Bay (Pto' Frances), the Spout (a cascade not far from Pta 
Bacalao). West Hay (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 W'l is a Special of Cumberland Bay, of which PI. XVII gives 
a good view, and PI. Will shows the Commodore's camp in the valley later 
named in commemoration of his visit. Masatierra was the rendezvous of Anson's 
scjuadron and brought salvation to the remnants of the crews, of which the greater 
[)art had fallen a victim to scorbut. The winter months of 1741 were spent here 
and the ships refitted. 

Am'omo 1)E Ui.i.o.v's narrative is 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 P'ernandez (Cumberland Bay) and Puerto de Juan 
P'ernandez (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 F^RANCISCO 
Am.\I)()R de Amava 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 Tho.mas Sutcijffe, 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; W^est Bay, 
Ulloa's Puerto del Ingles, Selkirk Bay, and East Bay F'rench Bay. Sugar-loaf Bay 
(V^aqueria) 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 S{)anish name, is known now as Bahia del Padre; La Punta is Pta 
de la Isla. The east ca[)e, 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. X'lllagra, 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. 
X'illagra 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 

' Abbrcridtions. I'>. r.ahia bay, C. - Cordon (range, ridge), Co = Cerro (mountain), L. = 
Loberia scaling grounds, M.-Morro small islet, rock), Pta = Punta (point, cape), Pto = Puerto 
(port, harbour, (). =Quebrada j-iarrow valley, gorge), V. = Valle (valley). 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 9I 

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 JOMN 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 Giinther 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 Giinther, 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 Cliilean charts, 
where, however, Punta is used for Cabo: C. del Padre, C. Tunquillar (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 Vifiillo, Bahia Chupones, C. O'Higgins. Some of the 
names on Friederichsen's map are now forgotten: Bahia de la Fe ( = B. Juanango), 
El Palillo (west head of Pangal), C. Madurgo (W of the east cape, here called 



92 C. SKOTTSBERG 

C. Giiasabullena, a corruption of Hueso de Ballena), Morro Caletas ( = EI 
Verciii<ijo), C. Chupones (now Pta Larga), Bahia Aguabuena [now Tierra Blanca, 
but modern charts liave Pta Aguabuena between T. Blanca and Carvajal (Coq- 
bajal of Friederichsen)j. The to{)ography 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. PoKlll.MANNs sliort description of the islands, with special reference to the 
geology, serves as an introduction to Johow's well-known work on the natural 
histor\-. Johow s map of Masatierra, based on "recientes trabajos recopilados per 
la ()hcina Ilidrogratica en 1895", gives a very unsatisfactory idea of the topo- 
graph)-. 

Amador dc Amaya's map of 1795, with additions and corrections by the 
British (no. 1383) and Chilean navies, remained the basis of all charts until 191 7, 
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 Xegros, 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. 
K 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. Me has taken up Herradura for Bahia del Padre, which is all right, but 
when he called Bahia Juanango "l^^nsenada 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 IVofessor 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 
195 I, I was going to |)ut 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 l-'orce. 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 
aidf of all this material a new sketch map, reproduced here in reduced size, was drawn 
{fig. i). 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, 
area 75.2 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. Frodin's photographs of Masatierra seen from the air in 2 000 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 i: 15000. We had the good luck to get the summits quite free 
from clouds . . ." This story is confirmed by Mr. Frodin's kodachromes which show the central 
and northern parts of Masatierra very clear. On the aerial map in i : 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. 



94 



C. SKOTTSHERG 




5^ 



O 

a 






A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



95 




Fig. 2. Eastern half of Masatierra, seen from the air. — Photo B. Frodin 8/4 1952. 




Fig. 3. End of Cordon Salsipuedes close to Pta San Carlos, showing- the dip of the lava beds. 
At the foot of the ridge the Cemetery. — Photo C. Skottsberg ^/^ 191 7. 



96 C. SKOTTSBERG 

eluding the Vunque 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 
5 — 9° 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 700 m 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 
disap{)earing 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 K 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 — 
900 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. Yery often the lower cloud limit is knife-sharp (Skottsb. 3 fig. 2 on 
p. 808). V'lg. 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. 

Xo extensive geological survey has been made in these islands. Our know- 
ledge is mainly based on OlKNSEL'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 mincralogical compo.sition 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 19 16 — 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 8/^ 1952. 



were described by Hagerman. The joint material served Ouensel 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.e. 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. \aqueria valley, Masatierra, showing bed of doleritic basalt, — After a photograph 

bought in Valparaiso. 




F'ig. 6. Spheroidal weathering of basalt, Valle Colonial, Cu.nbLiicUid Day, — Photo C. Skotts- 

berg 24 191 7. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



99 



Zz~ Hard basalt 
',',','/,' Less hard basalt 
?oV Agglomerate beds 
Dikes black 




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 consoHdated 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. 1 1 (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; I.e. 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, 9). 

The geology of Bahia del Padre was considered by POEIILMANN 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 (i p. 266, 2 p. 56); they "have been subjected to alterations 
in connection with thermal processes during some intermediate phase of volcanic 
activity". 



C. SKOTTSBERG 




Fig. 8. The escarpment between Vaquen'a and Juanango, Masatierra, showing the numerous 
vertical dikes. — Photo C. Skottsberg 26/^ i^qS. 







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Fig. 9. Diagrammatic sketch of part of the escarpment between Vaquen'a and Juanango; comp. 
hg. 8. Dikes black; in nature they do not run quite so regularly. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 




Fig. JO. El Puente seen from the shore in Bahia del Padre, Masatierra. 

15/j 1917. 



Photo C. Skottsberg 



Fig. 10 shows the profile of the Puente. The foot of the chff is more or 
less hidden under talus material. A tufif bed overlayers a sequence of basalt 
and agglomerate beds; one of these, rather distorted, can be followed righ across 
the slope on PI. 102: i, 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 CaCOg: "Die wahrscheinliche Deutung dieser Phano- 
mene ist wohl, dass mit Calciumkarbonat gesattigte thermale Gewasser iiber eine 
Vegetationsdecke geflossen sind, wobei Wurzeln etc. mit einer Kruste von oben 
angegebener Zusammensetzung iiberzogen wurden." This should bear witness of 
late volcanic activity contemporaneous with the existence ot 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 patclies 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. 



I02 C. SKOTTSHKRG 

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" 
(Oiiensel 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 (Cordon Chifladores 400 m, Portezuelo 500 m), feldspar 
basalts seem to predominate (Ouensel I.e.). 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 PI. 97:2 in Skottsb. 3, about 450 m 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 mineraiogical composition of the rocks forming the 
highest summit. Mt. Yunque rises about 350 m above Portezuelo de Villagra. 



A geographical reconnaissance of Masatierra. 

Pta Ilueso 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 f)assage along the foot of the escarpment. 

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

Pto I-nviccs does not deserve to be called a harbour; it offers no protection 
even as an occasional anchorage (Instrucc. naut. p. 226). It is a small, open cove 
facing N and IC, 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. Tlie lower slopes of the valley are very barren, the soil 
is exposed or covered with })atches 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 50 m 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 



03 




Fig. II. Most easterly part of Masatierra, seen from the air, looking east. The detached, coni- 
cal rock is El Verdugo. Ridges a (centre) and b. — Photo B. Frodin ^\^ 1952. 




Fig. 12. 



Pto Frances seen from a spur about 350 m above the sea. 

"/12 1916. 



Photo C. Skottsberg 



I04 



C. SKOTTSBERG 




Fig. 13. Rulge a overlooking the E branch of Frances valley. ^ Photo C. Skottsberg ^'\^ 1917. 




\. 14. Pto Frances, W valley-branch. Left, c\ right, d. — Photo C. Skottsberg 1^/4 iQi?- 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



105 




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 Cordon Chifladores. From left to 
right ridges a, b and c. — Photo B. Frodin 8/4 1952. 




t* " 



'^■'"'^f If^"- 



»^-. ^ 



Fig. 16. Continues fig. 15, to Pta Pescadores. From left to right Quebr. Lapiz, Pesca de ios 
Viejos, Laura and Piedra Agujereada; ridges a, b, c, d. — Photo B. Frodin s/i 1952. 

7 - 537351 TA^ Nat. Hist, of Juan Fernandez and Easter Isl. Vol. i. 



Io6 C. SKOTTSBERG 

eral quebradas belong, filled with inaqui-infested, grazed liima 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, b and c denote three conspicuous crests, easy to identify also on 
tigs. II, 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 Pta 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 Cliijiadores (Whistlers' ridge, fig. 16) is, as most of the ridges 
extending X — 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 35°. 
Scattered trees are remnants of the once closed forest. The interior is well wooded 
u[) 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 litde 
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 \l 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 O. de la Piedra Ai^ujereada (fig. 19) which got its name from a rock pierced 

' For some of the leading species the local names are used. Canelo, Drimys conferii- 
folia\ L'houU, Juf/fi/n auslralis\ Luma, Aof/iof>iyniti fcrnandesiana (Masatierra) and Myrceuge- 
nia Schulzei (Masafiiera;; Maqui, Aristotelin ;;/«7^//'/ chilensis; a macal is a maqui grove); Naran- 
jillo, l-agani mnyu Masatierra and /-'. ^.r/^/vm ; Masafuera; ; Pangue, C/^'w^/t-r^ /^//^//rt (Masatierra) 
and (/'. Masafuc?-nc Masafuera . 

' The text is incorrect. Tlie photograph was taken by the author 17.4. 191 7. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



107 




Fig-. 17. Quebr. Pesca de los Viejos, looking S; ridge d. — Photo C. Skottsberg 1^/4 191; 





'i'^i!i^''.S?f^. iu-'it 



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



io8 



C. SKOTTSBERG 




Fig. 19. The interior of Quebr. Piedra Agujereada. — Photo K. Backstrom Aug. 191 7. 



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 sathnis, 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 191 7 the dry soil was covered with the dead stalks and innumerable young 
rosettes of Silybum inaria7m)n (see Skottsb. 3 fig. 34 on p. 891). PI. 86 (I.e.) shows 
the same spot in August, when a vigorous new growth had sprung up. The shadow 
across the Silybctuui 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 (I.e. PI. 94: i). 

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 



09 







O 

^ o 



2 I 



c o 

c bo 

03 13 



C. SKOTl'SHERG 




Fig. 2 1. View W from the summit of Centinela ridge, c. 780 m, toward Mt. Yunque and Mt. 

I'inimide right', in front of Yunque Mt. Damajuana; below Piramide the crest between Quebr. 

Minero and Pangal. — Photo C. Skottsberg ^^4 ^9^7- 



Cordoji 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 to[) 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 
(I.e. PI. 90: 2). 

I^ta Loheria (lobo, seal; [)lace 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 Cionherland or San Juan Ihmiista is the only harbour in the islands 
where large ships find good anchorage; see the charts and descriptions by Walter, 
Sutclifie, Lopez, Vidal Ciormaz, Glinther etc. The bay is open toward N and NE» 

' Ace. to (iuzman p. 16 not used now. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 




Fig. 22. Cumberland Bay from Pta Loberia to Cordon Central, seen from the air in looo m 

elevation. From left to right, Pangal, Cordon Escarpado, Quebr. Minero-Damajuana, La Dama- 

juana, Valle Anson with El Yunque. — Photo B. Frodin ^/^ 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. Cordon 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 Piramide), Cerro de los Munoces, Picacho de la Mona 
(= she-monkey) and Cerro el Tope ("top"). Pico Central, a long established name, 
is not mentioned by Guzman. 

ii/ 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. Pis. 'j'^, 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 
(I.e. PI. 89: i). A branch with a patch of forest comes down from the Centinela 
ridge (fig. 23). 



C. SKOTTSBERG 




Fig. 23. Centinela ridge and Pangal, seen from the slope of Salsipuedes above the Cemetery. 

Photo B. Frodin 2»/i 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 Daniajuana (figs. 25, 26). The lower slopes have been cleared, but 
between 1 50 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 Damajiiana (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 1^2nglish 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. 

\ ^alle de Anson drains the loftiest part of Masatierra and is watered by two 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



»»3 




Fig. 24. Quebr. del Minero. — Photo C. Skottsberg 2^/^^ ,916. 



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 Yimque ("small 
square"), situated about 220 m above the sea (Skottsb. 3 PI. 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 as a 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 Rubus 
ulmifoUus when he visited the island a few years ago. 



1 14 



C. SKOTTSBERG 




Fig. 



frc 



25. Damajuana and Vunque from tlic ucui u> I'aiigal, looking SW. 

"/4 1917. 



Photo C. Skottsberg 



55° to 70°, in places almost perpendicular. The exact height is not known; 
the fit^iires vary from 1700 ft (537 m, Walpole) to 983 m (Viel); Branchi has 805 m, 
tiie English chart 3005 ft (913 m), the new Chilean 915 m, Instrucc. naut. 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 IJamajuana-Yunque saddle seems to lead to the summit. 
The figure 838 m (doubtless too high) on Friederichsen's map corresponds to a small 
{/mnacle set on the saddle, the "Camote" (= sweet potato, bulb). The ascent, which 
is difficult and dangerous, was described by Tp:nz, who was the first to give us 
any information on the topography and plant life of the summit; see Skottsb. 3 
pp. S97— 898. 

Cordivi Coitral (figs. 22, 27, 31, 99). From the broad gable of Mt. Piramide, 
F 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. Pico Central, c. 570 m, marks the end of the harder 
basalts Ijcloiiging to the upper horizons; I.e. PI. 89: 2, Johow PI. IV. 

\'allc ('dlofiiai (V. del l^olvorin 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 FERNANDEZ ISLANDS 



115 




Fie 26 Mt Damajuana seen from Anson's valley. In the foreground a section through an 
^* " agglomerate bed. — Photo C. Skottsberg 2^/12 1916. 



i6 



C. SKOTTSBERG 




Fig. 27. Valle Colonial seen from the Centinela ridge, c. 350 m above sea level. Left, Cordon 
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 

C. Skottsberg "/4 19 17. 

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. 

FA Pirdmidc (figs. 21, 29, 31), separated from Mt. Yunque by a narrow saddle 
(W l^ortezuelo del Yunque), towers above the colony. Johow's PI. 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. 
3 PI. 9i)- 

Portezuelo 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 




Fig. 28. Portezuelo de Villagra, seen from above the settlement in Valle Colonial. 

C. Skottsberg 3«/ia 1916. 



Photo 



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 (i 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 fallen den Talboden, was darauf hindeutet, dass es sich nicht um ein P>osionstal, 



C. SKOl rSBKRCr 




Fi^. 2(). South side of Portezuelo with tlic .Scikhk iiicinoriai 

I'hoto (". Skottsberj; ^^j^^ 1916. 



)ict, seen troni the pass. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I I 9 

sondern urn eine teilweise erhaltene Kraterbildung handelt." Finally, "Ob das west- 
lich von dem Portezuelo sich offnende cirkusformige Tal audi 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 Rucken des Portezuelo 
die Scheidemauer bildet." This was written after our visit in August 1908, and at 
that time I shared my companion's opinion (i 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 Bahi'a 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 Lavabruchstiicke, 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. 



r. SKOITSBERG 




F'ij^. 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. Frodin 2*/^ 1952. 




Fig. 31. Cumberland I'.ay in SSK, seen from the air. From left to right Quebr. Minero-Dama- 

juana, Damajuana, Portezuelo del ^'unque, Valle Anson, El Yunque, El Piramide with Cordon 

and Pico Central, .Salsipuedes with Pta San Carlos. — Photo B. Frodin »U 1952. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 




Fig. 32. From the trail across the Salsipuedes highland, looking SE. Photo C. Skottsberg ^i/^ 191 7. 



Cordon Salsipuedes forms the western boundary of Cumberland Bay and runs 
from the main range to Pta 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 wath crags. Already at about 600 m it is only 
I — 2 m wide, and the rise, quite gentle farther down, increases to 25 — 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 PI. 90: i from 19 16 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, O. Gutierrez and O. del Monte 
Maderugo, were thickly wooded in 19 1 7 (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. SKOTTSHERG 

rather abruptly into V. Ingles. The quebradas coming down from the main ridge 
are filled with a remarkable Dicksonia 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 luf^U's 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 {)ierced 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 I.e., 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 cordon 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. 
90: 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 PI. 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 [)erhaps 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. 

Ccrro Alto boldly terminates the dividing ridge between Pto Ingles and Vaqueria 
(fi^^- 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 cjuite [)erpendicular. Patches of forest are seen on the flanks of the cone. 

Bahia dc la Wiqucria 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. Mg. 39 is a general view of the valley seen from the air. 
The outer [)art 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 X'aquen'a have been left to run wild, and the visitor should look out for 
the bulls. 

As far as I could sec, the geological structure is the same as in Cumberland 
Hay, 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 



123 




liiilii 



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 20^1 1917. 




.V. ... 



^r-yr Vjs^-; 



Fig. 34. Ihe cave ("Robinson's grotto") in Pto Ingles. — Photo P. Quensel "/g 1908. 




SKOTTSBERG 




35. The surroundings of Pto Ingles, seen from the air; note the crescent-shaped saddle 
o\erlooking the south coast of the island. — Photo B. Frodin ^1^ 1952. 




Fig. 36. The gently sloping floor of V'alle Ingles with the main range in the background; right, 
the dividing ridge where, in 1908, the last living Satitalum grew. — Photo K. Backstrom 191 7. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



25 




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 ^'12 191 6. 




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 ^^/^ 1917. 



126 



SKOTTSHERG 




^ 'ft- 39- I 'i6 Vaquen'a cove and valley, seen from the air. Left, Cerro Alto, right, ridge 

between Vaqueria and Juanango; the top of Cerro Chumacera visible behind. — 

Photo B. Frodin 8/4 1952. 




I Co Agiulo 



7 » 9 

:i I'ir;'uni(ie; 3 Kl Yunque; 4 Co Chumacera; 5 Co Tres Puntas; 6 V. Villagra; 
7 \illa Alcmana; 8 M. Juanango; 9 V. Juanango. 

Fig. 40. I5ahia Juanango with Pta Negros and Morro Juanango, seen from the air looking SE. 

— Photo B. Frodin 8/4 1952. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I 27 

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, o^ 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, Pta de los Negros, and SE of this lies Morro Juanango (fig. 40). 

Bahia del Juanango 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. Villa Alemana (Germantown), accessible with difficulty, 
and Q. Juanango. 

Q. del Juanafigo. 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 Juanango to Bahia del Padre the coast escarpment trends SW in wide, 
slightly concave curves to Pta Letnos and Pta Tunquillar and thence to Pta 
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 PI. 102: i) 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 "risuenas representantes del mundp vegetal" praised 
by Guzman (p. 24), except a patch of salt-meadow [Salicornia) 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. El Puente was described and 
illustrated above (p. loi, fig. 10); the flat, sandy surface is seen on figs, 45 and 46, 
the former showing the wind-polished tufT 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 





Fig. 41, 







Jl 



Fig. 42. 

Hgs 4 1 --42. North coast of western Masatierra from h. Juanango to Pta Lemos, seen from 
<Juebr. Juanango. Cerros Cluimacera note patches of luma, Tres Puntas and PInrique. — Photo 

C. Skottsberg ^/^ 1917. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 1 29 




Fig. 43. Cerro Tres Puntas. — Photo Hans Frey 



C. SKOTTSHERG 






Fig. 44. The entrance to Bahia del Padre. — Photo C. Skottsberg ^'^j^ 191 7. 



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 aufvveist" (p. 26). 

On the naked sand we found many living beetles (no reference is made to them 
in vol. Ill) and empty shells of four species of landshells, Fernandezia tryoniY''\%- 
bry, Succinea fernandi Reeve (also in sand at Tierra Blanca), S. texia 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. loi) were formed? Or did dwarf trees such as Dendroseris 
I'ltoralis, Rea pruijiata and Clieiwpodinm Saiictae 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 Kea and Dendroseris 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 




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. Backstrom "/^ 191 7. 




Fig. 46. Mobile sand on Puente, in the foreground a dense growth of weeds, mainly Clienopo- 
dlum multifidum. C. S. as measure. — Photo K. Backstrom ^^/^ 191 7. 



32 



C. 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 w-e have 
to find out if landshells live on the plants mentioned above. 



With Pta dc la Is/a 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. 

P/a (y///[i^x///s, watched by a nameless morro (small skerry), and the inhosp- 
itable coast cliffs of BaJiia Carvajal are seen in fig. 47. With a NW wind 
boats engaged in langost fishing at Santa Clara find shelter here. From Pta 
Agiiabuoia to Co Xegro the coast shows several well-marked bights. 

Haliia Ticna Blanca or Tierras Blaiicas 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 I.a>\i!^a (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 Enri- 
que. From a distance it seems to be formed by basalts of the higher horizons, but 
no specimens were brought. 

Pallia Chupones derives its name from "chupon", in Chile a vernacular name 
for (ireii!;ia spliacelata Reg. (Bromeliaceae), which has edible fruits (chupar = suck), 
and ap|)lic(l by the fishermen of Masatierra to the extremely rare Hespevogreigia 
as well as to Ocliagaria; 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 likcl)- occurs here, even if we did not observe it W of Tres Puntas, where 
it co\ crs rock faces (see Skottsb. 3 PI. 97). The slopes round the bay are grass- 
land, in tile western half almost pure Aveiia barbata, otherwise with extensive 
|)atchcs of the native StipctiDii. All streambeds in this western section are dry 
most of the year. 

Ijuna Psiarpa(ta\ 385 m high where we crossed it near the main range, 
separates B. Ciiuj)ones from \'illagra Bay, this taken in a wide sense. Off the 
j)<)int lies Mono I'ihillo (vifiilla means "small hill planted with vines", but the 
name must refer to something quite different in this case); the gradient of the 
surface shows tiie tilt of the lava beds (fig. 49). 

Pallia (tr Mllagra and its valleys, the bay taken in a wide sense and 

\\\ my tield notes I called this ridge "Cordon ?2scarpado", and this name also appears 
on tlic Chilfan chart, and on my map I had used the same name for the ridge between Pangal 
and <^). Mincro, and as it was published in this sense Skottsb. 3 pp. 890, 915 , I have renamed 
the other ridge Loma Kscarpada. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



^33 




Fig. 47. View from Tuente toward liahia Carvajal and Pta O'Higgins. Behind, Santa Clara. 

Photo K. Backstrom ^^j-^ 1917. 




Fig. 48. Bahia Tierra Blanca seen from Pta Larga. — Photo C. Skottsberg ^/^ 191 7. 



34 



C. SKO'lTSBERG 




V\^^. 49. \'ie\v from Portezuelo de Villagra looking SW, with Santa Clara in the background. 

Morro V'inillo in the middle, right the south coast of Masatierra from Bahia Villagra to Pta 

O'Higgins. — Photo C. Skottsberg ^^l^^ 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 
Tuntas 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 
|). 896) with a small waterfall a grove of BoeJimeria. 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. 

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

X'illagra is watered 1)\' three permanent streams and densely wooded. The 
scener)- is even more grand than on the Cumberland side, with the sequence of 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



135 




Fig. 50. View from Portezuelo de Villagra lookiiig W. From W to E Cerros Enrique, Tres 
Puntas and Chumacera. — Photo C. Skottsberg "/^ 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 — 70° 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, I.e. PI. 85: i) and on the slope of Mt. Piramide 
(I.e. PI. 91) 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, PI. 85: i) I have discussed the nature of 
this timber-line. Below the forest degraded grass-land with foreign grasses and 
herbs dominates over the natural Siipeium. Along the streams a fringe of pangue 
runs down toward the sea (I.e. PI. 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 I.e. 
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 



^36 



C. SKOTTSHERG 




FiK- 5'- 1-1 N uiKiiie seen from the \'illagra slope c. 175 m above sea level. — Photo C. Skotts- 

berg «/i 191 7. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



137 




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; I.e. PL 92: 1). 

From El Yunqiie 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 Molina, an Italian-Chilean naturalist of the i8th 
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 clifi" walls and gorges; 
see figs. 2 and 11. The sinister name ir/ Fifr^//^^ (executioner, hangman, fig. 11) 
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. 



38 



C. SKOTTSRERG 




^* 'h- 53- 'Santa Clara from the air, looking" SE. — Photo B. Frodin ^/^ 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. Frodin'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 Jolioic 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 I 39 

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 Ai/ejia in dominance, all very dry 
during the summer. When Giinther says that there were "algunos arboles en la 
pendiente del este" he either did see some specimens o^ Dendroscrjs on iha c\\f(s 
above the water or refers to Morro Spartan (also called M. de los Alelfes; 
fishermen are said to have collected seeds of Matthiola 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 Doiia 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 PI. 103. The dikes project 
as flat slabs above the softer beds. 



Masafuera, former surveys and maps. 

Until our visit in 19 17 Masafuera was much less known than her sister island. 
It had been inhabited more permanently only during the period of the penal 
settlement 1909 — 19 13, 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 ofl"er sufficient shelter for any ship desirous of refreshing 
ashore" (Walter p. 134). As four ships of the squadron were missing the Com- 



140 



C. SKOTTSBERG 



MASAFUERA 




Vicente Forrai 



Kodado del Sindilo 



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 (I.e. p. 156). 
PI. X.XI is a view of the Xl^ side of "Masa-Fuero", as the name is spelled. It 
shows the table-land uj) 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, b, b, and c\ a seems to indicate a reef, 
b and b arc the entrances to two of the canyons. I believe that a also indicates 
(J. Casas and that b are (J. Ovalo and (,). Sanchez; c is the waterfall coming 
down from (). Larga. PI. XXII, pretending to show the west side, is less easy 
to read. 1 think tliat it re[)resents the north-west section of the coast from Cabo 
Xorte south toward Loberia X'ieja, 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 I 41 

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 afucra de Jiam Fer- 
ftajtdea 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. 6^) he must have included the Dkksonia 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 Sanialuni 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 191 7, 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 Hidrografica published the first map showing the prin- 
cipal topographical features. New dates had been provided by Johow. During 
our visit in 191 7 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). I he 
table-land is traversed by numerous parallel deep valleys running NK — L (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). 



42 



C. SKOTTSBERG 




Fi) 



%-*j 



W. Quebr. de las Casas at the extreme left. — Photo 
Backstrom Feb. 191 7. 



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 Httle 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 consecjucnce, 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 O. Sanchez and O. 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 
Loben'a Vcntana 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 ^/^ 
of the shore and to proceed from one canyon entrance to the next greatly 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I 43 

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; Gunther, 
9.25 by 6 km, and Guzman, 17(1) 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 Ouensel, 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 tuftaceous 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 191 7 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 Blocken 
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 tufi"aceous 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 1 100 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 iioo 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 



44 



C. SKOTTSBERG 



m-^^' 




V\^. 56. East coast of Masafuera between Pta Negra and Playa Ancha, with entrance to Q. del 
Varadero. — Photo C. Skottsberg 21/^ 19 17. 




Fig. 57. Part of the east side of Masafuera seen from the air. The highland is covered with 
thick fog. — Photo B. Frodin 8/4 1952. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISI-ANDS 



45 




Fig. 58. View from the high land above Las Chozas looking toward the Ovalo, Pasto and 
Sandalo valleys. — Photo C. Skottsberg 3/3 1917. 





^^*mA 



'"^f^mt^ 



Fig. 59. The shore at Loberia Vieja with a crag on the reef. — Photo K. Backstrom 1V2 ^9^7- 



146 



C. SKOTTSBERG 




Fig. 60. At the landing-place in Las Casas. Our party is about to embark, assisted by the crow 
of the schooner. — Photo K. Backstrom 1^/3 191 7. 




Fig. 61. As tig. 60, showing ijcach ot large boulders. — Photo K. Backstrom "/j 1917. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



f47 




Fig. 62. Slope of Mt. Inocentes seen from Cordon Atravesado, c. 1350 m ctouvc sea level. — 

Photo C, Skottsberg ^^/j 191 7. 



resemble the hard basalts between 400 and 600 m on Masatierra (I.e.). The sum- 
mit of Mt. Inocentes consists of a still different rock, a slaggy olivine basalt 
supersaturated with iron oxides (I.e. 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 (I.e. 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 'j'j 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 



148 



C. SKOTTSBERG 











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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 Gunnera 
Masafuerae. — Photo K. Backstrom Feb. 19 17. 




Fig. 65. Looking down into the interior of Casas canyon from the crest of Cordon del Barril. 

— Photo K. Backstrom 1/3 19' 7- 



ISO 



C. SKOTISBERG 





V **-. \ t " 

Fij(. 66. The grass covered table-iand ijetween 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 ^^/j 191 7. 

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 Inocefites 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 (Hranchi) 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. y\bove 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 



15 




Fig. 67. Cordon del Barril, looking W toward C. Atravesado. Note goat track along the ridge. 

— Photo C. Skottsberg 1/3 ^9^7- 



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 i 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. no. Johow called this place "El Pangal", and nowhere is the 
pangue [Gunnera Masafuerae) more luxuriant (I.e. PI. iii). About 2 km from the 



152 



C. SKOTTSBERG 




FiJ,^ 68. Entrance to Quebr. de las Vacas. — Photo C. Skottsberg ^^j^ 1917- 



entrance and 210 m 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. Q. 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 luirril. 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 tliis {)lace, and extending west c. 2 km, where the cordon 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, (.). ("hica, (). Hlindado and O. Cabreros. Q. Cliica (-= small) is very 
shallow and dry, but some luma trees are seen. Q. del Bliiidado (B. means 
ironclad cruiser, but may have a very different significance here) is much larger 
and tlie water comes down over a threshold about 500 m upstream and has dug 
out a miniature caiu'on. Tiiere is good forest between 400 and 500 m and some 
groups of trees a little farther clown. There is some forest also in Q. de los 
Cahreros ((ioat-hunters' valle)-). Where the narrowing rocky Barril ridge, which 
has its name from a barrel sha|)ed 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. G'j). The ridge slopes 50 — 60° on 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I 53 

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

Q. 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 19 17 
and were introduced again in 1927). Following the beach south from Casas 
past Pta 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 i 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 Play a Ancha, a level stretch of stony beach, from where Q. de los 
Inocentes 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 O. 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. i 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 Dicksonia 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—80° are the rule in these gorges. 

io~53735i ^^^ ^<^i- Hist, of Juan Fernandez and Easter I si. Vol. I. 



54 



C. SKOTTSBERG 






A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



55 




156 



C. SKOTTSHERG 




Fig. 71. A low threshold in the Vacas canyon. — Photo C. Skottsberg ^^1^ 191 7- 



^; 





I'ig. 72. 'rwo witches' cauldrons in the \'acas canyon. — I'hoto C. Skottsberg "/j 1917- 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



57 




Fig. 73. A view of the interior of Quebr. Angosta, seen from the ridge N ol lins v.uicy, 550 m 
above sea-level. — Photo C. Skottsberg ^j^ 191 7- 



T/ie sotitJi coast. The strip of beach is narrow S of Varadero, and the escarp- 
ment reaches an imposing height at Pta 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, Tierras Blancas, from the light colour of 
the deposits. Along Rodado del Sdndalo (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. ']^. At Vicente Porras we arrive on the west coast. The long and broad abra- 
sion terrace. Play a 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 i 1/2 cable (277.5 "i) ^^' 
cording to Giinther, 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 
Q. 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. 'j'i). 
The altitude of the valley floor is only c. 190 m. 

A short but impassable stretch prevents us from reaching the remaining 



158 



r. SKOTTSHERG 



^= 










FiK. 74; l^ntraiKc to Ouebr. del X'ar.ulcro, from where, up the south wall (left) an ascent was 
made. The knob on the extreme right is shown in fi(,^ 75. — Photo C. Skottsberg "/a 1917. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



159 




Fig. 75. Looking into the Varadero canyon. — Photo C. Skottsberg ^i/,^ 1917. 



1 6o 



;kottsberg 




P^ig. 76. Coast of Masafuera between Tierras Blancas and Vicente Porras. 

Strom ^^/a 191 7. 



Photo K. Back- 



"loberias", L. \^entaua (=^ window, a hole in the rock), L. Nueva and L. del Buque 
I 'arado. 

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 tlie barranca is of formidable height and steepness (fig. 78). A stream 
coming from the gullies above has cut a winding bed (fig. 82). 

I'jiseiiada loltthi 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 
Token is (juite unfit as a base camp, as the highland cannot be reached, nor 
is there a j)assage along the shore to O. Sanchez. 



The table-land N of 0- de las Casas. 

The Plauo 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 O. del Mono (another cjueer 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 



6i 




Fig. ^T. Interior of the Loberia Vieja canyon. — Photo C. Skottsberg ^'/a 19^ 7- 



C. SKOTTSHERG 



I•'i^^ 7<S. Pta del liii.in and Playa del Buque Varado, seen from the sea looking S. — Photo 

C. Skottsberg ^^/g 191 7. 



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 O. Pasto, 1130m 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 i)lain and to Las Torres (Towers), c. 1370 m, and Co Correspondencia, 
c. 1420 m. V\\r. 80 is a view of one of the Loberia gorges seen from Corres- 
pondencia. ihe 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 cla\-ey and the surface, which shows distinct signs of water erosion, 
strewn with stones and cracked in places. Some small boulders rested on short 
jMllars of cla>'. The ground is perhaps flooded during the winter. I was told that 
{)atches of snow have been observed here, but this needs corroboration. Wind 
erosion might be res[)onsible for the formation of the small "tables". 

Ai'ciiida dc las Cahras, "Goat Avenue", is a well-beaten track running along 
the very edge of the abyss from C. Atravesado to Co Verde. From about 
1100-125001 alt. one looks down on Buque Varado (fig. 82) and Q. del hndn 
(Magnet gorge, fig. 83). One of the former inmates of the convict settlement told 
me that he had been em{)loyed 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 



163 




] 



Fig. 79. View across Piano de la Mona toward the Larga and Sanchez valleys, elev. 940 m. 

— Photo C. Skottsberg 25/2 ^9^7- 




Fig. 80. Looking down into one of the upper gorges of Quebr. Loberia from an altitude of 
c. 1400 m. — Photo C. Skottsberg "/a ^9^7- 



i64 



C. SKO'lTSHERG 






Fi.l,^ 8i. 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 oS. Lophosoria. — Photo C. Skottsberg ^^/a 191 7- 



Lookins 



down on I'laya del liuque Varado from c. 1 200 m. 
^3 I9I7. 



Photo C. Skottsberg 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



165 




Fig. 83. Quebr. del Iman seen from c. iioo m. Left, Buque Varado. — Photo C. Skottsberg 

73 1917- 




Fig 84 Piano de la Mona, looking N, with Cerro Verde in the background. — Photo C. Skotts- 
berg 73 191 7. 



66 



C. SKOTTSHERG 




Fig. 85. Looking S tf)\var(l Loberia Nueva from Avenida de las Cabras N of Buque Varado. 

— Photo C. Skottsberg '/a 1917. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



167 



I 68 C. SKOTTSBKRG 

only scattered patches of trees on this side, while ferns, especially Lophosoria, 
pant^ue, <^rass etc. form a more or less continuous cover. A picture taken from a 
point between Buque \'arado and Iman gives some idea of the scenery (fig. 85). 
At the north extremity of the plain stands a green, rounded hill, Cerro Verde 
(fig. 84), mentioned as Pico \orte in Instrucc. naut. p. 230 and said to be 1 340 m 
high. This figure is too high, I believe. 

From Cabo Norte to 0- de las Casas. 

The coast cliffs from Cabo Xorte to the Sanchez shore are almost perpen- 
dicular and about 300 m high. Half way a cascade tumbles down to the beach 
in two lea[)s, coming from the gap of O. 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. 

0. lie Scvichez 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 0. Xegra (also called O. del Plan Negro) does not reach the shore. Q. del 
Saudalito and O. del Sdndalo do not seem to offer anything of particular interest. 
The talus slope permits a fairly comfortable passage as far south as to the large 
O. 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. del 
0/mlo, which derives its name from a conspicuous hill inside the entrance, a good 
exam[)le of columnar structure. The passage is a little fatiguing until we are op- 
posite O. 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, I^Y'rnandez 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 P\M-nandez' death the island was turned over to 
the ( )r(lcr 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 
ot the buccaneers" the islands served as a place of refreshment, but the damage 
was, I (laresa\-, confined to the surroundings of the harbours. Their first visit is 
said to lia\e taken [)lace 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. 127 — 131. 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 



69 




Fig. 87. Ruins of the old Spanish fort in Cumberland Ray. — Photo C. Skottsberg 28/4 191 7- 



in full swing between the islands and Perii. Toward the middle of the i8th 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 
II ""537351 T^h^ Nat. Hist, of Juan Fernandez and Easter Isl. Vol. I. 



lyo C. SKOTTSBERG 

a i^rcat asset durin*^ the era of the corsairs, and in order to stop this traffic 
mastiffs were landed as early as in 1686 or Sy (Hurney IV p. 210). The result 
was tiiat the t^oats 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. I lis 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 looo, 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 I.e.), but the tenants took no interest 
in farming, and in i860 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 
Hay 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 peoi)le like Governor Sutcliffe or Mrs. Graiiam could 
dream of Masatierra as fit not only to (ecd 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 1 7 I 

invaluable as a port." And Sutcliffe (i pp. 442—443) cites a letter written to him 
by one MANUEL DE Salas, Sept. i, 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, iio 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 loooo — 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 = Acaejta argentea, a noxious weed — was common and 



1 Translated from Spanish. 



72 



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 Vie! recommends raising cattle. 
The island could easily maintain looo, 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 })rotection: "the output was mediocre, a consequence of the 
settlers not being sufficiently intelligent and industrious"^ — he forgot that the 
j)oor 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, onl\' 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 
\'alparaiso Fkllexberg 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. 

Ai.KRKi) vox Root, 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, h'.ncouraged 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 |)osition of the place; he tells that there were 7000 wild 
goats and [)asture for 1000 head of cattle and that he intended to start "verschie- 
dene Industrien '. I'A'idently 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. 

I he 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 \ isitcd the islands; the sons of Alfred made their living as lobster ^ fish- 

^ 'I'ranslated from .Spanish. 

2 i'rofrssor Kari. I..\N(;, head of the Dept. of Evcrtebratcs in the Nat. Hist. Museum, Stock- 
hohii, on my ro(|ucst kindly made a dihgent search for an English equivalent to the Spanish 
word langosta as name of the large Decapod Jasus Lalatidci (formerly known as Palinurtis 
ln>n/a/is , which from a scientific viewpoint is no real lobster. He reports that Palinurus vul- 
i^uiris and related forms are called "rock lobster"; "spiny lobster" is another name. Yox the sake 
of brevity "lobster" or langosta" will be used here. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I 73 

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. Ill) 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.^ 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 fiir 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 PI. XIX shows that his "sea lions" were sea elephants. 
UUoa 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".^ 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 i, 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. SKOTTSHERG 



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. 

Hut let us return to l^rmel. l^ven if fishing ranked first, the wealth of the 
land was b\' 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 — 92 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 (jfasus Lalandei\ confined to 
Juan I^'ernandez and the Desventuradas Islands (San Ambrosio and San Felix), and 
the bacalao {Polyprion prog7tatiis), 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 w^orst enemies of Jastis. The waters abound in many other kinds of 
savoury fishes. When, however, the commission recommends to repeal the law of 
1 89 1, 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 sandalo 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 caracter comercial i economico" 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 I 7 5 

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, Antennaria scoriadea 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 erythrojioius 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 endemica 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. 



76 



C. SKOTTSllERC 




h ig. 88. The boat harbour in Cumberland Bay. — Photo C. Skottsberg ^/jg 1916. 




Fig. 89. Lobster fishers working for Recart y Doniez by their boats during the closed season. 
Cumberland Bay. — Photo K. Backstrom Dec. 19 16. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



77 




Fig. 90. The take is brought into the Recart y Doniez factory; left, Hve lobsters; right, the 
metal cooking baskets. — Photo C. Skottsberg »/i 191 7. 



P 



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/i 1917. 

12 —537351 The Nat. Hist, of Juan Fernandez and Easter Isl. Vol. I. 



C. SKOTTSBERG 




P'ig. 92. The village in Cumberland Bay, seen from the slope of Salsipuedes. 

'berg 21/^ 1 91 7. 



Photo C. Skotts- 



Masafuera got no supervisor, the management of the forests continued as before, 
the clionta 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 19 14 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—91) and to the export of the living animals, in 
the com()any 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 



79 






-C^ ". 



?y^. 



Fig. 93. Carica papaya in the garden of Mr. Charpentier, Valle Anson. 

berg i*/4 1917- 



Photo C. Skotts- 



if the passage could be completed in 2 V2 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, Albizzia, 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 cliinate 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. 



i8o 



C. SKOTTSBERG 




Fig. 94. Threshing the wheat with a tropilla of horses, Cumberland Bay. — Photo K. Back- 

strom 1917. 



m 


mmmmmemwsm 




'■■--■^m- ™ ■—  -— ,^^^i* . . 1 _^ 




■-— p--- 




^^^— jMjJfc , i. "■■./*;*»■ 1 



Fig. 95. Ch..;; ...A...^. ;.,c wind. Same place as lig. 94. - Tholo K. Backslrui;. uji] 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



8r 




Fig. 96. A rodeo in Cumberland Bay, — Photo K. Backstrom 191 7. 



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., Tropaeoluni 
ntajus, 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 slieep, and 
we were never offered mutton. 

After 19 1 7 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. 



C. SKOTTSP.KRG 




J 



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 — loi. A short summary is given here. Guzman begins with the 
following words: "Tal vez la descripcion 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-sup[)orting, 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 conviccion que tenian los creadores del presidio en la fe- 
racidad de la estcril Mas Afuera, que al alcalde se le designo con el singular titulo 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS 



183 




Fig. 98. The ruined forest and village of Las Chozas, Masafuera. — Photo K. Backstrom Feb. 191 7. 

de Director y ]e^e 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, catde 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 191 3 Masafuera was abandoned. The 
buildings in Casas were in tolerably good condition in 191 7 (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 prodiga en helechos y en plantas sub-alpinas, no fuera generosa tambien 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 



i84 



C. SKOTTSBERG 

Table I. 
Lobster catcli 1940—46. After Guzman. 



i<>40, 

i«(4i 

1042, 

i')43 

1044 

K'45 



Gross weight in kg 



Number 










Total 


Living 


Tinned 


log 250 


11^ 531 


15 533 


96998 


103 6()3 


100 590 


14415 


86175 


134 5«') 


II-: 531^ 


20 903 


91 628 


(,2 287 


64 160 


14929 


49231 


^7 959 


28 6oo2 


II 947 


16633 


^'3516 


24 220 


13643 


10557 


3^>43i 


58 120 


— 


— 



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

(3n January the i6th, 1935, President Arturo Alessandri signed a decree 
making the Juan Fernandez Islands (together with Easter I.) a National Park, and 
two (ierman 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 soUte 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. Ikit what happened? Guzman's interesting 
account gives the answer, I.e. p. 37. 

' 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 I kg, and it was very little 
below that weight in 1941. 

* On p. 202 r,u7.iii;'m cjuotes a very different figure: 47238 kg obtained during the period 
Jan. — Aug. The balance cannot, 1 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 



85 




^^S- 99- The central part of Masatierra, seen from the air and showing the cultivations and 

plantations around Cumberland Bay: from left to right V. Colonial, V. Anson and Quebr. del 

Minero. — Photo B. Frodin ^/^ 1952. 

In 1940 the population amounted to 434 persons (225 men and boys), in 

1943 77^ (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 99. 
In 191 7 the lower slopes of Q. del Minero were barren, in 1952 there were plant- 
ations; Anson's valley, where few people lived in 191 7, bears sign of much 
activity, and the aspect of the settlement in the Colonial valley has changed a 
great deal from 19 17 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 



l86 C. SKOTTSHERG 

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 191 7, 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 Rir.CKiEi., "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 ulmifolius), a wellknown pest on the 
mainland, was introduced on purpose to be used as living fences. The success was 
comj)lete. The thruslies 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 {)ersons in all). Their principal occupation is said to be lobster 
fishing. 

Basing his opinion on his [)crsonal knowledge of the islands and on a wealth 
of material from various sources, (Guzman discusses the present situation and the 
{)ossibilities to improve it. There can be no doubt that he is interested in the 
f)reservation 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 r^roduccd he differs cf)nsi(lerably from Johow who did not hold a very 
high opinion of the (juality, 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 I 87 

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 (I.e. 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" (I.e. 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 celebre 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 



I 88 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 Cordon 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 191 7 — better displayed. 



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 



189 




Fig. 100. 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. Backstrom 1917. 



90 



C. SKOTTSHERG 



everjthint;. To-day we can speak with Guzman of the "Crepusculo de la langosta". 
The situation is ahirniinir, he calls for measures to put an end to the decline, and 
he j)roposes certain ways and regulations. I would like to add that in a case as 
serious as this the best thing to do is to {)roclaim the whole year a closed season 
tluring a sufhciently long period. An industry based on the enormous supply 
of fisii, with the bacalao (hg. lOo) 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. 1 shall add one more remark. Cumberland 
l^a\- ought to be an ideal place for a biological station. The fauna and flora, 
terrestrial as well as marine, ofi'ers 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. 

S[)eaking as on behalf of the Chilean nation, I would like to say, with a 
slight alteration, what (iuzman said about the threatened langosta (p. 223): 

\o 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. 

Andkkson, Cj. \V. a new, authentic and complete collection of voyages round the 

world. I-ondon. No date. (Byron's and Carteret's visits to Juan Fernandez.) 
Anson, (). A voyage round the world in the years 1740 — 44. Compiled by Richard 

Wai.tkr. London 1749. 
Branchi, K. C. La Isla de Robinson. Valparaiso 1922. 
Brugcen, M.J. Inmdamentos de la geologia de Chile. Santiago 1950. 
liiRNKv, James. A ( hronological history of the discoveries in the South Sea or Pacific 

()(can. London. Vol. Ill (1813, L'Heremite), IV (1816). 
P.lr(;er, O. Die Rohinson-Inscl. Leipzig 1909. 

Krmki,, a. Line Rcise nach dcr Rohinson-Crusoe-lnsel. Hamburg 1889. 
I'RODIN, B. Kx|)eflition na svenskt uppdrag kartlade sagoon i Stilla havet. (Expedition 

on Swedish initiative map])cd the fairy isle in the Pacific Ocean.) Dagens Ny- 

hetcr 2« ^j 1(^-2. 
(Irak AL\rin, A. l'rol)lenias c( onoinicos de Juan Fernandez. Departamento General 

de la Pro(hu(i6n. ()fi( ina de Enlace Agn'cola. Memorandum 158. Santiago, 

29 dc Ahril de 1944. ('fypcwrittcn.) 
Cramam, Maria. Journal of a residence in Chile during the year 1822. London 1824. 
(iCNitn-.R, E. Derrotero de la costa de Chile. V. Anuario Hidrografico de la Marina 

de Chile. 32. 1920. 
(jfZMAN Parada, J. Cutiihres oceanicas. Santiago. No date. 



A GEOGRAPHICAL SKETCH OF THE JUAN FERNANDEZ ISLANDS I9I 

Hagerman, T. H. Beitrage zur Geologie der Juan Fernandez-lnseln. Nat. Hist, of Juan 

Fernandez and Easter Island. 1: i. 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 nauticas de la costa de Chile. Anuario Hidrogrdfico de la Marina de 

Chile. XX. 1896. 
JoHOw, F. Estudios sobre la Flora des Islas de Juan Fernandez. Santiago 1896. 
Juan, G. y Ulloa, A. de. Relaci6n historica del viage a la America meridional. Ft. 2, 

vol. III. Madrid 1748. 
Lopez, Juan E. Esploracion de las islas esporadicas de la costa de Chile. Anuario 

Hidrografico de la Marina de Chile. II. Santiago 1876. 
Philippi, R. a. Geognostische Beschaffenheit der Insel Masafuera. Neues Jahrb. fiir 

Mineralogie etc. Stuttgart 1857. 
Philippson, a. Die Erosion des fliessenden VVassers 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 jeolOjicas del 

Archipielago (de Juan Fernandez). In: Johow, Estudios. 
QuENSEL, P. (i) Die Geologie der Juan Fernandez-lnseln. Bull. Geol. Inst, of Upsala. 

XI. 1912. 
(2) Additional Comments on the Geology of the Juan Fernandez Islands. Nat. 

Hist, of Juan Fernandez and Easter Island. I: 3. 1952. 
Skottsberg, C. (i) The Wilds of Patagonia. London 191 1. 

(2) Till Robinson-on och varldens ande. Stockholm 191 8. 

 —  — (3) The vegetation of the Juan Fernandez Islands. Nat. Hist, of Juan Fernandez 

and Easter Island. II: 29. 1953. 
SuTCLiFFE, Th. (i) 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 ascensi6n emocionante a la cumbre del Monte Yunque. El Mercurio 

V3 192 T. Valparaiso. 
ViDAL GoRMAz, F. Jeografia nautica de la Repiiblica de Chile. Anuario Hidrografico 

de la Marina de Chile. VII. 1881. 
ViEL, O. Islas de Juan Fernandez. Anuario Hidrografico 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. 
Weber, Hugo. 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. Alberto Graf 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 Guzmiin 
for the same year, 771; both cannot be correct. Ciraf 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 Guzmdn. The average annual 



IQ2 C. SKOTTSBERG 

profit derived from the export of wool was about ^/e of the income from fishing, the 
number of sheej) 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 (iraf 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 
iq^i 16 — 309 — unnecessary. 

Although Ciraf 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 
Tark, 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 tnelanoxylon 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 
lohow in 1892 and by me in 1908 was ordered by the Subdelegado Vera 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 Antennaria (Limacinia) Graf Marin shares my 
opinion that Johow exaggerated its dangerousness. A more serious enemy of the luma 
is an insect, Saissetia 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 1 20 000 (considerably 
more to-day that 10 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 (iraf Marin's interesting report materializes, Juan Fernandez will stand as a unique 
e\am|)lc of a National Park, the home of a unique fauna and flora, offered for sale. 



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

By 
C. SKOTTSBERG. 

With I Map. 

Part 1. 
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. 

I. 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 Degeneriaceae, 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 Juncus may not be truly indigenous), Rubiaceae 5 (4 gen ), 

13 "~ 557857 The Nat. Hist, of Juan Fernandez and Easter Isl. Vol. I 



194 ^- 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 (i), (iunncraceae 3(1), Halorrhagidaceae 3 (i), Berberidaceae 2 (i), 
Ikomeliaceae 2 (2), Convolvulaceae 2 (2), Labiatae 2 (i), Leguminosae 2 (i), 
Plantaginaceae 2 (1), Rutaceae 2 (i), Scrophulariaceae 2 (2); represented by i 
species only: Aizoaceae, Horaginaceae, Callitrichaceae, Empetraccae, Ericaceae, 
luipliorbiaceae, Flacourtiaceae, Iridaceae, Lactoridaceae, Loranthaceae, Palinae, 
Ranunculaceae, Rliamnaceae, 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 [Azorella\ Caryophyllaceae, Cruci- 
ferae, Labiatae, Leguminosae (Adesniia, Cassm), Plantaginaceae, Scrophulariaceae 
[Calceolayia], Horaginaceae [Plagiobotrys and related genera), Euphorbiaceae, Iri- 
daceae [SisyrijuJiiuui], Ranunculaceae, Rhamnaceae, Verbenaceae. The absence 
of such families as Amaryllidaceae, Asclepiadaceae, Eagaceae, Geraniaceae, Lau- 
raccac, Liliaceae, Xolanaceae, 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, Table 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 difi"erence between 
them is [)artly explained by the difi"erence in altitude and thereby in climate — 
Masafuera has an ali)ine 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. '1 he difference becomes particularly obvious when endemism is considered. 

Of the 89 genera, 17 (19 ?{,) are endemic; of these 12 (70.6 %) are confined 
to .\L'isatierra, i (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 1 34. 3 "„), the corresponding figures for Masafuera are 18, I and 5.5 % 
and, for the genera occurring on both islands, 36, 4 and 1 1. 1 %. These figures 
serve to illustrate the great difference in the distribution of the endemic genera 
commonly looked upon as ref)resenting the most ancient element among the 
Angiosperins in Juan h'ernandez. 

Ot the total number of species, 147, 99 are found on Masatierra, 9 on Santa 



DERIVATION OF THE FLORA AND FAUNA 



195 



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 tugurionim R. Br 

Cardamine chenopodiifolia Pars 

— 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 coUina Phil 

DENDROSERIS litoralis Skottsb 

— macrantha (Bert.) Skottsb 

— macrophylla D. Don 

— marginata (Bert.) Hook, et Arn 



t 


rt 
rt 


t 








(U 


u 


% 








nS 


cj 




S 


a 


(§ 


% 


^ 


S 



196 



C. SKOTTSBERG 



j Dichoiulra repens l-'orst 

! Drim\ s confertifolia Phil 

Dysopsis hirsuta (Muell. Arg.) Skottsb. . . . 

Kleocharis fuscopurpurea (Steiul.) 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 Callcottiae Hook, et Am 

Euphrasia formosissima Skottsb 

Fagara externa Skottsb 

— mayu (Bert.) li^ngl 

Galium masafueranum Skottsb 

Gnaj)halium spiciforme Sch. Bip 

Gunncra bracteata Steud 

— • Masafuerae Skottsb 

— peltata Phil 

Halorrhagis asperrima Skottsb 

— masafuerana Skottsb 

— masatierrana Skottsb 

I Hedyotis thesiifolia St. Hil 

Hesperogreigia Herteroi Skottsb 

HESPEROSERIS gigantea (Johow) Skottsb. 

JUANIA australis (Mart.) Drude 

JuiK us capillac ens I. am 

- — dombev.'iuus (iay 

— imbri(atiis Laliarpc 

— l)lanif..lins K. I',r 

— ])ro(cnis I'.. Mvv 

Kock'ria iiii( ratlicra ( Dcsv.) Griseb 

LACTORIS fernandeziana Pliil 

Lagcnopliora nari..1i Iraiu h 

Libcrtia formosa C.rali 



1 



a 


2 

1 


_ 


+ 


— 


+ 


— 


+ 


- 


+ 


- 


+ 


- 


+ 


- 


+ 


- 


4- 


_ 


+ 


- 


+ 


- 


+ 


- 


+ 


- 


■+ 


- 


+ 


- 


+ 


- 


+ 


- 


+ 


- 


+ 


- 


+ 


- 


+ 



DERIVATION OF THE FLORA AND FAUNA 



197 



Lobelia alata Labill 

Luzula masafuerana Skottsb 

Margyricarpus digynus (Bitter) Skottsb 

MEGALACHNE berteroniana Steud 

— masafuerana (Skottsb. et Pilger) Hubbard ms. . . . 

Mimulus glabratus H. B. K. forma 

Myrceugenia Schulzei Johow 

Myrteola nummularia (Poir.) Berg 

Nertera granadensis (L. fil.) Druce var 

Nicotiana cordifolia Phil 

NOTHOMYRCIA fernandeziana (Hook, et Arn.) Kausel 

OGHAGAVIA elegans Phil 

Oreobolus obtusangulus Gaud 

Parietaria humifusa Rich 

Paronychia chilensis DC 

Peperomia berteroana Miq 

— fernandeziana Miq 

— margaritifera Bert 

— Skottsbergii C. DC 

Pernettya rigida DC 

PHOENIGOSERIS berteriana (Dene) Skottsb 

— pinnata (Bert, ex Dene) Skottsb 

— regia Skottsb 

Phrygilanthus Berteroi (Hook, et Arn.) Reiche 

Piptochaetium bicolor (Vahl) Presl 

Plantago fernandezia Bert 

— truncata Cham, var 

PODOPHORUS bromoides Phil 

Ranunculus caprarum Skottsb 

REA micrantha Bert, ex Dene 

— neriifolia Bert, ex Dene 

— pruinata (Johow) Skottsb 

Rhaphithamnus venustus (Phil.) B. L. Robins 

RHETINODENDRON Berterii (Dene) Hemsl 

ROBINSONIA evenia Phil 

— gayana Dene 

— gracilis Dene 

— Masafuerae Skottsb 

— thurifera Dene 



03 




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03 


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- 


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- 


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- 


- 


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+ 


- 


+ 


+ 


- 


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+ 


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- 


- 


+ 


+ 


- 


+ 


+ 


- 


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+ 


- . 


— 


- 


- 


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+ 


- 


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+ 


- 


- 


+ 


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- 


+ 


+ 


- 


+ 


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- 


- 


- 


+ 


+ 


- 


- 


+ 


- 




+ 


+ 


- 


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+ 


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+ 


- 




+ 


- 


- 


+ 


— 


— 


- 


- 


+ 


+ 


- 


- 



198 



C. SKOTTSBERG 



Riibus geoides Sm j 

Salicornia fruticosa l\unth j 

i Santalum fernandezianum F. Phil j 

Scirpus cernuus \'ahl 

j — nodosus Rottb ! 

I SELKIRKIA Berteroi (CoUa) Hemsl 

I Solanuni fernandezianum Phil 

! — masafueranum Bitter et Skottsb 

— robinsonianum Hitter 

; Sophora fernandeziana (Phil.) Skottsb 

I — masafuerana (Phil.) Skottsb 

I Spergularia confertiflora Steud 

I — masafuerana Skottsb 

i Stipa (Xassella) laevissima (Phil.) Speg 

— neesiana Trin. et Rupr 

SYMPHYOGHAETA macrocephala (Dene) Skottsb 

i 
Tetragonia expansa Murr 

: Trisetum chromostachyum Desv 

; Ugni Selkirkii (Hook, et Arn.) Berg 

j Uncinia brevicaulis Thouars 

i — costata Kuekenth 

— Douglasii Boott 

— phleoides Pers 

— tenuis Poepp 

Urtica fernandeziana (Rich.) Ross 

j — Masafuerae Phil 

j Wahlenbergia Berteroi Hook, et Arn 

j — fernandeziana (A. DC. p. p.) Skottsb 

j — Grahamae Hemsl 

— Larrainii (Colla) Skottsb 

i — Masafuerae (Phil.) Skottsb 

I YUNQUEA Tensii Skottsl) 



Clara, and 74 on ^Ma.safuera; 6<S - 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 Ma.satierra 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 °o are reported from both Masatierra and Masafuera, a surprisingly small 



DERIVATION OF THE FLORA AND FAUNA 



199 



Table II. 
Distribution of the endemic genera. 



Centaurodendron 
Cuminia . . . . 
Dendroseris . . 
Hesperoseris . . 

Juania 

Lactoris . . . . 
Megalachne . . . 
Nothomyrcia . . 
Ochagavia . . , 
Phoenicoseris . . 
Podophorus . , 

Rea 

Rhetinodendron , 
Robinsonia . . . 
Selkirkia . . . . 
Symphyochaeta , 
Yunquea . . . . 



I + 



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), loi are endemic = 68.7 %; of the 99 
species found on Masatierra, 66 (66.'j %) 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 (lOi), 50 are, as we have seen, confined to Masatierra (49.5 %), 3 (3 %) 
to Masatierra + Santa Clara, i (i %) to Santa Clara, 34 (33.7%) to Masafuera, 
12 (11.9%) common to Masatierra and Masafuera, and i (i %) 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 i (2.2 %), only on 



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 Joiiow. In his table 
"Continjente B, I^^species autoctonas, pero no endemicas", pp. 221-222, he men- 
tions several species regarded as doubtful natives: 3 species of Gnaphaihwi, Mimu- 
liis, Solanum furcatian ( = robinsonianum), Tetragonia cxpansa, Spergularia rubra 
(= confertiflora), Parietaria )iuiuifusa\ Trisetum c/iroifwstachyum and DantJioiiia 
colliiia. With the exception of GnapJialiian spp. I have listed them as native. 
There seems to be little reason to regard Tetrago7iia, a wide-spread thalassocho- 
rous plant, as anthropochorous, and the Spergularia is known only from Juan 
Fernandez and Desventuradas Is. Danthoiiia gave me the impression of being 
just as autochthonous as PiptocJiactiuin and the two species of Stipa, and I find no 
good reason to exclude either Mimulus, Parietaria or Trisetum (this never seen 
by us). With regard to Solanum "'furcatimi' (not furcatum of Dunal) I share 
Joi row's doubts. It was described by BlTTER as vS*. robi^isonianum, a Morella micro- 
species of 5. nigrum L. coll. and found on all three islands. This assemblage 
as represented on the mainland has never been seriously studied. Possibly vS". robifi- 
sonianum is a case of the same kind as the dandelion discovered on the islands 
in 19 1 7 and described as Taraxaaim ferna7idezianum 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 JOIlOW's 
list I have excluded Bahia, Amblyopappus, Erythraea, Monocosmia and Phalaris. 
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: J uncus capillaceus and pla7iifolius, Ce7itella and Hedyotis, perhaps also Paro- 
7iycJiia. 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 Erigero7i with 6, yu7icus, Robi7iso7iia, U7ici- 
7iia and \Vakle7ibergia with 5, and De7idroseris and Pepero77iia 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. (Bkws), widespread both hemisph., subtrop.-temp. 

7ieesia7ia Trin. et Rupr. Mcx. to Boliv. and Chile, Braz., Argent., Urug. Poly- 
morphous (^^p. 771). 

Iae?rissi7na (Phil.) Speg. Peru, Chile, Argent. With a number of S. Amer. sp. 
referred to a separate genus [Xassella). 



DERIVATION OF THE FLORA AND FAUNA 



Piptochaetium Presl. 20; N. Amer. to extratrop. S. Amer. 

bicolor (Vahl) Desv. Chile: Valdivia; Braz., Argent. 
Podophorus bromoides Phil. In 22y I referred to Brachyely thrum 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 Brachyely thrum is disjunct after a well-known pattern: B. erectum (Schreb.) 
Beauv., N.E. Amer., and B. japonicum Hack, ex Honda, China, Korea, Japan. 
PiLGER (/p^) brings this genus to Festuceae-Festucinae next to Aphanely thrum, 
placing Podophorus with the Brominae. 
Chaetotropis Kunth. 2 (?3). 

imberbis (Phil.) Near the following. 

chilensis Kunth. Peru, 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 ipy). This is recorded from 
Magell., Falkl., N. Zeal. (55). F. B. H. Brown (j5. I. 84) says that his A. rapensis 
from Rapa Id. is "very closely allied" to magellajiica, 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. 

collina 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., better oniana Steud. and masafuerana 
(Skottsb. et Pilg.) Hubbard ms. Reduced to Bromus by PiLGER {iQj), but restored to 
generic rank in his posthumous paper on the system of the Gramineae [igS). 
Chusquea Kunth. About 100 (.?) sp. Mex. to Argent, and S. Chile. 

fernandeziana Phil. Related to Chilean species. 

Cyperaceae. 

Cyperus L. A world-wide, essentially tropical genus of about 600 sp. (KuKEN- 
THAL). 

er agrostis 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 

cermius \'ahl. Subcosmopolitan and of variable habit; south to Fueg. and 
Falkl. 
FJeocharis R. Br. A world-wide genus of about 150 sp. (Svenson). 

fiistopiirpurea (Steud.) H. Pfeifif. S. Chile. Listed before as a subspecies or 
variety of //. 7>iaculosa (Vahl) R. Br. (W. Ind.-Braz.). 

Oreoholus R. Br. An austral-circump. genus of 6 sp., extending north to Malaysia 
and Hawaii. 

obtusaugiilus Gaud. Andes of Colomb. and Ecuad. to Fueg., Falkl. 
Cladium R. Br. 48 (Ki kkxtii.\l); widespread, trop.-subtrop. with preponderance 
S.I'^. Asia-X. Guin.-Austral.-X. Zeal., north to Hawaii, W. Ind., Braz. 2 boreal sp. 

scirpoideum (Steud.) Benth. et Hook. f. Nearly related to C. angtistifolium 
(Gaud.) Benth. et Hook. f. (X. Guin., Tahiti, Hawaii). 

rnciuia Pers. 32; circump.-austr.-subantarct. with outposts north of the P^quator 
in Centr. Amer. and Philipp. Is.; greatest concentration of species in N. Zeal. 

brevicaulis Thouars. S. Chile to Fueg., P'alkl.; Trist. da C; St. Paul and 
Amsterd. Is. 

Douglasii Boott. 

costata Kuekenth. Related to the former. 

pJihoidcs 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. 

Haiiksii Boott. Andes of Centr. Chile to P'ueg. Belongs to the boreal section 
I^Vigidae-P^uliginosae of KlKENTIlAL. 

berteroniana Steud. Sect. Echinochlaenae of KCkenthal, 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 Dkl DE, but perhaps nearer to the Morenieae, or possibly 
regarded as type of a separate subtribe. HUTCIIINSON brought Juaiiia next to 
Ctroxylo}!, following Bextiiam and HooKER, Croizat (7/) remodelled Morenieae, 
including Jutuiia and the Mascarene IlyopJiorhe, otherwise linked with Chamae- 
doreae. 

Bromeliaceae. 
Ilcsperogreigia Berteroi Skottsb. 2 or perhaps more sp. Subtrop. Andean, related 
to (ireigia («S sp., Costa Rica, Venez.-S. Chile). LOOSER called Hesperogreigia 
monotypical, because my {)ai)er (2^0) had escaped him; if not, he would not have 
written "^'Ouiza dc la afinidad de Greigia.^" [ijo. 291). Inspired by L. B. SMITH 
he sup{)ressed I fispcrognigia 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 irrej.;ular, 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 Rliodostachys\ see 22g. iio 
and 24g. jy^. 

Juncaceae. 

Luzula 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. 

Junciis L. Mainly temp.-subtrop.; about 225. Well represented S. hemisph. (about 
50), especially Austral, and N. Zeal., less so temp. S. Amer. 

imbricatus Laharpe. Ecuador-S. Chile, Braz., Argent., Urug. 

capillaceus Lam. Ecuador, Centr. Chile, Braz., Argent., Urug. Introduced? 

procerus E. Mey. S. Chile. 

dombeyanus Gay. Peru, Chile, Braz., Argent., Urug. 

plaiiifolius R. Br. S. Chile, Austral., Tasm., N. Zeal. Introduced.^ 

Iddaceae. 

Libertia Spreng. 5 sp., 3 Chile, 2 N. Zeal. 
formosa Grah. S. Chile. 

Piperaceae. 

Peperomia Ruiz et Pav. Possibly over 1500; pantropical. 

berteroana Miq. Extremely like P. tristanensis Christoph. from Gough I.; 
see 24^. 

margaritifera Bert, ex Hook. Possibly related to P. Reineckei C. DC. (Samoa). 

Skottsbergii C. DC. Allied to the former. 

These species form a separate subgenus Tildenidium Skottsb. (^^7), which 
seems to stand closer to palaeotropical than to neotropical groups. P. berteroayia 
and tristanensis show affinity to P. urvilleana A. Rich. (Austral., N. Zeal., Lord 
Howe I., Norfolk I.) and P. Vente7iati Miq, (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 [/. 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 



Parictaria L. Some 30 sp.; in all parts of the globe. Several medit. -orient., others 
N. and S. Amer., one [P. debilis Forst.) supposed so be widely distributed. 

/luniifiisa Rich. Chile. Until now listed as debilis which, in Weddell's mon- 
ograph, is a collective species. I have shown [2^1) 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 debilis. Pending a revision of the genus, Jinmifusa is a cor- 
rect name for the Chilean plant. 

Loranthaceae. 

Phrygilantlius hjchl. About 30; bicentric: Mex.-Braz. and Chile, Argent., Urug.; 
Austral., X. Zeal., X. (kiin., Philipp. Is. 

Berteroi (Ih^ok. et Arn.) Reiche. Related to Andean sp. 

Santalaceae. 

Sautaluui L. 18; Austral. ~X. Guin. 5, Melan. 2, Polyn. 2, Hawaii 7, Bonin i, and 
the following. See 2?^ and 270. 

fernandezianutn ¥. Phil. P^xtinct. Belongs to sect. Polynesica Skottsb.; not, 
as has been said (///), related to S. freycinetiaimni of Hawaii which belongs to a 
different section {2'/6). 

Chenopodiaceae. 

Clie)iopodiu})i L. Over 250; world-wide, but essentially temp. 

Sanctae Clarae Johow, crusoeanum Skottsb. and nesodendron Skottsb. On 
my request Dr. P. Aei.LKX, who created sect. Skottsbergia to receive the three 
island endemics, sent me the following remarks: "Ich wiirde sagen, dass die drei 
Arten zu kciner der uns heute bekanntgewordenen Arten irgendwelche verwandt- 
schaftliche Zijge und l^eziehungen aufvveisen. P^s sind vollig isolierte Typen, Re- 
Hkte eines im Meere versunkenen Plorenreiches." Evidently Dr. Aellen does 
not support the idea that they are related lo pajiiculatum Hook. (X. Amer., Peru- 
Chile) or oahuejise Meyen (Plawaii), which latter has the same arboreous habit. 
Salicorjiia L. Widely distributed, about 30 sp. 

fruticosa L. (peruviana Kunth). Taken in a wide sense Mediterr., S. Afr., W. 
Ind., Polyn.; peruricvia, \V. coast of S. Amer. to S. Chile. 

Aizoaceae. 

Tetragojiia P. About 25 s[). Strongly represented in S. Afr., several sp. in Chile. 
expajisa Murr. Coasts and islands of the Pacific; Braz. 

Caryophyllaceae. 

Spergularia Presl. A widely scattered genus of about 40 sp.; numerous in Chile. 
eojifertijioya 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. 
chilensis DC. Centr.-S. Chile, S. Braz., Argent. 

Ranunculaceae. 

Ranunculus L. Probably over 300; world-wide, particularly boreal, numerous sp. 
N. Zeal. 

caprarum Skottsb. Apparently with distinct relations in New Zealand, not 
near boreal or S. American species (22^. 125; jyj). 

Berberidaceae. 

Berberis L. 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., i 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 Piperales. GUNDERSEN 
(/20) asserts that it belongs to the latter, but it differs from this in very important 
characters. 

Cfuciferae. 

Cardamine L. About 130; world-wide, essentially temperate, many Chilean sp. 
chenopodiifolia 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. [ij^]- 

Rosaceae. 

Rubus L. A very large, temp, and trop. -montane genus, richly developed in the 
N. hemisph. 

geoides Sm. S. Chile to Fueg., Falkl. 
Margyricarpus Ruiz et Pav. 4 or 5, trop. Andes to Patag. 



2o6 C. SKOTTSBERG 

digynus (Bitter) Skottsb. Near J/, pinnatus (Lam.) O. K. (Peru, Chile, S. 
Hraz., Argent.). 

Acaoia L. An austral genus of over lOO sp., the majority in S. Amer.; 12 
Austral. -X. Zeal., single sf). Calif., Hawaii, the Cape, etc. 

masafuerana I^itter. Near A. antarctica Hook. f. (Magell., Fueg.). 

oralifolia Ruiz et Pav. subsp. australis l^itter. S. Chile to Fueg., Falkl. Per- 
haps accidentally introduced to Juan P"ernandez. 

Leguminosae. 

Soplioni L. 70-80, Old and New World; sect. Tetrapterae ("Edwardsia") austral- 
circump.; 17 closeh' related species. 

fernandeziana (Phil.) Skottsb. and masafuerana (Phil.) Skottsb. Very near 
the Chilean ''tetraptera^ [S. iiiacuabiaua Phil.), not identical with vS". tetraptera 
Ait. from X. Zeal. 

Rutaceae. 

lui^s;ara L. Over 200, pantrop.; a large palaeotrop. group well represented in the 
S. Pacific (Austral., X. Caled., Polyn.), extending north to Hawaii. The two Juan 
I-'ernandez species F. mayu (Bert., Hook, et Arn.) Engl, and externa Skottsb., 
which are very closely allied, form the sect. Mayni Engl. Related to W. Pacific sp. 

Euphofbiaceae. 

Dysopsis Baill. 3; i in Pxuador \D. paucidentata (M. Arg.) Skottsb.], i in S. Chile 
\D. gleclio})widcs (Rich.) M. Arg. p. p.], and the following. 
hirsiita (M. Arg.) Skottsb. 

Callitrichaceae. 

Calliiriche L. Widely distributed in the N. hemisph., austral-circump., south to 
Austral, and S. Chile but absent from the southern half of Afr. 
LechUri (Hegelm.) Fassett. S. Chile to Magell. [24^. 781). 

Rhamnaceae. 

Colli tia C'onim. 17, Andean and extratrop. S. Amer. 

spartioides Ik-rt. ex Colla. A well-marked species, related to Chilean and 
other S. Amer. forms. 

Flacourtiaceae. 

Az(ir(7 Ruiz et Pav. 20~2 1 sj).; 18-19 Chile, i Braz., I Argent. 

fernandeziana (iay. Closely related to A. scrrata Ruiz et Pav. (S. Chile). 

Myrtaceae. 

r^'-;// 'I'urcz. 15, Mex., Centr. Amer., Andes, south to Chiloe, Venez., Braz. 

Selkirkii (Hook, et Arn.) Berg. Probably related to species in Centr. Amer. 
and \'enez., perhaps also to f\ Ccvidollci (Barn.) Berg from Chile; not very close 
to r. Molijiae Turcz. (S. Chile; Kauskl in litt.). 



DERIVATION OF THE FLORA AND FAUNA 207 

Myrteola Berg. 12, Colomb.-Chile. 

mimmularia (Poir.) Berg. Chile, Cord. Linares to Fueg., F'alkl. 
Nothomyrcia Kausel. 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 Chiloe. 

Schulzei Johow. Related to M. planipes (Hook, et Arn.) Berg (S. Chile). 

Gunneraceae. 

Gunnera L. 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 F'alkl.; 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. kaMaie7isis Rock 
than to S. Amer. species. 

Halorrhagidaceae. 

Halorrhagis 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. 

asperritna Skottsb., masatierrana Skottsb. and masafuerana Skottsb. belong 
to Subsect. Cercodia and are closely related to H. erecia (Murr.) Schindl. (N. Zeal.) 
and other Austral, and N. Zeal. sp. All J. Fern, forms were formerly incorrectly 
identified with erecia. 

Umbelliferae. 

Cejttella L. 29; 19 Afr., Madag., i China, 2 Austral., 5 trop. Amer. and the 
widely dispersed C. asiatica (L.) Urb. 

triflora (Ruiz et Pav.) Nannfeldt. Centr. and S. Chile, formerly included under 
asiatica. Introduced .r* 

Eryngium L. 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. Fruticosa, but differ mainly in being arborescent from 
the species occurring on the mainland. 
Apium L. About 30; in all parts of the world. 

fernandezianutn Johow. A well-marked species, probably not nearly related 
to the Chilean species, but showing some affinity to A. prosiratiini 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.; i Galapag. Is., 2 N. Zeal., 2 Tasm. 
rigida (Bert.) DC. A well-marked sp. [2_$2). 



2o8 C- SKOTTSBERG 

Empetraceae. 

Kmpctruin L. A bipolar genus, E. ?iij{runi I., north, /:. rubrum Vahl south, but 
according to Gool) the latter is represented in the Subarctic by special forms. 
See _v^. 781. 

rubriun Vahl. Andes of S. Chile and Argent, to Fueg. and Falkl.; Tristan da C. 

Convolvulaceae. 

Dichondra Forst. 5 trop. -subtrop. Amer., i X. Zeal., and the following. 

)cpe)is Forst. Widely spread over both hemispheres, north to N. Amer. and 
China, south to S. Chile and N. Zeal. 
Calystegia R. Hr. 10-20; temp. -subtrop. in all parts of the world. 

tugiiriorum R. Br. S. Chile (Hantelmanni Phil.) and N. Zeal. 

Boraginaceae. 

Selkirkia Berteroi (Colla) Hemsl. According to Johnston (148) very near Hackelia 
()j)iz, a genus centering in western N. Amer., with outposts in S. Amer. and 
I'Airas.; //. revoluta (Ruiz et Pav.) Johnst. Peru to Boliv. and Argent. 

Verbenaceae. 

Khaphitliaiinius Miers. 2, one in Centr. and S. Chile. Near CitJiarexylon 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 Prasioideae by El'LlNG (go), a 
palae()tr()j)ical group best developed in Hawaii, but whereas Cumiuia has the drupe 
of this tribe, it has the corolla of Stachyoidcac-Me7itJiiuae, where BRIQUET placed 
it; lUkciKKS statement that the flowers are "Lippenbliiten" (./j. 23) is erroneous. 
fernandezia Colla and eriantha Benth. Perhaps united by intermediate forms. 

Solanaceae. 

Sola)iu))i L. Prcjbably over looo; in all parts of the world, richly represented in 
trop. Amer, man\- in Chile. 

fernandezianiitn Piiil. Related to .S". tKhcrosuui L. coll. 

masafiieranutn Hitter et .Skottsb. A \ery well-marked Morella. 

robinsonianutrj Bitter. .See above j). 200. 
XicotidJKJ L. 60; 45 Anicr., C"alif.-Mex., And. S. vXmer. (hxuad.-Peru-Chile, ]3raz.- 
Argent.-Patag.), 15 Austral, (i \. Caled., etc.). 

coydifolia IMiil. Belongs to the Rustica group, confined to trop. Amer. and 
Australas., and related to X. Raijiioiidii ^lacbr. (Andes of S. Peru), solaJiifolia 
\\'al[). (.\. and Centr. Chile) and piviiculata L. (Perii, \. Chile). Dr. GooDSl'KKD 
ex[)resscd his opinion (in litt.) that the Rustica complex originated in the region 
now occupied by western ]^oli\ ia and l\>ru and extended to the "J"^'^ Fernandez 
land", becoming isolated during the final uplift of the Andes or possibly even earlier. 



DERIVATION OF THE FLORA AND FAUNA 209 

See also 112. F. B. H. Brown (jj. III. 262) suggested an affinity between cordifolia 
2ind fatuhivensis F. B. H. Brown (Marquesas), but they belong to different subgenera 
and the latter is referred to N. fragrans Hook, from N. Caled. as a variety {112). 

Scfophulariaceae. 

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 {114). 
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 y8.22/\. 

formosissima Skottsb. The fact that this species is very unlike the Trifidae 
of Chile makes it particularly interesting. In 22g. 169 I emphasized the differ- 
ence between formosissima and the Australes of N. Zealand and placed it nearer 
to the boreal Semicalcaratae. Wettstein, in his contribution to my paper (I.e. 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 RiETZ 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. priiiceps Cham, et 
Schld. of Hawaii. Both belong to sect. Palaeopsy Ilium, scattered over the S. hemisph. 
and extending north to N. Amer., Hawaii and S. Eur.: N. Amer. 5, S. Amer. i, 
S. Eur. 2, Afr. 5, St. Helena i, Madag. i. Lord Howe I. i, Auckl. Is. i, Rapa 2, 
and Hawaii 9. 

iruncata 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. 

Hedyotis L. A large pantrop. genus. 

thesiifolia St. Hil. Trop.-subtrop. S. Amer., in Chile south to Chiloe. 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 24.4, 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 



2IO C. SKOTTSBERG 

buted from Australia to Malays., Melan. and Polyn. with a secondary centre in 
Hawaii (i8); entirely absent from Amer. 

Hookeri (G. Don) \V. R. B. Oliver {1^2). Forms a separate monotypical section. 

pyrifolia (Hook, et Arn.) Skottsb. Belongs to a section of 8 Polyn. sp. (4 
Tahiti, i Cook Is., i Rapa, 1 Pitcairn, i J. Fern.). 

CaliiDH L. World-wide, essentially boreal. Over 500 have been described, almost 
^'4 of these Medit.-Orient.; about 50 in S. Amer., mostly along the Andes and 
extending south to Fueg., Falkl. and S. Georgia. 

masaftierantim Skottsb. Related to C criocarpuin Bartl., DC. and tricJiocar- 
pu))i DC. (both Coquimbo-Cord. Linares), the fruit as in masafueraimm covered 
with straight bookless hairs. 

Campanulaceae. 

Wahlciibergia Schrad. l^ssentially S. African; about 230 sp. (9 S. Amer., 10 S. 
Fur.-Oricnt.. 2 or 3 St. Helena, 20 trop. Afr., 6 Madag.-Mascar., 150S. Afr., 9 As., 
I \. (iuin., 8 Austral., 8 N. Zeal, 5 J. Fern,). 

Larrainii (Bert, ex Colla) Skottsb., fern andeziana 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 /['. Masafuerae and a couple of perennial S. African species such as 
//'. lickloiiii Buek and oxypJiylla 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 
tiiey arc 3. Hkmsi.ev {i2y. 61) regarded the St. Helena species as allied to African 
and Juan I^Y'rnandez species. 

Lobelia L. 350-370; particularly numerous in Amer. and Afr., less so in As., 
Austral, and Oceania, 2 in Juir. 

alata Labill. S. Chile, S. Afr., Austral. A sea-side sp. 

Compositae. 

Lagenop flora Forst. A biccntric austr.-subantarct. genus of 16 sp., most numerous 
in Austral. -N. Zeal.; Fiji, Rapa, extending north to Philipp. Is. and Hawaii. 

JIario/i I^Vanch. Andes of S. Chile to Fueg. 
]-j-io;ero}i L. A large bor.-temp. genus. Ind. Kew. lists about 700 sp. as valid, 
half of them in X. Amer. and about lOO in S. Amer., where the genus is richly 
developed along the Andes, south to Patag., Fueg. and Falkl. Not few are known 
from tro{). mountains in the W. and P2. hemispheres; very few reported from 



DERIVATION OF THE FLORA AND FAUNA 211 

Australia. One species is found on Rapa, related to the Juan Fernandez species 
according to Brown [jj. 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. 

ViERllAPPER (see 22p. 1 82) suggested that the island species are allied to 
Andine species, but also that E. fruticosus comes very close to E. lepidotus Less. 
of Hawaii; this is, however, now referred to Tetramolopiu7n by SllERFF. There is 
no Erigerori in Hawaii. 
Gnaphalium L. A large subcosmopol., essentially temp, genus needing revision. 

spicifonne Sch. Bip.; comp. 22g. 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 e.\t\\Qr americanum W\\\., ptirpureum L., 
spicatuin Lam. or mucronatum Phil, which, together with consanguineuin Gaud., are 
regarded as forms of a single polymorphous taxon. 

Abrotaiiella 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 
I in Austral., 2 Tasm., i N. Guin., i Rodriguez I., and the following. 

crassipes Skottsb. Very near A. Moseleyi Skottsb. nom. (W. Patag.); see 
22g. 189-190. 

Robinsonia DC. Bentiiam (20.460) remarks on Robinsonia 2iW^ Rhetinodendroni 
"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 Dubautia and Raillardia, but they are not dioecious. Re- 
cently an undisputable relative was discovered in New Guinea, Brachionosiylunt 
Mattfeld (j/c?. 27-28) : 

Von Senecio, dessen pacifische Arten unserer Pflanze recht nahe kommen, unter- 
scheidet sich diese Gattiing wesentlich durch die Zweihausigkeit . . . und waiter 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 Robinsonia 
und Rhetinodendron 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 verhaltnismiissig sehr 
breiten, etvvas vorspringend gestreiften Zungen, die Form der durch ein kleines Be- 
cherchen gekronten 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 driingt. ... So bleibt als einziger Un- 
terschied der durch die sehr verschiedene Blattform bedingte habituelle Findruck. . . . 

1 Described as a variety of the former and now raised to specific rank. 



212 C. SKOTTSBERG 

Die Rliitcnkopfe sind bei Robinsonia erheblich zahlreicher und kleiner, kurz und breit 
glockenformig, bei unserer Pflanze aber schmalglockig. 

Wiirden diese Sippen demselben pflanzengeographischen Gebiete angehoren, so 
wiirde man sie sicher nicht generisch trennen konnen. Was hier aber wesentlich dazu 
zwingt, ist, dass die Zweihiiiisigkeit, die ja hier die wesentliche Ubereinstimmung be- 
dingt, bei den Kompositen zu den verschiedensten Malen in den verschiedensten und 
auch in denselben (iru})pen entvvickelt wiirde und daher keineswegs als Kriterium 
fur eine generische Verwandtschaft angesehen werden kann. 

To this I shall remark that nobody would think of uniting Rohhisonia and 
Rlicti}wdeudron, because they differ in essential floral characters. Brachionostylmn 
is, to judge from the description and plate XC: B (only ? known), closely related 
to Robi)iso7iia but not to RJietinodendron. The mode of growth is not quite the 
same, for in Robinsonia (as well as in Rheti7iodendroii) 2 or 3 innovations are 
developed, and none of them continues the mother axis. Besides, Brachionosiylimi 
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 Robinsonia. 

gayana Dene and thurifera Dene are a pair of related species. 

evenia Phil, and Masafuerae Skottsb. form another pair. 

gracilis Dene stands more isolated. 
Symphyochaeta (DC.) Skottsb. See 2^g. 785. 

macrocephala (Dene) Skottsb. 
Rhetinodendron Aleisn. Berterii (Dene) Hemsl. See above. 

Centaurodendron dracaenoides Johow. Not, as has been supposed, just an ar- 
boreous Centaurca, but differing materially in flower structure [24J). 
Yunqiiea Tenzii Skottsb. See 2jy. 163. 

Dendroseris C. Don. An isolated genus, perhaps distantly related to TJiavmo- 
sevis Phil, from Desventuradas Is., but probably not to Fiichia Hook. fil. Ben- 
lllAM 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 Dendroseris s. str., and also 
of PJioenicoscris, whereas they are of a rather normal type in Rea and Hespe- 
roscris. All are devoid of paleae, which are present in Tlianinoseris and Fitchia, 
a distinctive feature pointed out by 1^p:ntiiam, but the awned achenes of /7/f///^ 
differ from those of all the other genera mentioned. In spite of the profound 
discrepances in sucli im[)ortant characteristics as inflorescence, involucre, recep- 
tacle and stigma Tlianinoseris has more in common with the Dendroseris as- 
semblage than with other genera. 

macrophylla \). Don, macrantha (Bert, ex Dene) Skottsb., margitiata (Bert, 
ex Dene) I look, et y\rn. and litoralis Skottsb., all lumped together by JOIIOW, are 
well-marked s{)ecics; comp. 22^1.201-204. 
Phoenicoseris Skottsb., 2./9. 787. 

pinnata (Bert, ex Dene) Skottsb. and fter^er/ana (Dene) 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 24^. 788. 

neriifolia Dene. Little is known of this, but it is undoubtedly a very good 
species; micrantha Bert, ex Dene and pruinata (Johow) Skottsb. are near rela- 
tives but good species (comp. 22g. 207). 
Hesperoseris Skottsb. gigantea (Johow) Skottsb. See 24p. 788. 

Vicarious species. 

VlERHAPPER (2yd) 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 Myrceuge7iia Schulzei, 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 VlER- 
HAPPER were called "substitute species". 

WULFF [2gi. 66-62) devotes considerable space to a discussion of vicarism. 
He agrees with VlERHAPPER: 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 (I.e. 276) calls polytopic species, i.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 {ijo. 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 ^' SKOTTSBERG 

These were (the species from Masatierra mentioned first): Dendroseris niicrantha- 
:gigantca (belong to different genera), WaJiloibergia Bcrteroi-tuberosa {= Masa- 
fuerae, not very closely related), JMyrceugcnia fernandezia7ia-Sc)iulzei {^\^^x^x\\. 
genera), liryngium buplcuyoides-sarcopJiyllujn (only distantly related), Cardamiue 
alsophila ( = Jlaccida) - Kruessi'/ii{v\ot nearly related), and Urtica glo^neruliflora ( = fer- 
7iandt'zia)ia)~Masafucrae (belong to different sections). There is not among them 
a single example of either vicarism or pseudo-vicarism in the sense of ViERllArPER. 
In 1914 (-?-?/) I distinguished 4 pairs of vicarists in Juan Fernandez: /^<f;/^/r^- 
stris inicraiitha-gigajitea, Myrceugenia fcruandeziana-Schulzei, Giinnera peltata- 
Masafuerac, and Pepcrouiia luargaritifera-Skottsbergii; 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 I am not prepared to argue that they arose out of one 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 \icarious or substitute: Berberis corymbosa-masafjierana, CJienop odium crii- 
soeanum-nesodendron, Dendroseris macrayitha-macrophylla, F agar a mayu— externa, 
Gunner a peltata- Masafuerae, Halorrhagis masatierrana-masafuerana, Peperomia 
inargaritifera-Skottsbergii, Robinsonia eve^iia — Masafuerae, and Sophora ferna7i- 
deziajia- masafuerana. As regards Berberis, Fagara, Halorrhagis and Sophora, 
perhaps also Gumiera, Peperomia and Robinsonia, the two members of a pair are 
closely related, but in the case of Halorrhagis the situation is complicated be- 
cause there is a third species, endemic in Masafuera, H. aspe^^rima; masaftiera7ia 
seems, however, to be a better match for masatierrana than asperri7na. As we 
have seen, there is also a third species of Che7iop odium, endemic to Santa Clara, 
very likely formerly occurring also on Masatierra, but extinct there. Of the 3 
species of Dendroseris inhabiting Masatierra, 7narginaia and litoralis 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. I have mentioned this 
above (|). 194) pointing out that most of these families have numerous genera 
and sj)ecies on the opi)osite 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. JOIIOW 
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, Amaryllidaceae 
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 2^1. 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 19 14 [22^] 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 JOIIOW 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 Santalum, a jeneros representados, si no 
en Chile, a lo menos en alguna parte de la costa occidental de Sud-America" — 
this did not, however, apply to the isolated Compositae, to Lactoris 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 (22^), 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 i non-endemic 
{Halorrhagis). Group II, called Neotropical, contained 6 endemic species; Cuminia 
and Juania 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, 1 1 ; 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 
JOllOW'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 [2jg), of which a 



2l6 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 si)ecies; 47. 

a) in Central or S. Chile, 20. 

b) in the Magellan region, but not of subantarctic character, 3. 

c) in South America, but not in Chile, i. 

d) endemic species of non-endemic genera, 21. 

e) ,, ,, ,, endemic ,, , 2. 

2. Element consisting of wide-spread species, also inhabiting Chile, 5. 

3. 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 s])ecies 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, i 7 . 

c) species of isolated endemic genera with supposed transpacific relations, 18, 

d) sj)ecies of endemic genera with unknown relations, 3. 

This analysis was based on my memoir on the Phanerogams published in 
1922 (22^). 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, OcJiagavia and NotJiomyrcia, 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, P^scallonia 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, P>igeron fruticosus, luteoviridis, Ingae, Innocentium, turricola and 
rupicola. 

b. 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, Danthoniacollina, Koeleria micra- 
thera, C"\-perus 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 21 7 

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. 

b. Not endemic, in several cases extending north along the Andes (11): Oreo- 
bolus obtusangulus, Uncinia brevicauli? 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. 

b. 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-Hawaii; 
all endemic, 

a. The genus endemic (9): Cuminia fernandezia and eriantha, Robinsonia 
gayana, thurifera, evenia, Masafuerae and gracilis, Symphyochaeta macrocephala, 
Rhetinodendron Berterii. 

b. 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. — i 2 (8.2 %). 

Isolated endemics of unknown parentage, the Cicoriaceous genera forming a 
natural group. 



2l8 C. SKOTTSBERG 

Lactoris fernandeziana, Dendroseris macrophylla, macrantha, marginata and 
litoralis, Phoenicoseris pinnata, berteriana and regia, Rea neriifolia, micrantha and 
pruinata, Hesperoseris gigantea. 

The ditierences between this arrangement and the one of 1934 are consider- 
able, but partly at least more apparent than real. Group i 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 Selkirkia has now been placed in III, a group 
correspontling 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 [Thyrsoptcris] is endemic in Juan P'ernandez and found on Masatierra 
and Masafuera. 

Of the 53 s{)ecies 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, ii.i and (^^.'jX, 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- 
cctUage 11.4, 22.9 and 65.7, respectively. The difi"erences 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 \ery 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 Vandenhoschia 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 Ti'ichomanes . . . 
its cellular structure quite unique" (CllRlSTENSEN ^^.3). — Also in S. Chile? (see 
I.e. 2). 

Ingae C. Chr. Belongs to the neotrop. pyxidiferum group. 

exsectum Kze. S. Chile, south to Chiloe. Near the neotrop. T. tener'utn Kze. 
Serpyllopsis v. d. Bosch. A monotypical genus without near relatives (Copeland 

^9' 37). 

caespiiosa (Gaud.) C. Chr. (var. fernandeziana C. Chr. et Skottsb., slightly 
different from the forms described from the mainland). Falkl. and Fueg. to S. Chile. 
Hymenoglossum Presl. Monotypical and without near affinity to any known genus 
(Copeland I.e.). 

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. 

cuneaUwt Kze. S. Chile, Valdiv.-W. Patag.; Marquesas Is., Rapa. Near 
H. polyanthes Sw., a variable pantrop. sp. 

caudiculatum Mart. Peru, S. Braz., S. Chile, Valdiv.-49° s. lat. 

fuciforme Sw. S. Chile to W. Patag. A very distinct sp., referred to Meco- 
dium with considerable doubt (CoPELAND ^<5. 95). 

b. HyfnenophyllMH s. str. About 25, austr.-circump. with outlying stations in 
W. Eur. and Japan; H. peltatum (Poir.) Desv. widely scattered in slightly different 
forms. 

pectinatum Cav. S. Chile, Valdiv.-Fueg. An aberrant species, CoPELAND 6g. 34. 

falklandicmn Bak. W. Patag., P'ueg., Falkl., S.Georgia. Related to /'^//^?///w 
which, \{ falklandicum really belongs to Mecodium, where CoPELAND placed it 
-<5^. 94, also must be brought here; peltatum is known from extratropical regions 
of both hemispheres and also reported from Chile (perhaps = var. Mejiziesii '(^x^^X) 
C. Chr. 61. 4), by Ciiristensen also quoted for Juan Fernandez — he must have 
forgotten that in 62. 1 1 all material from there was referred to falklandicum. 

rugosum C. Chr. et Skottsb. Related to //. tunbridgerise (L.) Sm., W. Ind., 
Venez., Chile, Atl. islands, Mediterr., S. Afr., Austral., Tasm., N. Zeal. 

c. Sphaerocioyiium 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 H. ci- 
liatum Sw. and belonging to a small austral-tricentric group of closely allied spe- 
cies, H. Frankliiiiae Col. (N. Zeal, probably = ferrugineum, CllRlSTENSEN 61. 5), 



C. SKOTTSBERG 



Table III. 

List of species. 
Endemics bold faced, endemic genera in capital letters. 



Adiantum chilense Kaiilf 

Arthropteris altescandens (Colla) J. Sm, 
Asplenium dareoidcs Desv 

— obliquum Forst. var 

— macrosorum Bert, ex Colla . . . 

— stellatum Colla 

Blcchiinm auriculatum Cav 

— chilense (Kaulf.) IVIett 

— cycadifolium (Colla) Sturm .... 

— longicauda C. Chr 

— Schottii (Colla) C. Chr 

— valdiviense C. Chr 

Cystopteris fragilis (L.) Bernh 

Dicksonia berteriana (Colla) Hook. . . 

— externa Skottsb 

Dryopteris inaequalifolia (Colla) C. Chr. 
Elaphoglossum Lindenii (Bory) Moore . 
(ileichenia cf. litoralis (Phil.) C. Chr. . . 

— pedalis (Kaulf.) Spr 

— quadripartita (Poir.) Moore .... 
Histio[)teris incisa (Thunb.) J. Sm. . . 
Hymenoglossum cruentum (Cav.) Presl 
Hymenophyllum caudiculatum Mart. . 

— cuneatum Kze 

falklandicum Bak 

— ferrugineum Colla 

— fuci forme S\v 

— pectinatum Cav 

— plicatum Kaulf 

- rugosum C. Chi. et Skottsb. . . . 

—  secundum Hook, et Cirev 

— tortuosum Hook, ct Crev 

Hypolepis ru.t^osuia (Labill.) J. Sm. . . 
Lophosoria (|uadripinnata ((Imcl.) C. Chr. 

Lycopodium ma.uellanicum S\v 

— • scariosum l-'orst 

Ophioglossum fernandezianum C. Chr. 

Pellaea chilensis I'ee 

Polypodium magellanicum (Desv.) . . 



Mt 



SC 



DERIVATION OF THE FLORA AND FAUNA 



— intermedium Colla 

— lanceolatum L 

— Masafuerae Phil 

— trichomanoides Sw 

Polystichum berterianum (Colla) C. Chr, 

— vestitum (Forst.) Presl 

Pteris berteroana Ag 

— chilensis Desv 

— semiadnata Phil 

Serpyllopsis caespitosa (Gaud.) C. Chr. . 
THYRSOPTERIS elegans Kze . . . 
Trichomanes exsectum Kze 

— Ingae C. Chr 

— philippianum Sturm 



Mt 


sc 


Mf 


+ 


- 


+ 


+ 


- 


+ 


- 


- 


+ 


+ 


- 


- 


+ 


- 


+ 


+ 


- 


+ 


+ 


- 


+ 


+ 


- 


+ 


+ 


- 


+ 


+ 


- 


+ 


+ 


- 


+ 


+ 


- 


+ 


+ 


- 


- 


+ 


- 


- 



aeriigi7wsimi (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., 
N. Zeal., Fiji. 

secundum Hook, et Grev. S. Chile, Valdiv.-Fueg. 

plicatum Kaulf. See 2/i.g.'j6i. 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 
Cyatheoides thyrsopteroides Berry is remarkably similar to Thyrsopteris; it was, 
however, found sterile, and he adds that most of the fossils formerly referred to 
Thyrsopteris are based on too slender evidence. Seward (2.?o. 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 Dicksonia and Cyathea had a common ancestor" ((5^. 48) and he thinks 
that it is allied to Culcita Presl. He brought Thyrsopteris to his large and possibly 
very heterogeneous family Pteridaceae, where also Dicksonia is placed. 

Winkler's statement that Thyrsopteris is found on Masafuera only [28y. 472) 
is erroneous. 
Lophosoria Presl. One polymorphous species. 

quadripinnata (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- 



222 C. SKOTTSBERG 

fercnce between the plants of Masatierra and Masafuera) is certainly unlike the 
form inhabitint^ Chile; see 62. 16. 

l)ickso7iia Lllcrit. 24; i Mex., 3 Centr. Amer. to Colomb., i Ecuad.-Braz., 2 Peru; 
1 St. Helena, i Malays., 4 X. Guin., 2 Austral.-Tasm., 3 N. Zeal., 3 X. Caled., 
I Mji-Sanioa. 

berteroana Colla and externa Skottsb. are very near each other and closely 
related not to the Andean group but to I), laiiaia Col. from X. Zeal. CllRlST 
{■,-9.154) further cjuotes as relatives 1). antarctica Labill. (Austral.), Bluffiei {)^zt) 
Moore (Indones., Philij)p.), gnvidis Ros. (X. Guin.) and species from X. Caledonia 
and the South Sea islands. 

Polypodiaceae. 
Cvstopii-ris Bernh. 15; i boreal, 10 Ind.-China, Japan, X. Zeal., 2 Eur., i 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). 
/>';-)Yy'/r;7> Adans. Sens u C. Chr. about 1 200 sp.; s. str. Coi'KLANl) about 150; "an 
assemblage of genera that must be segregated in their entirety before of use for 
a study of distribution" (Coi'KLAXD 6g). 

inaeqtialifolia (Colla) C. Chr. Belongs to a neotrop. group and comes nearer 
to tropical forms than to the south Andean I), spcctabilis (Kaulf.) C. Chr., which 
extends south to CUiile. 
PolysticJiuui Roth. About 225; widely spread, numerous in E. As. 

berterianum (Colla) C. Chr. Xear P. adiantiforrnc Eorst., spread through the 
S. temp, zone, north to W. Ind. 

I'i'stituni (k'orst.) I'resl coll. S. Amer. (also Chile), Austral., X. Zeal. 
Artliyoptcris ]. Sm. About 20; j)alaeotrop., recorded from trop. Afr., north to Arab.; 
Madag., Australas., north to l^hilipj). Is.; Xew Caled., Fiji, Samoa; best developed 
X. (iuin., X. Caled. and Madag. 

altescandens (Colla) J. .Sm. Closely related to Pacific forms (62.21). 
Asplciiiii))! L. 650-700; cosmopolitan. 

obliquiiDi Eorst. var. cJioidrophyllum (Bert.) Mett. Typical obliqutwi reported 
from S. Chile, Austral, and X. Zeal. 

tnacrosorutn Bert, ex Colla. Distantly related to neotrop. sp.; see 62. 23. 

stellatutn Colla. Belongs to the pantrop. ///;/;^/<r? /;/;;/ group; see ^2. 24. 

day f aides Desv. (magcllanicum Kaulf.). S. Chile, Eueg., Ealkl. Closely related 
to A. alrarczciise Rudm. Ikown of Diego Alvarez (Tristan da C); see 61. 13. 
lUiilniuui L. 180-200; essentially southern. 

auriculatuni Cav. Temj). S. Amer., common in Centr. and S. Chile and related 
to //. anstralc L. (S. Afr.). 

raldinioisc C. Chr. Chile, south to Chiloe. Related to B. lanccolaium (R. Br.) 
.Sturm from Austral, and \. Zeal. As regards the nomenclature, see i6g. 54, and 

Schottii (Colla) C. Chr. Stands near B. aitcnuatiwi (Willd.) C. Chr. (S. Afr., 
IC. Austral., Polyn.) and nieridejise (Kaulf.) C. Chr. (trop. Amer.); see 62. 27. 



DERIVATION OF THE FLORA AND FAUNA 22$ 

chilense (Kaulf.) Mett. Centr. Chile to W. Patag.; Falkl. Belongs to the capense 
group (S. Afr., Indomal.), see 62. 2^] and i6g.^']. 

cycadifolium (Colla) Sturm. Almost too near B. viagellmiicum (Desv.) Mett. 
(S. Chile to Fueg. and Falkl.); both related to B. tabulare (Thunb.) Kuhn (S. and 
E. Afr., Madag., Mascaren.). 

longicauda C. Chr. Very near B. Sprucei C. Chr. from trop. S. Amer. (Ecuad., 
Boliv., Braz.), see 2^0. HiCKEN records it from Tucuman in Argent. (133. 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 Fee. Very close to P. nivea (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. -Poly n. 

rugosida (Labill.) J. Sm. Centr. and S. Chile; the typ. sp. Austral., N. Zeal., 
var. villosoviscida (Thouars) C. Chr. [61. 6) on Tristan da C; Polypodiu7n viscidum 
Roxb. from St. Helena very likely is another variety. 
Adiafitum L. 200-225; widely distributed but most numerous in S. Amer. 

chilense Kaulf. Peru-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.). 

semiadnata Phil. S. Chile, Valdiv.-Huafo I. A distinct sp., possibly related 
to P. pidchra Schlechtd. from Mexico. 

berteroana Ag. Related to P. comans Forst. (Austral., Tasm., N. Zeal.) and 
particularly to P. endlicheriana Ag. (Norfolk I.). 

Hisiiopteris (Ag.) J. Sm. An assemblage of trop. and austr.-circump. forms which 
according to CllRlSTENSEN cannot claim to be regarded as different species 
[62. 36), whereas Copeland [6g. 60) speaks of local, derived species (a concentra- 
tion in Indomal.-Polyn.) differing sufficiently from the following polymorphous taxon. 

incisa (Thunb.) J. Sm. Chile, S. Afr., Tristan da C, Austral, Tasm., N. Zeal., 
Polynes. 

Polypodiwn L. coll. World-wide; Ind. Fil. registers more than iioo sp. The genus 
is now generally broken up, most radically by CoPELAND, who segregates numer- 
ous genera partly restored from synonymy. 

a. Granimitis 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. niagellanician 
(Desv.) C. Chr., but later restored to specific rank by CHRISTENSEN 61. 18. Distri- 
bution of Billardieri: Austral., Tasm., N. Zeal., Auckl. and Campb. Is., Lord 
Howe and Norfolk Is., St. Paul and N. Amsterd. Is., Kerguel. 

b. Synammia Presl, According to Copeland 6g. 184 only one species, wS. 
Feuillei (Bert.) Copel. [Polypodiwn trilobmn Cav.), but I fail to see how we could 
exclude the island species. 



224 



C. SKOTTSBERG 



intermedium Colla [translucens (Kze) Fee]. JOHOW united P. irilobimi Cav. 
and califoyjiicuDi Kaulf. with traiislucens, and they seem to belong to the same 
small group, to which also P. Espiiiosae W'eatherby (Chile, Atacama) must be 
rcferretl. 

c. PolvpodiioJi s. str., about 75 sp., mainly N. hemisph. and neotrop. 
Masafiierae Phil. S. Peru to N. Chile (Antofagasta), Argent.; comp. i^g. 14 

and 16S. 33. \>ry close to P. pycnocarpiiui C. Chr. (Mex.-N. Chile). 

d. Xiplwptcris Kaulf. About 50, pantrop. 

tyichonianoides Sw. Trop. Amer. A puzzling record (see 2^g. 766). 

e. Plcopeliis Humb. et Bonpl. About 40, pantrop.; the following sp. wide- 
spread. 

la}iceolatii>n L. Mex. and \\\ Ind. to subtrop. S. Amer. (also in Chile), St. 
Helena, Tristan da C, Afr., Madag., Indomal., Hawaii. 

lUapJioglossiiui 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. 

Li)idenii (Bory) Moore. Mex. to Ecuad. and Braz. 

Gleicheniaceae. 
GleicJienia Sm. About 130 sp., the majority of them referred to 5//f//<f?7/i- (Kaulf.) 
Ching, to which also the species in J. P'ern. belong; about a dozen sp. spread 
over the austral zone; 5 Chile, one extending to Falkl.; i S. Afr., i Madag., 
Mascaren. and Seych. Is., i Tasm., i Tasm. and N. Zeal., 2 N. Zeal. 

quadripartita (Poir.) Moore. S. Chile to Fueg. 

pedalis (Kaulf.) Spreng. S. Chile, south to Chonos Is. 

cf. litoralis (Phil.) C. Chr. Hardly identical with this little known species and 
perhaps only a form of G. pedalis. 

Ophioglossaceae. 
OpJiioglossuDi L. 28 sp. recognized by Clausen {64); the genus "scattered with 
remarkable uniformity over the habitable globe" (COPELAM) 6g. 12). 

fertiandezianum C. Chr. Not very near 0. ypaiicnse Mart. (Colomb., Braz.) 
as Chris ri-:NSEX thought, but close to 0. scariosuni Clausen from Peru, Dept. Junin. 

Lycopodiaceae. 
Lycopodiuin L. A large world-wide genus, badly ill-treated by Nessel, who listed 
several undistinguishable species in J. P^ern.; see 2/i.g. ^66. 

j/Kii^r/Iimiiinn Sw. .S. Chile to P\ieg., P'alkl., S. Georgia, Marion I., Kerguel. 

scariosiiiH I^'orst. Chile, Valdiv. -Guaytecas Is. [L. gayajiuni Remy), N. Zeal.; 
Tr(){). Andes, Braz. [L. Jiissicui Desv.). See I.e. J^S'] . 

In his treatment of the geographical groups distinguished by him, JOIIOW 
included the ferns — no 1^'ern Allies were at that time known from Juan Fernandez. 
All were classified as American exce[)t 3, Dicksonia berteroafia, said to be similar 
to a species from T'iji, Ptrris bcrtcroaiia (identified with comans), and '' Aspleniimi 
/oJigissiuiNiN Illume", the s[)ecies now known as BlecJinimi longicatida. In 1914 
[22^) I went a step further; to my "Altpazifisches P^lement" 5 species were referred, 
to the Trop. American 5, and all the rest to the Chilean. In 1934 I attempted 
a more detailed subdivision (279, here translated from P>ench). 



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. 

2. Element consisting of wide-spread species, also inhabiting Chile, 2. 

3. Neotropical element, not represented in Chile, 7. 

a) non-endemic species, i. 

b) endemic species of non-endemic genera, 6. 

4. Magellanian-Old Antarctic element, 9. 

a) non-endemic Magellanian species, 2. 

b) ,, ,, Old Antarctic but not Magellanian species, 4. 

c) endemic species allied to subantarctic species, 2. , 

d) ,, ,, not allied to subantarctic species, i. 

5. Pacific element, 5. 

a) isolated endemic species of wide genera, i. 

b) endemic species belonging to genera or sections of west to central Pacific distribu- 
tion, 3. 

d) species of endemic genus with unknown relations, i. 

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. 

d. 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 (75%). 
Not endemic, extending north but not beyond Centr. Chile: Serpyllopsis caes- 
pitosa, Hymenophyllum falklandicum, Asplenium dareoides, Lycopodium magel- 
lanicum. 

III. Neotropical element. — 9 (17 %). 

a. Endemic species (7, 13.1 %): Trichomanes philippianum and Ingae, Dryo- 
pteris inaequalifolia, Asplenium macrosorum and stellatum, Blechnum longicauda, 
Ophioglossum fernandezianum. 

b. Not endemic (2, 3.8 %): Polypodium trichomanoides, Elaphoglossum Lindenii. 

IV. West Pacific element. — 5 (94%). 

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. — i (1.9%). 

Endemic genus witliout 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 w^ould expect the opposite, 
because ferns ought to travel across the ocean much more readily than flowering 
plants. 

III. Musci. 

The following list is based on Brotiierus' paper in vol. II ( ?^) and on his 
treatment of the group in the 2nd edition of Natiirl. 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 (i2g) and in pt. 2 of 
Irmscher's book (i4j). I am indebted to Dr. Herman Persson for kind assistance. 

Mi =- Masatierra, Mf = Masafuera. 

Ditrichaceae. 

l^Uiiridium Brid. (about 30). Wide-spread, mainly temperate. 

Robinsomi (Mont.) Mitt. Chile, Urug. Belongs to a S. Amer. group. — Mt. 
DitricJiuni Timm (about 50). In all parts of the world. 

affine (CM.) Hampe. S. Chile and Patag., E. Austral., N. Zeal, Auckl. Is. 
Not listed in 210, where D. elongatiim (H. f. et W.) Mitt, is quoted for Chile, Austral., 
Tasm. and N. Zeal.— Mt, Mf. 

lougisetum (Hampe) Jaeg. S. Chile to Fueg. — Mf. 
Ceratodoi Brid. (2, one trop.). 

puypurcus (L.) Brid. Cosmopol. — Mt, Mf. 

Pottiaceae. 

llymc}wst()))iu))i R. Br. (50-60). All over the world. 

kuuzcanum (CM.) Broth. S. Chile.— Mt. 
(jyuiuostannufi Hedw. (10). All over the world. 

calcariUDi Jkyol. germ. Eur., Afr., As., N. Amer., S. Amer.: P3cuad.-Chile; 
Austral., Tasm., X. Zeal.— Mt, Mf. 
Tricliostomum Hedw. (81). Cosmopolitan. 

hracliydoiitiuin Bruch. Eur., Caucas., N. Afr., Macaron., Mascar., Japan, N. 
Zeal., but not mentioned in 210. — Mf. 

Dicranaceae. 

AmpJiidium Xees (10). Very widely distributed. 

cyatJiicarpum (Mont.) Broth. Ecuad.-Chile, S. Georgia, Afr., E. Austral., Tasm., 
N. Zeal.— Mt, :\If. 



DERIVATION OF THE FLORA AND FAUNA 227 

Dicranella Schimp. (6o). All continents, numerous S. Amer. and As. 

costata Broth. — Mt, Mf. 
Oncophorus Brid. (8). Southern S. Amer., Eur., E. As., Ceylon. 

fuegiaims Card. Patag.-Fueg. — Mf. 
Dicranolonia Ren. [']6). Almost exclusively S. hemisph,, austr.-subantarct., i S. Afr. 

fernandezianum Broth. Near D. Dusenii Broth. (S. Chile). — Mt, 

capillifoliwn Broth. S. Chile-Fueg. — Mt. 

capillifolioides Broth. Near the former. — Mf. 

Menziesii (Tayl.) Par. S. Chile, E. Austral., Tasm., Norfolk I., N. Zeal., 
Chatham Is., Auckl. Is. — Mt. 

Billardieri (Schwaegr.) Par. Peru-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. 

mtroflexus (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. 

trimcatus CM. Chile.— Mt. 

polytricJwides De Not. W. and S. Eur., N. Afr., Mascar. — Mt. 

aberrans Broth. A very peculiar sp. — Mt, Mf. 

areodiciyon (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. 

Richardi Schwaegr. Centr. and S. Amer. to Chile. — Mt, Mf. 

leptodiis (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 (i). Austral (i), N. Caled. {4), N. Zeal + N. Caled.(i). 
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.crass/pes Wi\s. (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 Peru, S. Chile and Braz., W. Afr., 
Macaron., Tristan da C, Indomal, Queensl, N. Zeal. — Mt. 



228 C. SKOTTSBERG 

LeptodoJitiuDi Mampe (80). Most numerous in Amer., 40% And. 

fernandezianum Ikoth. Related to L. luteum (Tayl.) Mitt. (trop. And.). — Mf. 
Diiiymodon Hedw. (91). Subcosmopol., mainly temp., essentially Amer.: Mex,— 
Centr. Amer. -Peru and Chile; i Antarct. 

calytnperidictyon Broth. — ^Mt. 

linearis l^roth. — -Mf. 
Tottula Hetlw. (220). Subcosmopol., mainly temp. 

scabriutrris (C. M.) Mitt. Chile.— Mt. 

flaij^cllayis (Schimp.) Mont. Chile.— Mt. 
G}i)H)}iia IChrh. (230). Subcosmopol. but rare in the trop. zones. 

phyllorhizans Broth. — Mt. 
Kluicomitriu))i Ikid. (80). As Grimmia. 

subuiirritum (CM.) Par. W. Patag.-P\ieg.— Mf. 

sy)}iphyodo)itu})i (C. M.) Jaeg. S. Chile to P\ieg. and Falkl., S. Afr., Kerguel., 
Tasm., \. Zeal. Not listed for N. Zeal, in 210. — Mt, Mf. 

striatipiliDJi Card. S. Chile to Fueg., S. Georgia; N. Zeal. [^216). — Mf. 

lajiugiiiosuni (Hedw.) Brid. Cosmopol. — Mf. 

loriforme Dus. W. Patag. — Mf. 

conrolutuni Mont. S. Chile to W. Patag.— Mf. 

Ptychomitfiaceae. 

Ptycho))utrin))i (Bruch) P'urnr. (62). Of wide distribution in temp, zones. 
fernandezianum (Mitt.) Jaeg.^Mt, Mf. 

Orthotrichaceae. 

Zyo^odoii Hook. f. et Tayl. (about 100). N. and Centr. Amer. (6), trop. S. Amer. 
(40), .S. Chile and Argent. (20), \\\\x. (4), trop. Afr. (2), trop. As. (6), \l. Austral.- 
X. Zeal. (10). 

i)itcn)ii'dius B. et S. S. Amer., Afr., Monsoon reg., Austral., Tasm., N. Zeal. 
— Mt. [26^). Z. oboi'alis Mitt., a doubtful species (,?/?. 139). is supposed to be iden- 
tical with i?ifrn>ifdii(s. 

Moiziesii (Schwaegr.) W. Arn. S. Chile, Austral., Tasm., N. Zeal. — Mt. 
Sfoioi/iitriinn (Mitt.) l^roth. (2). Viiru, Chile. 

pojtastichuni (Mont.) Ikoth. Peri'i-W. Patag.— Mf. 
riota Mohr (43). X. and S. temj). zones, ^3 Chile, some N. Zeal. 

fernandeziana Malta (.?/?). Xear ('. rufula (Mitt.) Jaeg. (Chile-W. Patag., 
Argent., X. Zeal.).— Mf. 

Macromitriion Brid. (415). Troix-subtro|)., special gr()uj)s Chile-Patag. and P2. Austral.- 
Tasm.-X. Zeal. 

Iiyinoiostouiu))! Mont. S. Chile-P'ueg. — Mt. 

saxatile Mitt. — Mt, Mf. 

fernandezianum Broth. Related to J/, aspcruluni Mitt. (Tasm., N. Zeal.; not in 
210).— Vii. 

Masafuerae l^roth. Xear the former. — Mf. 



DERIVATION OF THE FLORA AND FAUNA 229 

Funariaceae. 

Funaria Schreb. (200). A world-wide genus. 

hygroinetrica (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. 

Lechleri C. M. Chile.— Mf. 

fernandezianum Broth. Related to B. Cruegerz Hampe (nGotrop.). — Mt, Mf. 

Leptostomaceae. 

Leptostomutn R. Br. (i i). S. Chile (2), Indomal., Austral., N. Zeal., Norfolk I., Campb. I. 
Menziesii (Hook.) R. Br. S. Chile to W. Patag. and Fueg.— Mf. 

Eustichiaceae. 

Eustichia (Brid.) Mitt. (8). 6 Mex.-S. Chile, i S. Afr., i islands of E. Afr. 
Poeppigii (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.; i pantrop. 

Nouae Hollandiae Brid. v2iX. patagonicum Card, et Broth. W. Patag.; the typical 
sp. Austral., Tasm., N. Zeal.— Mt. 

mnioides (Hook.) Schimp. Colomb.-Fueg., E. Austral., Tasm., N. Zeal. — Mf. 

Bartramiaceae. 

Anacolia Schimp. (7). W. N. Amer.-trop. Andes (5), i Medit., i Ethiop. 

subsessilis (Tayl.) Broth. Mex.-Ecuad.— Mt, Mf. 
Bartramia Hedw. (iio). 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 210, where B. halleriajta 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., 
Tasm., N. Zeal, Auckl. and Campb. Is.— Mt, Mf. 

vagans (Hook. f. et Wils.) Mitt. Forms a separate section. Chile-Fueg., S. 
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. hcmisph. 

Iliiuiboldtii \\\(^oV) Lindb. Me.x., \V. Ind., Colomb.-And. Patag., Fueg., Falkl., 
Centr. Afr.. Madag., Reunion, Austral., Tasm., N. Zeal. A polymorphous sp. — Mf. 

Cryphaeaceae. 

Deudrocryf^haea Par. et Schimp. (4). Austr.-subantarct., 2 Chile, I Argent.-Patag., 
I Tasm.-\. Zeal. 

cuspidata (Sull.) Broth. S. Chile. — Mf. 
Cvptodon I'ar. et Schimp. (6). 2 Austral.-N. Zeal, 1 X. Caled., 2 Fiji-Samoa-Tonga, 
I J. Fern. 

crassinervis Broth. Close to C. (210 under CrypJiaea) dilatatus (Hook. f. et 
Wils.) Par. (h:. Austral., X. Zeal.).— Mf. 

Lepyrodontaceae. 

Lcpyrodou Hampe (6-8). i bicentric (Chile-Fueg., P3. Austral., Tasm., N. Zeal., 
Canipb. I.), 2 or 3 trop. Andes to Braz. and Argent., 2 or 3 Chile, I N. Zeal. 

pannilus Mitt. Centr. and S. Chile.— Mt, Mf. 

tojnffitosus (Hook.) Mitt. Colomb.-Perii, Braz., Argent., W. Patag. — Mt. 

iuiplexus (Kze) Par. Chile, south to Fueg.- — Mt. 

Ptychomniaceae. 

PlycJioDuuH))! 1 look. f. et Wils. (9, or 8 if cyg)iisetu))i = acicularc). Austral-subant- 
arct., e.xtending north to Braz. and Hawaii; Argent, and Chile to Pueg., \\. Austral., 
Tasm., X. Zeal, and subant. islands. Lord Howe L, X. Caled., Polyn. 

suhaciculare Besch. Chile, south to I^\ieg. — Mt. 

falcatiilum Broth. Related to the former. — Mt. 

ptychocaypian (Schwacgr.) Mitt. Chile, south to \V. Patag.— Mf. 

Neckeraceae. 

WeyDioutliia Ikoth. (3-4). Austr.-circump., Hawaii. 

iHol/is (Hedw.) Broth. Centr. and S. Ciiile to Magell., PI Austral., Tasm., 
X. Zeal.- Mt, Mf. 
Ltplodo}! Mohr (4I Afr. and islands (3), and the following. 

Sniithii (Dicks.) Mohr. Argent., Chile; S. ICur., Caucas., Afr., Macaron.; Y.. 
Austral., X. Zeal.^Mt, Mf. 

Xcckcra Hedw. (127). Widely distributed troi).-subtrop., extending south to Chile 
and X. Zcal.-Auckl. I. 

rotundata Broth. A very distinct sp. — Mf. 
PoyothainjiiuDi IHeisch. (51). i W. X. Amer., 41 Centr. and S. Amer., extending 
south to Patag., 7 trop. Afr., i Ceylon, i X. Zeal. 

fasciculaiion (Sw.) Meisch. W. Ind., Colomb.-Peru, Ikaz. — Mf. 

arbuscnlans (C. M.) Broth. Chile, Patag. — Mf. 



DERIVATION OF THE FLORA AND FAUNA 23 1 

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 rigidum. — Mt. 

Ingae Broth. Very near the former. — Mt. 

crassinervium (Mitt.) Broth. — Mt. 

proboscideum Broth. Near the former. — Mt. 

assitnile Broth. Very near the former. — Mt. 

confertum (Mitt.) Broth.— Mt. 
Piimatella (C. M.) Fleisch. (37). Pantrop. 

tnacrosticta 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. 

assitnile Broth. Near the former. — Mt. 

fernandezianum Broth. Related to D, rotundifoliuin (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.- 
N. Zeal, 2 of these also S. Amer. 

anomalum (Schwaegr.) Mitt. F'ueg. — Mt. Perhaps only an extreme water form 
of the following (jjj. 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 210, where 
P. dentatum (Hook. f. et Wils.) Mitt, is supposed to occur in Chile and P'uegia. 
Eriopus (Brid.) C. M. (25). Some trop. Andes, i S. Afr., numerous N. Zeal. 

leptoloma Broth. Related to E. apiculatus (Hook. f. et Wils.) Mitt. (S. Chile- 
Fueg., Austral, Tasm., N. Zeal).— Mt, Mf. 

grandiretis Broth. Near the former. — Mf. 
Lamprophyllum 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). 

concinnum (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 

Thouvii (Schwaegr.) Mont. S. Chile-Fueg. Belongs to a group of 4sp. (3 Chile, 
I N. Zeal.).— Mf. 

Rhacopilaceae. 

Rhacopilum I'alis. (51). Pantrop., mainly southern, south to X. Zeal. 

fcDiaudeziiDiuin Card. S. Chile. Related to R. toinentosum (Sw.) Brid. (trop.- 
subtrop. Amer.). — Mt, Mf. 

Thuidiaceae. 

riiuidiion Bryol. eur. (161). All over the world, especially in humid mountain climates. 

Masafuerae l^roth. Related to 7\ fuluastrum (Mitt.) Jaeg. (Tristan da C, 
X. Zeal, but not included in 210) and to the following sp. — Mf. 

I'aldiviae Broth. S. Chile.— Mt, Mf. 

Amblystegiaceae. 

Sciayouiiuni Mitt. (22). Mainly austral, the majority southern S. Amer., i Cape, 2 
Ciiina, 2 1^. Austral., i X. Zeal.; the single X. Amer. sp. forms a separate section. 
Sect. Aloma 6 (i Boliv., 2 Fueg., 2 \l. Austral., i X. Zeal.). 
pacJiyluDia (Mont.) Par. S. Chile-W. Patag.— Mt, Mf. 

Hypnaceae. 

Stcrcodoji Mitt. (7). 2 Mex., i Chile, 4 mountains of Asia. 
LccJilcri (C. M.) Mitt. Centr. Chile to W. Patag.— Mt. 
Isoptervi^iuui Mitt. (170). Widely distributed, preponderately trop.-subtrop. 
fernandezianutn Broth. Related to /. teuerum (Sw.) Mitt, (neotrop.). — Mt. 

Sematophyllaceae. 

RhaphidostegiiiDi (Bryol. eur.) De Xot. (106). Subcosmopol., trop. and temp. The 
C'hilcan species sometimes regarded as belonging to a separate genus RJiapJiido- 
rJiyiichiion. 

Masafuerae Ikoth. Similar to R. cyparissioides (Hornsch.) l^esch. (Braz.). — Mf. 

aberrans Broth. Similar to R. callidion (Mont.) Jaeg. (S. Chile to W. Patag.). — 
Mt, Mf. 

cacspifosuiii (.Sw.) Jaeg. \V. Ind., trop.-subtrop. S. Amer. — Mt, Mf. 

caespitosoides Broth. Related to the former. — Mf. 

brachycladulum J^roth. — Mt. 
Rij^odiion Kzc. (19). Trop.-subtrop. 7\mer., 2 Afr. 

toxcniuni (Schwaegr.) Schim[). Trop. S. Amer. to W. Patag. — Mt, Mf. 

arboresicus (C. M.) Broth. S. Chile-\V. Patag.— Mt, Mf. 

hylocoDiioidcs Card, et Broth. Patag. — Mf. 

robustutn l^roth. Mt. 

Looseri Thcr. Related to A', gracile Ren. et Card. (Costa Rica). — Mt. 

tcDnarix C. M. (elegantulum Card.). S. Chile-W. Patag. — Mt. 
R/iyuc/iostf^i^iian Ikyol. eur. (130). Temp.-subtrop., almost cosmopol. 

coiuplcvium (Mitt.) Jaeg. Centr. Chile. — Mt, Mf. 

toiuifoliuiu (Hedw.) Jaeg. Urug., S. Chile, E. Austral., Tasm., X. Zeal.— Mt. 



DERIVATION OF THE FLORA AND FAUNA 233 

Catagoniopsis Broth. Monotypical, related to Catagonium (S. hemisph.) 
berteroava (Mont.) Broth. Centr. Chile. — Mt. 

Hypnodendraceae. 

Hypnodendron Lindb. (28). Chile (2), IndomaL, Austral., Polyn., Hawaii. 
microstictum Mitt. Chile. — Mf. 

Polytrichaceae. 

Oligotrichmn Lam. et DC. (13). Widely scattered: 2 W. N. Amer., 4 Chile; single 
sp. Braz., trop. Andes, Arct.-alp., Eur., bor.-circump. 

caiialiculatum (Hook.) Mitt. var. The typical sp. in S. Chile. — Mf. 
Psilopilum Brid. (17). A bipolar genus, tricentric in the S. hemisph. 

antarcticu7n C. M. Boliv., Fueg., Falkl., S. Georgia, Kerguel. — Mf. 
Polytrichadelphus (C. M.) Mitt. (22). Centering in the Andes (17), i W. N. Amer., 
I Braz., 2 Magell., i 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 onMasafuera, 
and vice versa. 

The 131 species of mosses hitherto recorded from Juan Fernandez belong to 65 
genera (2: i), 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 Thamnium 
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 tlie 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 1 1 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. Xo 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%). 

ci. luidemic species (15): Dicranoloma fernandezianum and capillifolioides, Fissi- 
dens pycnotylus, Didymodon calymperidictyon and linearis, Ulota fernandeziana, 
Philonotis glabrata, Ptychomnium falcatulum, Distichophyllum fernandezianum, 
Pterygo[)hyllum tenuinerve, Eriopus leptoloma and grandiretis, Thuidium Masa- 
fuerae, Rhaphidostegium aberrans, Rigodium robustum. 

b. Also known from Chile, rarely extending to the extreme south {57): 
Pleuridium Robinsonii, Ditrichum affine andlongisetum, Hymenostomumkunzeanum, 
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 con volutum, Zygodon 
intermedins and Menziesii, Stenomitrium pentastichum. Macromitrium hymeno- 
stomum, Ikyum Lechleri, Leptostomum Menziesii, EustichiaPoeppigii,Rhizogonium 
nmioides, Hartramia aristata, Philonotis krauseana and vagans, Rhacocarpus Hum- 
boldtii, Dcndrocryphaea cuspidata, Lepyrodon parvulus and implexus, Ptychomnium 
subacicularc and j)tyciu)carpum, Weymouthia mollis, Leptodon Smithii, Porotham- 
nium arbusculans, Lamprophyllum splendidissimum, Lopidium concinnum, Hypo- 
|)tcrygium Thouini, Khacopilum 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, Gyinnostomum calcareum, Cam- 
pylof)us introflexus, Rhacomitrium lanuginosum, l^\inaria 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 P'alkland 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 ( r i . 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. 

b. 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,Distichophyllumsubelimbatum 
and assimile, Thamnium rigidum, latinerve, Caroli, Ingae, crassinervium, probosci- 
deum, assimile and confertum. 

V. Atlantic element. — 3 (2.5 %). 

a. Endemic: Fissidens crassicuspes. 

b. 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 I ^ may have had the same history though, 
at present, they do not reach so far south. 

The Atlantic element is artificial. Fissidens crassicuspes is, if Brotiierus 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 polytrichoides \'s> diXv 
Atlantic species with its nearest locality on Madeira; its presence on Masatierra 
is indeed surprising. Trichostomum 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 Reunion, 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, Dicrmiella costata (many in S. Amer.), 
PtvchomitriuDi fevnaudeziamini (other sp. in the Andes), Macroniityiiim saxaiile (many 
in S. Amer.), Bartrauiia fernandcziima and Breutelia Masafuerae (many austral sp.) 
may turn out to belong to an American element. The occurrence in Masatierra 
of 8 endemic species of Thatiniium 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. 

Xo handbook equal to Brotherl's' comparatively modern account of the mos- 
ses exists of the Hepaticae. ScillFKNERs treatment of this group in the 1st edition of 
Xatiirl. Pflanzenfamilien is too antiquated to be of much use. Much important infor- 
mation is, however, found in Hkrzog's work [i2g) as well as in Domin's paper (7^), 
and also in this case Dr. Pkrsson kindly helped me, but to search the voluminous 
s[)ecial literature of the last thirty years was not to be thought of. The number of 
species given is, in many cases at least, too low, but I don't think this matters 
very much. 

The following list is based on PlVAXs' and Herzog's papers (gj, ijo) with 
a few alterations (/-?/). Several species credited to Juan Fernandez by Stephani 
( i^j-,^) but not mentioned by Herzog are included here; some of them are, perhaps^ 
identical with other species. 

Marchantiales. 

Plagiochasuia Lehm. et Lindenb. (about 20). Mostly trop.-subtrop. 

ntpcstrc (Forst.) St. Cosmopol., also Chile. — Mt, Mf. 
Rehoulia Raddi (i). 

Iieinispliaerica (L.) Raddi. Cosmopol., also Chile. — Mt, Mf. 
LiDiularia (Mich.) Adans. (i). 

cruciata (L.) Dumort. S. Amer., also Chile, Medit. Fur., Atl. islands, Afr.^ 
Austral.— Mt, Mf. 

Maycliautia L. (about 50). Subcosmopol., numerous trop. 

polyifwypha L. Cosmopol., also Chile.- — Mt, Mf. 

hertcrpiDia Lehni. et Lindenb. Chile south to I^'ueg., Falkl., St. Helena, S. Afr.,, 
Marion I., Kerguel., Austral., Tasm., N. Zeal. — Mt, Mf. 

foliacca Mitt. S. Chile, Tasm., N. Zeal.— Mt, Mf. 

Metzgeriales. 

Riccaydiii )^.V . (iray (140-150, but much higiier figures are given). N. temp. 6^ 
trop. Amer. 43, Afr. 14, trop. As. -Oceania 53, austr.-subantarct. 35. 

fucgiensis Massal. S. Chile to Fueg. — Mf. 

brcniramosa (St.) I^vans. Falkl. — Mt, Mf. 

adglutinata Fvans. — Mt, Mf. 

jusularis SchitTn. St. Paul and New Amsterd. Is. — Mt. 

pariabilis Fvans. 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 M. decipiens as a variety by HoDGSON [136. 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. 

Hochstetteri Nees et Mont. Falkl— Mt, Mf. 

hymenophylluni (Hook.) Nees et Mont. Trop. Amer..^ Chile, Austral., Tasm., 
N. Zeal. — Mt, Mf. Probably identical with vS. podophylla (Thunb.) Mont, et Nees 
from Centr. and S. Africa (S. Arnell, ms.). 

Pallavicinia S. F. Gray (about 25). Trop.-subtrop., Colomb., Indomal.; Kerguel., 
N. Zeal. 

xiphoides (Tayl.) St. N. Zeal.— Mf. 
Monocloa Hook. (2, i trop. Amer.). 

Forsteri Hook. S. Chile-W. Patag., N. Zeal.— Mt, Mf. 
Androcryphaea Nees (i ; 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 L. (about 140). Mainly trop. 

Skottsbergii (St. ex p.) Evans.— Mt, Mf. 
Megaceros Campb. (about 40). Mex., W. Ind., S. Amer. south to Chile, Reunion, 
Indomal.-N. Zeal., Oceania. 

ftiegiensis St. W. Patag.-Fueg. Related to neotrop. sp. — Mt, Mf. 

Jungermanniales . 

Solenostoma Mitt. (30). Subcosmop., but now partly referred to Aplozia Dum., partly 
to Plectocolea. 

crassuhim (Mont.) St. S. Chile.— Mf. 

obtusiflorum St. — Mt. 

rostratum St.— Mf. 
Jmnesoniella Spruce (20). Mainly trop.-subtrop. (i Engl.) but also important in the 
austral zone. 



238 C. SKOTTSBERG 

coloyata (Lchni.) Spruce. Centr. Amer., S. Amer. south to Fueg., Falkl.; Tristan 
da C, S. Afr., Marion I.. Kerguel.. \\. Austral., Tasm., N. Zeal, Auckl., Campb. and 
Antijiodes Is. — Mt, Mf. 

)nalui}ia St. Fall<l. — Mt. 

oeuops St. S. Chile, south to Fue^^., S. Georgia. — Mt. 

gyamiiflora (Lehni. et (k)ttsche) Spruce. Colomb.-Chile, south to Fucg.; Tristan 
da C, Marion I., Tasni.^ — Mt. 
Auastrop/iyliinn Xees (31). Very widely distributed. 

leucocephaliini (Tayl.) Spruce. X^enez., Peru, Fueg. — Mf. 
AcrobolbHS Xees (11). Essentially austral: 2 S. Amer., i subantarct., i Himal., 7 
Austral., X. Zeal., X. Caled. 

cxcisus (Mitt.) Schiffn. W. Patag., Fueg., Kerguel. — Mf. 
A)iasirepta (Lindb.) Schiftn. (4). 2 southern S. Amer., i Alaska-P^ur.-E. As.-Hawaii, 
I Himal. 

bifida St. S. Chile: Valdivia, Magell.— Mt, Mf. 
Plai^iochila Dumort. (about 1000). Very few bor. zone; trop., particularly neotrop., 
but well represented southern S. Amer. 

i:;aya}ia Gottsche. S. Chile. — Mt. 

fasiiata St. Chiloe.— Mt, Mf. 

hyadesiaua Bcsch. et Massal. S. Chile, P\ieg.— Mt, Mf. 

deform if olia St. W. Patag.— Mf. 

chilocusu St. S. Chile to W. Patag.— Mf. 

rcctani!;ulata St. W. Patag. — Mf. 

rciiiotidfiis St. S. Chile, Magell.— Mf. 

fuscobriinnea St. Almost too closely related to P. riibescens Lehm. et Lindenb 
(Syn. l\ chilcusis St. according to Hkkzog 770).— Mt, Mf. 

pudftcusis St. Chiloe. — ^It. 

hoDiomalla St. W. Patag. — Mf. 

mcsiivia Lindenb. S. Chile south to Magell. — Mt. 

riparia St. W. Patag.— Mt, Mf. 

squarrosa St. W. Patag.— Mt, Mf. 

robusta St. W. and \\. Patag., Fueg. — Mf. 

i'/ahj Hook. f. et Tayl. Inieg.— Mf. 

Xo/arisii Lehm. Falkl. — Mt, Mf. 
'ryliouDitJius Mitt. (25). .S. hemisph., best represented Australia; i \V. Ind. 

liwbalus St. W. I'atag.-- -Mt. 

silvaticus St. MW 

densiretis Her/.. Mf. 

bilobatus St. — Mt. 
My/in S. V . (ira>' (40-50). Tiie majority neotrop. mountains (Mex., W. Ind., Centr. 
and S. Amer. south to Magell.), 4 i\fr., i .S. Georgia. 

rrpe?is (Mitt.) I lerz. Magell., X. Zeal.— Mf. 

t'uscovircus (Tayl.) St. See / ,'0.714. Centr. and S. Chile to Fueg. — Mf. 

li^s^ulata (St.) Ilerz. \V. Patag.— Mt. 



DERIVATION OF THE FLORA AND FAUNA 239 

Lophocolea Dumort. (250-300). World-wide; main concentrations subantarct. S. 
Amer. (60), S. As.-Oceania (35), and Austral.-N. Zeal. (70), 

rotu7idifolia St. W. Patag., Fueg., Falkl. — Mf. 

fertiandeziaria 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. 

textilis Tayl. W. Patag.-Fueg., Falkl.— Mf. 

diver genticiliata St. S. Chile, W. Patag., Fueg.— Mt, Mf. 

mtiricata Nees. W. Ind., S. Braz., Centr. and S. Chile to W. Patag., S. Afr., 
Reunion, Java, N. Guin., E. Austral., N. Zeal.— Mf. 

angulata St. — Mt. 

cJiilensis St. Chile. — Mf. 

submuricata Herz. Near muricata, but also very close to L. fragrans Tayl. 
(atl.-mediterr., 130. 720). — Mt, Mf. 

Ckiloscyphus 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 i, Eur.-Macaron. i, Indomal.-Polyn. 2, 
Tasm.-N. Zeal. 3; Hawaii.? 

sqiiarristipula Herz. S. Chile to W. Patag. — Mt. 
Marsupidium Mitt, (about 40.?). Austr.-circump., subantarct. Amer., Austral.-N. 
Zeal. 

piljferiim St. S. Chile, E. Austral, N. Caled.— Mt, Mf. 
Adelanthiis 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. Peru, 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. (7^7.51). — Mt. 

fernandeziensis St. S. Chile (/jo. 725). Near L. plumulosa and Lindenb ergii 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. 

Jacquemontii St. Centr. Chile-Magell, Fueg. — Mt, Mf. 



240 C. SKOTTSBERG 

Uerberta S. F. Gray (15). Widely scattered. 

riiuciuata (Tayl.) Herz. S. Chile to W. Patag.— Mf. 
LepicoUil Dumort. (5). Scattered S. hemisph. 

ochyoleucii (Spreng.) Spruce. S. Braz., S. Chile to Fueg., Cape. — Mt, Mf. 
Lepidolaeua Dumort. (12). S. temp, and cold zones. 

mai:;ella)uca (Lam.) Schiffn. S. Chile to Fueg., E. Austral., Tasm., N. Zeal. 
— Mf. 

Trichocolfa Dumort. (32). i l^or. zone, 19 trop. Amer., 7 trop. As.-Oceania, 5 
Austral. -N. Zeal. 

opposita St. Near 7\ australis St. (X. Zeal.). — Mt. 

rettiallata St. S. Chile to W. and E. Patag.— Mt, Mf. 
Scliistochila Dumort. (83). P^ssentially southern: austral-subantarct. (20), Afr. (7), 
Indomal.-Oceania (35), Austral. -N. Zeal. (21). 

berteroana (Hook.) St. S. Chile.— Mt, Mf. 

Skottsbergii St. Related to 5. stratosa (Mont.) St. (S. Chile to Fueg.). — 
Mt, Mf. 

pachyla (Tayl.) St. W. Patag.-MagelL— Mf. 

splacliuophylla (Tayl.) St. W. Patag.-Fueg., N. Zeal. — Mf. 
Balantiopsis (Xees) St. (17). i Braz., i Oueensl., the remainder austr.-subantarct. 

caucellata (Xees) St. S. Chile to W. Patag.— Mf. 

cJiilensis St. S. Chile to W. Patag. — Mf. 

purpiirata Mitt. S. Chile. Determination uncertain. — Mf. 

hians Ilerz. — Mt. 

lancifolia St. — Mt. 
Radula Dumort. (220). Mainly trop.-subtrop. Bor. zone (Eur., N. Amer., 7), trop.- 
subtroj). Amer. (66), Afr. (37), Indomal.-Oceania (69), Austral.-X. Zeal. {29), sub- 
antarct. (15). 

hastaia St. S. Chile to luieg.- Mf. 

niicyoloha St. S. Chile. — Mt, Mf. 

Duscnii St. S. Chile to \V. Patag.— Mf. 

Mittvnii St. I^'alkl.— Mt, Mf. 

Madothcca Dumort. (153). Trop.-subtrop. Amer. (45), trop. As.-Oceania [6"]), 
Austral. -\. Zeal. etc. (12). 

chiloLsis Lchm, et Lindenb. var. feriiandeziensis Herz. Centr. Chile (Co- 
(juimbo; main sp. S. Chile to W. I'atag.). — Mt, Mf. 

subsquarrosa Xees et Mont. S. Chile to Fueg. — Mt. 
I'fulliuna kaddi (;oo-6oo). World-wide, but mainly trop. and southern: X. Amer. 
(35)- 

lickloui Spr. (crassa Herz.). Ccntr. Chile to W. Patag.; W. and S. Afr.— Mt. 

cluloisis St. S. Chile to W. Patag. — Mf. 

lobulaia Hook. f. et Wils. W. Patag., P\ieg. — Mf. 

})uio;cllanica (Sprcng.) Web. et Xees. Centr. and S. Chile to Fueg., Tristan 
da C. Tasm., Campb. I. — Mt, Mf. 

stipatiloba St. Centr. Chile to W. Patag.— Mt. 



DERIVATION OF THE FLORA AND FAUNA 24I 

Lopholejeunea Spruce (74). Trop. and austral; trop. Amer. (13), trop. Afr. (17), 
trop. As.-Oceania (38), Austral.-N. Zeal. (6). 
spinosa St. — Mt, Mf. 

Brachiolejeioiea Spruce (65). Trop. Amer. (26), Chile (i), Afr. (10), trop. As.-Oce- 
ania (22), Austral.-N. Zeal. (6). 

spruceaita (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. (i). 

nudicalymia Herz. Centr. Chile (Coquimbo, iji. 65). — Mt, Mf. 
Lejeiinea Lib. (190). Pantrop., scarce toward the south. 

reticulata Herz. Related to Chilean sp. — Mt, Mf. 
Apha7iolejeunea 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. (i), Afr. 
(2), Indomal. (16). 

bulbosa Herz. W. Patag. (j?7. 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 thallose 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 6^] 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 9 only on Masatierra, 5 
only on Masafuera and 6 found on both islands. 

Of the 59 species recorded for Masatierra i 5 are endemic in the islands (25.4 %); 
the corresponding figures for Masafuera are 67, 1 1 and 16.4 %. After the discovery 
of Siphonolejeunea on the mainland of Chile there is no endemic genus in Juan 
Fernandez. 

Of the total number of species, 124, 80 occur on Masatierra and 90 on Masa- 

i6 - 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 ']6. 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. luulemic (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- 
nuiricata, Lepidozia fragillima and disticha, Schistochila Skottsbergii, Balantiopsis 
hians and lancifolia, Strepsilejeunea squarrosula and macroloba, Lejeunea reticulata. 

b. 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, chilocnsis, rectangulata, remotidens, pudetensis, homomalla, neesiana, 
riparia, S([uarrosa and robusta, Tylunanthus limbatus, Mylia repens, fuscovirens 
and ligulata, Lophocolea fernandeziana, attenuata, divergenticiliata, chilensis and 
muricata,Chiloscyphusintegrifolius,Saccogynasquarristipula,Marsupidiumpiliferum, 
Bazzaniacerinaand peruviana, Lepidozia bicuspidata, pseudozoopsis, fernandeziensis, 
plumulosa and Jaccjuemontii, Herberta runcinata, Lepicolea ochroleuca, Lepidolaena 
magellanica,Trichocolea verticillata, Schistochila berteroana, Balantiopsis cancellata, 
chilensis and purj)urata, Radula hastata. microloba and Dusenii, Madotheca chi- 
lensis and subsfjuarrosa, Frullania hxklonii, chilensis, magellanica and stipatiloba, 
Brachiolejeunea spruceana, I larpalejeunea oxyota, Strepsilejeunea acuminata, Sipho- 
nolejeunea nudicalycina, Aphanolcjeunea asperrima, Colura bulbosa. 

c. Cosiuopolitan (4): Plagiochasma rupestre, Reboulia hemisphaerica, Lunu- 
laria cruciata, Marchantia polymorpha. 

II. Subantarctic-Magellanian element. — 18 (14.5 %). 

All non-endemic: Riccardia breviramosa, Symphyogyna Hochstetteri, Mega- 
ceros fuegiensis, Jamesoniella maluina and oenops, Acrobolbus excisus, Plagiochila 
data and Xotarisii, Lophocolea rotundifolia, pallidevirens and textilis, Chiloscyphus 
lobatus, Adelanthus sphaleriis, Schistochila pachyla and splachnophylla, Radula 
Mittenii, Frullania lobulata, I larpalejeunea setifera. 

III. Neotropical element. — 5 (4.0 %). 
a. Fndemic (4): Fossombronia fernandeziensis, Anthoceros Skottsbergii, Lo- 
pholejeunea spinosa, Aphanolcjeunea dia[)hana. 
h. 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. 

b. Not endemic: Pallavicinia xiphoides. 

V. Known from St. Paul and New Amsterdam Is. — i (0.8 %), 
Riccardia insularis. 

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 P 

Angiosperms 46.9 10.2 12.9 17.7 

Pteridophytes 64.1 7.5 17.0 9.4 

Mosses 64.2 10.8 1 1.7 10.8 

Hepatics 78.3 14.5 4.0 2.4 

The lower percentage of Mosses in A, as compared with the Hepatics, may 
be due to imperfect knowledge of the distributioh and to the fact that 1 1 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 [2(^6, 2^^), 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 Sticta 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. 
11. 886). Later he went over the proof-sheets of this paper and added further 
corrections. I 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 (7, J2, /j, jj^, iy6, lyy, 182, 184, 2g^, 
J2y-j2p) 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. 

I'errucaria Schrad. (about 2/0). All parts of the world. 

microspora Xyl. X. Amer., Chile, l^^ur., Jap. — Mt. 
Microglaeua Koerb. (37). luir., except i Hraz., i Socotra, and the following. 

fernandeziana Zbr.^Mt, SC. 

Dermatocarpaceae. 

X01 )>ia}tdi)ta X\'l. (i). 

piilchella Xyl. X. and S. Amer., south to Fueg., Eur., Afr., St. Paul's I., As., 
Hawaii, X. Zeal.— Mt. 

Pyrenulaceae. 

Arthopyroiia Mass. (150). Tro[).-subtrop. 

Ci>icho}iac M. Arg. Widely distributed in the trop. zone; also Hawaii. — Mt. 

adnexa M. Arg. var. leptosperma Zbr. The typ. sp. Braz. — Mf. 

pliDiorbis M. Arg. Troj).-subtrop., also Hawaii. — Mt. 
Poriua M. Arg. (235). All over the globe, south to X. Caled. and N. Zeal. 

fernandeziana Zbr. Belongs in the vicinity of P. cJdorotica (Ach.) M. Arg., 
a cosniop. sp. — Mt. 

rufocarpella 7An. — Mt. 

depressula Zbr. Possibly related to P. exscrta M. Arg. (Braz.).— Mt, Mf. 
Pyrcnula Ach. (170). Widely distributed in the trop. zone. 

aspisiea Ach. Trop., also Hawaii. — Mt. 

juanunillaua Trev. Trop.; Hawaii, S. Chile. — Mt. 

Kioithii h'ee. Trop. — Mt, Mf. 

Astrotheliaceae. 

Pyroiastnoii ICschw. (18). Trop. 
chiloise Mont. Chile. — Mt. 

Sphaerophoraceae. 

SpJiacrophonis Pers. (S). Of wide distribution, centering S. hemisph. 

)Hela)iociirpus (Sw.) DC. Almost cosmop.; Chile, south to I'^ueg., P^alkl.; N. Zeal., 
Auckl. and Campb. Is. — Mt, Mf. 

Arthoniaceae. 

Artliouia Ach. (about 370). The majority trop.-subtrop. 

Cytisi Massal. var. meridionalis 7An. The typ. s[). luir. — Mt. 

subnebulosa Zbr. Related to A. rphelodes Xyl. (X. Caled.) and scitula Krmph. 
(I^raz.)— Mt. 

berberina Zbr. Related to .-/. rar/a (Ach.) Xyl. (trop. Amer.). — Mt. 

coftipliDiaia l*'ee. I'antroj)., also Hawaii; Chile. —Mt. 

Graphidaceae. 

Grapliis Adans. (about 280). The majority tr()|).-subtrop. 
intricata Fee. Widely distributed trop. zone. — Mt. 
Dumastii 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. 

Dirma Fr. (12). Of wide distribution. 
limitata Nyl. Chile. — Mt. 

Lecanactidaceae. 

Schismatomma Fw. et Koerb. (80). Mainly warmer countries. 
accede7is (Nyl.) Zbr. Chile.— Mt. 

Chfysotrichaceae. 

Byssocaiilon Mont. (5). Austral.-Oceania. 

nivetirn 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. 

sc7'uposus (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. 

jenejisis (Batsch) Zbr. N. Amer., Eur.; N. Zeal. — Mt. 
Pachyphiale Loennr. (4). 2 S.W. Eur., i 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. 

velutmum Zbr. S. Chile (comm. by R. Santesson). — Mt. 
Racodium Fr. (i). 

rupestre Pers. N. and S. Amer. (Staten I., comm. by R. Santesson); Eur. — Mt. 



246 C. SKOTTSBERG 

Collemaceae. 

Lemniopsis (\'ain.) Zbr. (4). i temp. X. Amer. 4- Eur., 2 S.W. Eur., and the 
following. 

polychidioides Zbr. — Mt. 
/V/ys///(i Massal. (10). W. Ind., trop. As., Japan, Austral., N. Caled. 

chilinse Hue. Chile. — Mt. 
Lepf(\i^iii»i (Ach.) S. V. (iray (over 100). All over the world. 

niolucciviuui (Pers.) \'ain. Widely scattered; Hawaii, also Chile. — Mt, Mf. 

tronelloides (L. fil.) S. F. (iray. Trop. -tern])., in Chile south to W. Patag., 
S. (icorgia. — Mt. 

cvivifsiffis (Ach.) Koerb. Tenijx-subtrop., e.g. Hawaii, S. Amer. — Mt, Mf. 

p/iyl/ocarpu/n (Pers.) Mont. Trop.-subtrop., S. Amer., Chile, S. Afr., Philipp., 
Australia. Mt. 

Moiziesii (Sm.) Mont. Mount, of trop. Amer., Hawaii, Chile south to Fueg., 
Ealkl., S. (Georgia.— Mf. 

callitluDnniou (Tayl.) Xyl. Trop. Amer. — Mt. 

Pannariaceae. 

Pcinfiiliflla M. Arg. (40). Widely scattered, warm and cold climates. 

}iii:^roci}icta (Mont.) M. Arg. Chile, south to Fueg., W. Afr. islands, Hawaii, 
Au.stral., X. Zeal.— Mt, Mf. 

symptychia (Tuck.) Zbr. — ^\t. 

pycuophora (Xyl.) R. Sant. var. subdivisa (Zbr.) R. Sant. W. Patag. [32^]. The 
typ. sp. X. Zeal. — Mt. 
I\vi}ia]ia Del. (about 60). Widely spread over the w^orld. 

fia\i^i(usis Zbr. P\ieg. — Mt. 

hilaris Zbr. Mt. 

nihiiriuosa Del. .Scattered; e.g. Hawaii, Chile, Falkl., St. Helena, Campb. I. — Mt. 

ruhijriuosa var. vulcanica Zbr. l^erhaps specifically distinct. ^ — Mt. 
Afiiss(i/o}ij^ia Koerb. (2, i Iv As.). 

catuosii (Dicks.) Koerb. Mount, of N. Amer. and Eur., P'alkl, X. Zeal.— Mf. 
I\s(>ro)fia X\-l. (al)out 60). Mainlx' cold and temp., centering in N. Zeal. etc. 

vulcanicum Zbr. — Mf. 

cephalodinutn Zbr. Mt. 

p/io/ido/iofi (Mont.) M. Arg. Chile, south to Fueg. — Mt. 

spliiuctriuum (Mont.) Xyl. S. Amer., also Chile, south to P'ueg.; X. Zeal. — 
Mt, Mf. 

dasycladuni Zbr. — Mt. 

atigustisecttim Zbr. — Mt. 

Stictaceae. 

Lobaria Schreb. (70). Mainly warmer countries. 

croiulata (Del.) Trev. Trop. Amer., south to Chile, Hawaii, Austral-Oceania. 
— Mt. 



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. 

fragillima (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. 

hirsiita (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. 

mougeotiaita (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. 

Durvillei (Del.) Vain. S. Chile to Fueg., Falkl., N. Zeal.— Mt. 

Jiavicans (Hook. f. et Tayl.) Vain. W. Patag.-Fueg.; Philipp., Hawaii, Australia. 
— Mt. 

Freycmetii {De\) 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. 

Weigelii (Ach.) Vain. Widespread, also Hawaii, S. Chile to Fueg.; Austral, 
N. Zeal— Mt. 

liyieariloba (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. 

plumbeu7n 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, Mt. 

Uucoplaca M. Ar^^ C'hile. — Mt. 

leucozonata 7hx. — Mt. 

euttrolema X>1. Temp. Kur., etc. St. Helena, etc. — Mt, Mf. 

laiYpea Ach. Temp, widely distributed. — Mt. 

viridiins Lamy. luir. — Mt. 

mutabilis Fee. \. and S. Amer., also Chile, W. Eur. — Mt. 

iiterica Tayl. S. Amer., also Chile. — Mf. 
Catillaria (Ach.) Th. Fr. (about 150). Widespread N. and S. hemisph. 

i)iter)nixta Arn. Widespread, south to N. Zeal. — Mt. 

mclasttiria (Nyl.) Zbr. S. Amer., Chile south to F^'ueg., N. Zeal.— Mt, Mf. 

eudochronia (Fee) Zbr. N. and S. Amer.^Mt. 

Umochlora (Mont.) Zbr. Chile.— Mt, Mf. 

theobromina Zbr. — Mt. 
Mi\i^ii/os/>()rii Mey. et Flot. (about 50). Warmer regions. 

rersicolor (I'"ee) Zbr. var. tnicrocarpa Zbr. The typ. sp. S. Amer., N. Zeal. — Mt. 
liiuidia (De Not.) Zbr. (at least 200). World-wide. 

ouiolcuca Kickx. Almost cosmop., also S. Amer., N, Zeal. — Mt, Mf. 

arciutifia (Ach.) Arn. var. hyposcotina Zbr. The typ. sp. F^ur. — Mt. 

df/iips(2?is Zbr. Hawaii. — Mt. 

subluifola (Xyl.) A. Zbr. l^raz.— Mf. 
T(>)U)iia Til. Vx. (about 80). Mainly temp. 

hullata (Mey. et Flot.) Zbr. Fxuador: Chimborazo. — Mf. 
I.of'iuiiutn Koerb. (about 50). Almost cosmop. 

U'UioxautIiu))i (.Spreng.) Zbr. var. albidius Zbr. The typ. sp. subtrop., south 
to Austral, and X. Zeal.; Hawaii. — Mt. 

s/>., different from the former, described as JMyxodictyon lopadioides by ZahL- 
r.RrCKNF.K p. 3<S3. Chile.— Mt. 
l\hizocaypo)i DC', (about 90). Cosmopol., cold to temp, climates. 

i::c()ij;)apliicu))i (L.) DC. Cold and temp., both hemisph.; Patag., F\ieg., Falkl., 
Kerguel., W. Antarct. Mf. 

microspermum 7An. .Similar to the following sp.— Mf. 

dbscuration (Ach.) Massal. var. deminututn Zbr. The typ. sp. cold and temp, 
climates. Mf. 

Phyllopsoraceae. 
Phyllopsora ^F Arg. (25). Trop.- subtrop. 

part'ifolia (I'ers.) AF Arg. Widesi)read, also Chile, Hawaii and N. Zeal. — Mt. 

Cladoniaceae. 

Baeoinyccs Pers. (34). Majority trop.; 7 X. Zeal. 

chiUiisis (Mont.) Cromb. Chile. — Mt, Mf. 
C/adonid (Hill) Vain, (about 280). All over the world. 

pycnodada (Pers.) Xyl. Holiv., S. Chile to Fueg., Falkl.; Tristan da C.?—Mt, 



DERIVATION OF THE FLORA AND FAUNA 249 

Mf. To this belongs C. alpestris of Zahlbruckner zgd. 370, a species not occur- 
ring in the south hemisphere (Santesson J2y 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. 

<^gg^^g<^f<^ (Sw.) Ach. N. and S. Arner., Chile to Fueg., Falkl.; St. Helena, 
S. Afr., Madag., Auckl. and Campb. Is., Macquarie I.; Asia; Hawaii. — Mt, Mf. 

fiircata (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. 

finibriata (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. 

patagoniciiDi M. Lamb. S. Chile and Patag. to Fueg., Falkl. — Mt, Mf. 

ramtilosum (Sw.) Raensch. N. and S. Amer., Chile south to Magell., N. Zeal. 
— Mt, Mf. 

implexum 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. 
xanthophana (Nyl.) Jatta. S. Amer. mountains. — SC. 

Pertusariaceae. 

Coccotrenia M. Arg. (i). 

granulatiim (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. 

leioplaca (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. 
Mela7iaria 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 (Fee) Vain. (Braz.). — Mf. 



250 C. SKOTTSBERG 

coarctata (Sin.) Ach. Widespread temp.; Chile. — Mf. 

atra (Huds.) Acli. Cosmop.; Chile, Falkl.; Hawaii. — Mt, Mf. 

Ingae Zbr.— Mt, Mf. 

albilli)U} M. Arg. var. validior Zbr. The typ. sp. Fueg. — Mf. 

dis/^ersa (Pers.) Flk. Widespread N. hemisph. — Mt, Mf. 

polytropa (Fhrh.) Ach. Cosmop.; Chile, Falkl., W. Antarct.; Hawaii. — Mf. 

thrysolcuca (Sm.) Ach. W. Arct., Fur. mountains, Antarct., etc. — Mt. 

saxkola (Poll.) Ach. X. Amer., p:ur.; .^ Chile.— Mt. 
Placopsis Nyl. (31). Widely distrib., but mainly austral, 18 southern S. Amer. 

chilcua M. Lamb. Chile.— Mf. 

fuscidula M. Lamb. S. Chile to P^ueg., Tristan da C. — Mt, Mf. 

pareUina (Xyl.) M. Lamb. Boliv. And., Chile to P'ueg., Java, S, Austral., 
X. Zeal, Plawaii.— Mt. 

gelida (L.) Ach. Circumpol. Arct. and temp, oceanic; Chile, Tristan da C, 
Kerguel., Java, X. Zeal., Hawaii. — Mf (var. subreagens M. Lamb, but identity 
doubtful). 
Caudclariella M. Arg. (27). X. and S. Amer., Eur. 

intellina I\L Arg. Widespread; Chile, Falkl., Hawaii. — Mt, Mf. 
Myxodiityon Massal. (3, i Australia). 

chrysostictum (Tayl.) Mass. Chile, X. Zeal— Mt. M. lopadioides Zbr. 2g6. 383 
is a species of Lopadiiim (Santesson in letter). 

Parmeliaceae. 

J\iruie/ia (Ach.) De Xot. (about 400). World-wide. 

laevigata (Sm.) Ach. Widespread temp, and trop.; also Hawaii and Chile. 
— Mt. 

lacrigatula Xyl. Ikaz.- — Mt. 

rei'oluta P1k. Widespread temp. -trop. Also Hawaii. — Mf. 

cctrata Ach. W^y widespread temp, and trop., also Hawaii. — Mt, SC, Mf. 

saxatilis (L.) Ach. Widespread, in Chile south to F'ueg., Falkl; W. Antarct. 
— Mt. 

couspeysa (P'.hrh.) Ach. Cosmop., also Hawaii, P^alkl— Mt, SC, Mf. 

ahstrusa \'ain. Hraz. — Mf. 

polata Ach. Widespread; also Hawaii and S. Chile. — Mt. 

)tilglicrr€)isis Xyl Trop.-subtrop. — Mf. 

pilosella Hue. X. Amer. to Mex., PLur. — Mt, Mf. 

piloselloides Zbr. — Mt. 

cetrayioidt's Del Very widespread.— Mf. 

uncrosdcta M Arg. 'Prop. Amer. — Mt. 

caperata (L.) Ach. Temp, zones; Chile, Hawaii. — Mt, Mf. 

sored ica Xyl. Calif., Mex. — Mf. 
Meiiegazzia Mass. (30). P'ew north hemisj)h., majority S. Amer. (11) and Austral- 
'Pasm.-X. Zeal. (14). 

sauguinascens (Raes.) R. Sant. [^^28. i i, Parmelia pertusa (Schrad.) Schaer. in 
2g6. 389), Valdiv. to Fueg.— Mt, Mf. 



DERIVATION OF THE FLORA AND FAUNA 25 1 

Ratnalina Ach. (about lOo). World-wide. 

linearis (Sw.) Ach. Warmer regions; Chile, Fueg., Falkl., N. Zeal. — Mt. 

us7tea (L.) Howe Jr. Trop.-temp., N. Amer., Chile. — Mf. 
Usnea Wigg. (about lOo). Cosmop. 

dasypogoides Nyl. Rodriguez I. — Mt, Mf. 

florida (L.) Hoffm. Very widespread, also Chile. — Mf. 

subtorulosa (Zbr.) Motyka. Piaster 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 ace. 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 I.— 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 
to temp. N. and S. hemisph. — Mt, SC, Mf. 

orthoclada Zbr. In the vicinity of C Feliponei Zbr. (Urug.). — Mf. 

Teloschistaceae. 

Teloschistes Norm. (12). World-wide. 

flavicans (Sw.) M. Arg. Widespread trop.-subtrop., in S. Amer. south to 
Fueg., 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 falk- 
.landica Darb. (P^alkl.). — Mt. 

stellulata (Tayl.) Mudd. Cosmop. Chile; Kerguel— Mt, SC, Mf. 

halophila M. Arg. Australia.— Mt. 
• halophiloides Zbr. Easter \. — Mt. 

fei'nandeziana Zbr. Easter I. — Mt. 

masafuerana Zbr. — Mf. 

barrilensis Zbr. — Mf. 

Physciaceae. 
-Pyxine (Fr.) Nyl. (16). Warmer regions. 

curvatula Zbr. — Mt. 



C. SKOTTSBERG 



Pliyscia (Schreb.) X'aiii. (100-150). Cosmop., most numerous temp, regions. 

picta (Sw.) Xyl. Widespread trop.-subtrop. Also Hawaii. — Mt. 
Anaptychia Koerb. (30). Widespread, mainly warmer regions. 

Inpolcuca (Muhlb.) Massal. Widespread, also Hawaii. — Mt. 

pccti}iata (Zbr.) R. Sant. Patag. (Nahuelhuapi), Fueg. — Mt. 

Hymenolichenes. 

Cora Fr. (8). Trop.-subtrop. 

pai'onia (Sw.) Fr. IMex. -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 (i %) 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 Zaiilbruckner 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 
C'lara. The figure for Masafuera is 10 = 27.7%; of these 5 belong to the highland 
above 1 000 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. 
r\iture 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 liclien flora of Chile is not very well known. I have tried to find out if a 
s[)ecies called cosmopolitan has been recorded for Chile. Many world-wide lichens 
have been found in Juan l-'ernandcz; if also found in Chile they were referred 
to the Chilean element. 



DERIVATION OF THE FLORA AND FAUNA 253 

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. 

b. 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-Magellaniati element. — 14 (8.2 %). 

a. Endemic: Pertusaria hadrocarpa. 

b. 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 (153%). 
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. 

b. 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- 
padiuni leucoxanthum (endem. var.), Parmelia nilgherrensis, Caloplaca subcerina 
(endem. var.). 

IV. Pacific element. — 7 (4.1 %). 

Pscudoc\phellaria subvariabilis, l^acidia delapsans, Usnea subtorulosa, Calo- 
placa rubina, Huellia halophila, fernandeziana and halophiloides. 

V. Boreal element. — 16 (9.4%). 

if. luulemic: Leniniopsis polychidioides. 

/;. Not endemic (15): Arthonia cytisi (endem. var.), Gyalecta jenensis, Massa- 
longia carnosa, Lecidea enteroleuca, latypea and viridans, l^acidia arceutina 
(endem. var.), Riiizocarjxni obscuratum (endem. var.), Acarospora smaragdula, 
Lecanora dis{)ersa, chrysoleuca and saxicola. Parmelia pilosella and soredica, Buel- 
lia concinna (endem. var.). 

VI. Only reported from Juan Fernandez and Rodriguez I. 

Usnea dasy{)ogoides. 

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, 
Caloi)laca clandestina, Selkirkii and isidioclada, Buellia siphoniatula, masafuerana 
and barrilensis, I^yxine curvatula — further, Lopadium sp, and the dubious Blastenia 
sp. have been excluded. The {percentages were calculated w'ith 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 Basidiouiycctcs about 40 species were 
identified by RoMi:[,i, {2of), including those enumerated by JoilOW (several spe- 
cies doubtful), luidemic species few. Keisslkr's list of Ascomycetes (759, i6o\ 
with additions by AkWiDssoN (.?//), 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 I from N. Amer- 
ica; 5 are wides[oread and 5 endeiuic, one of these belonging to an endemic 
genus. Liimniiiia scoriadca (Berk.) Keissl. is also known from Chile, Java and New 
Zealand. Three of the endemic genera of Compositae have their special rusts, 
the endemic I'lupluasia is attacked by the same Urcdo that is found on two species 
of sect. Irifidac in Chile, Azara fcy)ia)ideziana by the rust known from A. integrifolia 
in Chile, and Rubus geoides is accompanied by the same parasite as in P^uegia (j_5(?). 

Of (lasteroDiycetes only 2 named s[)ecies were reported; o^ i\\QSQ lieodictyon 
jrraci/e l^erk. is of geographical interest: S. America, S. Africa, Australia and New 
Zealand, the second species was known before from N. America and Samoa [102). 



DERIVATION OF THE FLORA AND FAUNA 255 

The Myxophyta, i8 species, are more or. less cosmopolitan [loi). 

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 tjbergangsprovinz", i.e. Cen- 
tral Chile and the Valdivian forest region, Engler's transitional belt between the 
Andean and Magellanian provinces. Good [log) 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 JoilOW (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 (2.?^) 
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 [118. 
931): "Sur 142 Phanerogames indigenes, la moitie sont endemiques mais appa- 
rentees aux especes chiliennes; les autres sont cosmopolites ou Sudamericaines, 
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 difference between the 
islands and the continent. Ermei., who got his impression from a short visit to 
Masatierra, wrote (^7.48): 

. . . muss die Zusammengehorigkeit dieser Insel zum amerikanischen Festlande, zu 
welcher Ansicht deren geringe geographische Entfernung von selbst hinleiten konnte, aufs 
nachdriicklichste in Abrede gestellt werden, weil die beiderseitige Flora and Fauna zu 
grosse Verschiedenheiten aufweist, wo wir an geeigneter Stella die notigen Beweise bei- 
bringen werden. Die Gestaltung der Flora ordnet die Inselgruppe vielmehr dem aiistralischen 
Weltteile zu, von dem der grosste 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 tlie distribution of the non-endemic spe- 
cies, to state where the endemics have their relatives, if any^ and to distinguish 
a number of geograpliical elements. Now 1 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 (2^1. 203), until now we occupied our- 
selves with the geograpJiic elements, now we shall try to trace the ^r?/r//r 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 Stipa, necsiana is distributed from Mexico through the 
tro{)ical Andes to Central Chile and east to Brazil, Argentina and Uruguay, lae- 
nissinia (Xassella) a typical Andean species; the former is neotropic, the latter 
Chilean, extending north into Peru and east into Argentina. Almost the same 
area is occupied by riptocliaeiiuin bicolor, a genus limited to extratropical South 
America. 

rodopJioiHs is a unicjue anomaly without known neotropical affinities, as it 
were, a far-travelled member of an Arcto-tertiary flora, if its affinity with Brachy- 
clythruDi hits the mark; in Pil(;er's opinion it sides with MegalacJine. 

The sj)ccies of L'haetotropis 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 subtro[)ical and even extend into a temperate zone. 

Agrostis inasafucyana and the bicentric A. viagcllmiica were linked together, 
but f)nly provisionally, because in a large and world-wide genus like Agrostis the 
relationships cannot be safely judged without a thorough taxonomic-genetic study 
of the whole genus. Assuming that PllXiFR 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 chromostachymn is Chilean, but the genus is both N. and S. temper- 
ate, perhaps originally Arcto-tertiary? Danthonia collhta and Koelerm micrathera 
also are Chilean, but the distribution pattern of the genera indicates that Antarc- 
tica eventually was involved in their history. 

Megalackne, 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 Chusqiiea belongs to the neotropical element in the flora of Chile, 
and the same is true of the species of Cyperus. Scirpiis 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. Eleo- 
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 Oreobolus can hardly be dis- 
puted; it is often referred to as a classical example of an Antarctic genus. Of the 
6 species recognized by KUKENTIIAL, 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, i in N. Sumatra and Malacca, i in Hawaii, and O. obtusangu 
lus in Chile from the Cordilleras of Valdivia to Fuegia, the Falkland Is. and Juan 
Fernandez. [A seventh species, 0. pfeifferianus Barros, was identified by KUKEN- 
THAL [16^. vill) with pjunilio var. pectinaius\ 

Cladiiim (16^. Xll) is wide-ranging but it is not cosmopolitan in spite of the 
large areas occupied by C. 7nariscus L. and its varieties, among v^\\\q\\ j amaiceiise 
is circumpolar and distributed also south of the equator. The main distribution 
of subg. Machaerina (inch Vincentia), where C. scirpoideum 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 scirpoideum 
is not an American species but C. angustifolium (Gaud.) Benth. et Hook. fil. (New 
Guinea, Tahiti, Hawaii). Subg. Baumea, 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-Tasmania- 
New Zealand and i 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 Cladium may well lead back to the Mesozoic, and 
it seems natural to refer the genus to the Antarcto-tertiary element. The same 
applies to Uncinia, 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 



238 C. SKOTTSBKRG 

south, one species ^oin<^ nortli as tar as Mexico and the West Indies, i extending 
east from I^^ie^ia to the Falkhmd Is. and Tristan da Cunha. On the opposite side 
of Antarctica are 18 species. 14 of tliese indit^enous in New Zealand, where 8 are 
endemic, the remainder scattered alon^ tiie 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 1. has an endemic species. Finally, 2 species are found in New 
(iuinea, one of them also reported from Borneo, and the Philippines have I en- 
demic species, l^oth sections of sub^. liuriicijiia are represented in America, only 
Stenandrae in the opj)osite sector. The monotypical subg. Pseudocarcx is Magel- 
lanian. 

It is not easy to find one's way through the labyrinth of the enormous and 
still (growing ^cnus Carcx, 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. hfrttrojiiivia and the south Andean Baiiksii, 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.), 
I in Australia, 1 in Tasmania; the little known Chilean C. laniprocarpa Phil, and 
L . bfrti-yojiiaiia are far-flung outposts, but it lies near at hand to assume that they 
or their ancestors migratetl across Antarctica. C. Bavksii belongs to PVigidae- 
I'"uli<4inosae, a boreal group centering in luu'asia with one species in Pacific North 
America, but I cannot tell if luviksii comes near this species. 

The s\stematic position of Juiuiia austraiis was briefly discussed in 22g. 
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. HUTCIIIN- 
S(>N followed Hi:n'1 HAM and HooKKR in placing it next to Ceroxylon, but this 
genus is |)ol\gam()-monoecious and the stigma becomes basal in fruit. To me 
Ckni/A 1 s opinion lacks foundation; he solves what he calls "a hopeless conflict 
among taxonomists" (7/. 85) by deriving Juajiia from "a massive center of origin 
of angiospermy at the Mascarenes " (p. 103). 

Ochai:;ai'ia and I It'sf^crooyeiiria belong to an Andean-neotropical assemblage 
of genera, with close relati\es in Chile. 

To judge from Hr( IIK\ \i s monogra[)h of the Juncaceae in Pflanzenreich 
f.uzula uiasafucnvia nuist be referred to a grouj) of Andean species, L. racentosa 
Desv. (.Mex. S. Chile), cxcelsa Huch. (Boliv.), ///^vyw]'/;// Buch. (Argent.), /.^'^^^//V 
Buch. (C^hile), and cliiloisis .\ees et Mey. (Chile, south to l^'uegia), but in the same 
grou}) we find A. spicata (L.) DC. (Arct. circumj). and Alpine) and ahyssinica Pari. 
(ICthiop., Brit. \\. Afr.), and the |)()ssibilit}' 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 Marsippospennuui dind RostkovJay 
the isolated rnoinuiu in South Africa, endemic Andine genera like Oxychloc and 
Patosia, well marked endennc species of I.uzula in New Zealand and the Magel- 
lanian region, and the subantarctic-bicentric Jinicns scluuchzcrioides group. It is 
true that 4 of the 5 J// ;/r//.s- 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 25 9 

guay, the fifth, J. planifolius, shared by Chile and AustraHa-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 {281), 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 
tells a story of an Antarctic past, and the group Sisyrinchineae has a stronghold 
in the South, where the genera concentrate; Sisyrinchium itself centres in South 
America, where 4 small genera are endemic [Symphyosteinon, Chamaelum, Sole- 
fiomelus and Tapeinia); South Africa is another stronghold [Aristea, Wilsenia^ 
Bobartia, Klattia and Cleanthe), and '^[Orthosanthus, Diplarrhena and Paiersojiia) 
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. fernandezia?ia (Chile) is neo- 
tropical, while the other species appear to be more nearly allied to palaeotropical 
ones (Java, Australia, Oceania); P. berteroana 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 masafuerajia^ one of the few indigenous annuals, is related 
to a species from Ecuador, U. feriiandeziana with the habit of a miniature tree 
{22g. 862) appears to lack near relatives. The family is, I suppose, of tropical 
origin, but the actual centre of Urtica 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., i Australia, 2 N. Zeal, and i Auckl. Is. 
U. fernandeziana seems to represent an ancient type. 

Boehmeria excelsa was described by BuRGER (^7) 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 hiimiftisa (Chile); see above p. 204. Belongs to a neotropical group 
which has not been cleared up [241) but is also closely related to the Australian- 
Polynesian debilis of FORSTER. Weddell's debilis is a mixture of varieties scat- 
tered over the globe and most likely consists of several good species. 

Phrygilanthus Berteroi. The family Loranthaceae is \xo\i\Q.di\,h\xX.Phrygilanthus 
is of Antarctic origin. Of the 7 sections, 4 (about 20 species) are American and 
range from Lower California to S. Chile, i with 2 species belongs to Australia 
and New Zealand, i (i species) to New Zealand, and i has one species in Aus- 
tralia, another in New Guinea and a third in the Philippines — a typically austral- 
bicentric genus. 

The genus Santalum 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 (^70), Of the 4 sections distinguished by 



26o C. SKOTTSBERG 

TrVAMA (who divided my section Eusanialum in 2 and referred the Hawaiian 
tLusantala to a sejxirate section SolenaJithus), Fjisantalinu s. str. is the largest with 9 
species, SoloiaJithus and sect, llawaiiensia (together 8) confined to Hawaii (incl. Lay- 
san), and sect. Poly)icsica (2, with varieties) to Polynesia; in this section the extinct 
.V. fi-riiajideziajnofi occupies a rather independent position. "Aber woher kam der 
frenulartige l'>eniit Santaluni.-" lU'RC.KR [4.1. 22) exclaims, "zweifelsohne fern aus 
ostindischen Meercn, wo die W'iege seines altberiihmten 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, Mida in New Zealand, Jixocarpus ranges from Australia across 
Polynesia to Hawaii, and several endemic genera are at home in temperate S. 
America [Arjojia, lodiiia, Ouidia, QuincJianialiuui, and Xa7iodea in the extreme 
south and in the Falkland Is.); one is tempted to regard the family as of Antarctic 
ancestry, but with Thesiiini in mind it might be safer to speak of a special 
Antarcto-tertiary centre. 

The 3 endemic species of CheJiopodiuin were commented upon above (p. 204). 
In general appearance they are very like the Hawaiian oahuense but the special- 
ist oj)ines 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. 

SalicoDiia fruticosa, taken in a wide sense, is a thalassochorous plant scattered 
along tropical and subtropical coasts; the same is the case with Tetragoiiia ex- 
pausa in the S. hemisphere, but while Saliconiia is a world-wide genus, Tetra- 
i:;o)iia has a stronghold in South Africa and several endemic species in Chile. 

Sptrj^idar/a 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 .V. co7ifertiJiora is also found on San Ambrosio, are 
closely related to Chilean species. Paronychia has about the same distribution 
j)attcrn as Spcyo^ularia, but is poorly represented in S. America. 

RamDuulus capraniui. A very large essentially boreal-temperate genus with 
Avell stocked branches in S. America and on New Zealand. The Masafueran en- 
demic stands a|)art from its American congeners and approaches certain New Zea- 
land sj)ecies, j)crhaps also the Hawaiian ones. An Antarctic migration route seems 
probable. 

Fuyheris coryinbosa and niasafuerana 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 \\. and S. Asia and ranges far south only in America, where 3 
species reach I'\iegia. 

Of the six genera of W'interaceae (<\?), Bubbia has 2 species in Australia, 
I on Lord Howe I., <S in New Caledonia and 19 in New Guinea, Belliohun 4 
in New Caledonia and 4 in the Solomon Is., Pseudowintcra 2 in New Zealand, 
Jixospoimiin 2 and Zy^^^ogyjiu))! 6 in New Caledonia, and Drimys 6 in Australia 
{i also in Tasmania), 29 in New (Guinea and i on Borneo, Celebes and in the 
Philippines, all these belonging to sect. Tasma7iia\ the other section, Eudrimys, 
is American with 4 species, J), confcrtifolia 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 Drimys is proved by the Tertiary fossils 
discovered in West Antarctica. 

Lactoris 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 Wiriteraceae, of which it is probably a reduced 
derivative" [140. 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 07te 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. 

Cardamifte 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 Escallojiia, E. Callcotiiae stands a 
little apart from the rest {is8)\ the genus is spread along the Andes and extends 
to Brazil and Uruguay. The subfamily Escallonioideae is austral-circumpolar: Tri- 
beles (i S. Chile-Fueg.), Valdivia (i S. Chile), Forgesia (i) and Berenice (i), 
Reunion, Anopterus (2, Tasm., E. Austral.), Cuttsia (i E. Austral), Argophylliim 
(10 E. Austral., N. Caled.), Colmeiroa (i Lord Howe I.), Carpodeius (i N. Zeal., 
N. Guin.), Quijitinia (15 Austral. -N. Guin.-N. Caled., Philipp.), Potiingeria (i 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. Ill 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. 

Acae7ta. Nobody if not Croizat would argue that Rosaceae are a southern 
family, but this cannot prevent us from assuming that the Sanguisorba assemblage 
of genera has gone through part of its evolution at least in the far south. BiTTER 
{31) distinguished 10 sections: I, 13 species, S. Amer.; II, i. The Cape; III, 2, 
I S. Amer., i Hawaii; IV, i, S. Amer.; V, 8, 6 S. Amer., i J. Fern., i Tasm.; 
VI, I, S. Amer.; VII, 28, 26 S. Amer., i Cahf., i Tasm.; VIII, 64, 58 S. Amer., 
I Tristan da C, i N. Amsterd. I., 3 N. Zeal., i N. Zeal.-Tasm.-Austral.-N. Guin.; 
IX, I, N. Zeal.; and X, 2, N. Zeal. Two Magellanian species occur on S. Georgia 
and I 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. I^e this as 
it may, 7 sections are represented in America, 5 in the Australian-X. Zealand 
area, 3 are common to both, one of them ranging to the African sector, where 
another section is endemic. As regards A. i)iasafuera7ia see p. 206 above. I sup- 
j)ose we can draw no other conclusion from this distribution than that Acaeiia is 
an Antarcto-tertiary genus, 2 sections having developed numerous species in the 
Andes and Patagonia. 

Mari:;yricaypus, a small Andean genus, is so closely allied to Acaciia that they 
have j)roduced a bigeneric hybrid in Juan Fernandez. Two more genera are found 
in the Andes, TetvaglocJii}! and Polylepis. In S. Africa we have the large genus 
Cliffortia. The remaining genera Sangiijsorba, Poteriuiu and Beiicomia (Macarone- 
sia) belong to the X. hemis{)here. 

Sophoni sect. Tetrapterae is austral-circumpolar: New Zealand (3 species), 
Chatham I. (i). Lord Howe I. (i). Austral Is. (i), Rapa (i), Marquesas (i), Hawaii 
(i), I'^aster I. (i), Juan Fernandez (2), Chile (2; S. Diacrocarpa, however, rather unlike 
all the others), Diego Alvarez (i), and Reunion (i). With the exception o'i macrocarpa 
and the Hawaiian cJirysopJiylla 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 
subs{)ecies. Unless we believe that kS. tetraptera 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 
ada|:)tc(l to float, assisted by the four narrow wings, Joiiow 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. 

I'agara inayu and exterjia form their own section. When HC'RGER said [4.1. 19) 
that I'agara had migrated to Juan Fernandez from the primeval forests of Peru 
and Colombia he oxerlooked that the affinity is with palaeotropical rather than 
with neotropical si)ecies; there are numerous species scattered from Australia and 
Xew Caledonia to Polynesia and Hawaii, where many are endemic. Rutaceae were 
})erhaps represented in the Antarctic in Tertiary times, and we have too look for 
a route across to the American sector. 

The family liuphorbiaeeae is pantroj)ical, let alone that Euphorbia has at- 
tained a world-wide distribution and flourishes also in temjierate climates. Dysopsis 
is Andean, Seidelia (2) and Leidesia (i) South African, the fourth genus of the 
JSPercurialis grouj), Mercurialis (<S), ranges from Xorth l^urope to the Mediterranean 
and is found in \\. Asia. The southern genera seem to be more closely con- 
nected mutually than with Mereiirialis. 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 Callitriche 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- 
leri, 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 (Xaturl. Pflanzenfam. 2nd ed.); except Adolphia 
(Mexico-U.S.A.) the genera are distributed over Andean and extratropical S. Amer- 
ica, centering in Chile, and Colletia spartioides finds its place with the Andean 
element. It should be mentioned that Discaria (ii) extends south to Fuegia and 
reappears in Australia (i) and New Zealand (i), suggesting transantarctic migra- 
tion from America. 

The family Flacourtiaceae is tropical; Azara is neotropical with about 19 
species in Chile, i in Brazil and i 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 [Xylosma] 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 Tepualia, 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 Ugni 
Selkirkii ^v\d Myrteola Tiuniinularja see p. 206. NotJiomyrcia, now restricted toMasa- 
tierra, may or may not have inhabited a larger area. Myrceugenia 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; 6'/<';/;7^r<^ is composed of 6 subgenera. 
Pangue is the largest with 10 species ranging from Costa Rica to S. Chile, i in 
Brazil, 3 in Juan Fernandez and 2 in Hawaii. Perperisuvi is monotypical with 
separate varieties in S. Africa, British E. Africa and Madagascar, Ostenia an aber- 
rant monotype endemic in Uruguay. Misandra includes 3 species, one extending 
from Colombia to Fuegia and Falkland, one restricted to the S. Chilean Andes 
and one to subantarctic America; Milligajiia has 8 species in New Zealand and 
I in Tasmania. The monotypical Pseudogmmera inhabits New Guinea, Java, Su- 
matra and the Philippines. No subgenus is found in more than one sector. It is 



264 



C. SKOTTSBERG 



surprisinj^ that the geographically isolated Hawaiian species are closely related to 
the species of Juan I^>rnandez. We know other examples of this connection, but 
to construct a route between Hawaii and Juan Fernandez meets with serious obsta- 
cles, and Pa)igue may have reached Hawaii along a quite different route. The 
evolution and differentiation of the subgenera very likely took place in Antarctica. 

Scilixni-KR (Pflanzenreich) expressed his opinion of the history of Halor- 
rhagidaceac thus: "Aus der geographischen Verbreitungder ursprlinglichsten 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 Futterptianze erklart." But there was no Ilaloi'rJiagis on the mainland, 
and the species indigenous in Juan Fernandez were not used as forage, let it be 
that the introduced animals eat them. Halorrhagis, eminently Australian (59 out 
of 80 species) and with 7 species in New Zealand, extends north to Indomalaya, 
IVIicronesia, S.F. China and Japan and east to Rapa (not known to Schindler) 
and Juan h^rnandez — see Ti'YAMa's map of distribution [2yi]. 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" {log. 108). HalorrJiagis shows the same distribution pat- 
tern as ScDitaluDi. 

Coitclla is essentially African, see above p. 207; the distribution is tricentric 
with some remote stations. The widespread C. asiatica 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. triflora 
may well have reached Juan Fernandez with the traffic from Chile. 

The peculiar endeinic species of Eryjigiuui differ so much in habit from all 
other species of this large and widespread genus that they have been referred 
to a se{)arate genus, a rank to which they are not entitled, liryvgiuni concen- 
trates in tiie Mediterranean region and in tropical South America, where also the 
island species Ijelong, in sj)ite of their arborescent habit; to quote TURMKI. [26g. 
130): "Se rattachant a la Region chilienne, on cite les especes de Juan Fernandez 
li. hupleuvoidcs, sai'copJiylluiii et hiaccessuni . . . , plantes, du moins pour les deux 
premieres, arbustives s'o[)posant radicalement aux autres especes des territoires 
voisins"— but viacccssuni also is a dwarf tree, more so I would say than sarco- 
pliyllufti. I 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-Amnfineae is very widely spread, with a concentration 
in the \orth hemisphere, Apiuni distributed also in the south temperate zone, and 
A. gyai'eoh)is L. is frecjuently 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. fcDicDidcziajiuin 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, i Galapagos Is., 2 in Tasmania and 2 in New Zealand; P. rigida 
is linked to Andean species, but very distinct [2^2). 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 Gaultheria, 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 E. rubrum see 2^g.jd>\. 

Of the 7 Dichondra species 5 are neotropical, i endemic in New Zealand 
and D. repefis (incl. sericea) 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. sepiiwi 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. Hantelman7ii Phil, 
and later identified with tugurioriwi from New Zealand; see 2^g. 783. If this is 
correct, C. tugiiriorurn 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 Berteroi, the only representative of Boraginaceae, so richly devel- 
oped on the mainland, was regarded as an isolated, Old Pacific type [22"/. 31,.?^^. 
593) until Johnston {148) showed that it comes close to Hackelia and differs from 
this principally by its arboreous habit. With Urtica fernandezimia, the species of 
EryngiwH etc. I refer it to the neotropical element. 

Rhaphithannius vc7iustiis is of neotropical ancestry (see above p. 208); the 
second species is common in Centr. and S. Chile. 

Briquet placed Cuminia 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'Herit. (Canary Is., Colomb.-Peru, different sections). Ep- 
LING (using the fancy name JoJioivid) referred Cuminia to Prasioideae, an ancient 
group showing great disjunctions: Prasitmi Mediterranean, Stenogyne, Phyllostegia 
and Haplosiachys in Hawaii, Bosirychmtthus 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 ^. robinsonia?ium , see above p. 200. 

Nicotiajta cordifolia has, according to GOODSPEED [112. 347), its closest resem- 
blance, in flower structure as well as in general habit, to N. Raimondii. 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 (I.e.). 

Mi})iulus is essentiall)' a western N. American genus with few species else- 
where; the somewhat polymorphous M. glabratus ranges from N. America to 
Bolivia, Chile and Argentina. 

The different opinions on the systematic \)os\t\on o{ Euphrasia forifios is siifi a 
have already been referred to above. Within the area of the geographically isolated 
Chilean-Magellanian 'Irifidac occurs the semicalcarate E. pcrpitsillaV\\\\. (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 Dl' RiETZ thinks (77. 536): "The Euphrasia popu- 
lation of Juan r>rnandez may therefor very well have formed the northern end 
of a population so far south that the lack of close relationships between E.for- 
niosissiDia and the species of Middle Chile is fully explained, and the semical- 
carate anthers of the more southern E. perpusilla may be the last South American 
remnant of this old connection." Another question, not yet answered, is this: are 
the very well-marked Trifidae, reaching from subantarctic America to the Andes 
of S. Chile, likewise descendants from a remote southern population or did they 
originate in Chile? 

Plantago fernaiuiezia 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. Palaeo- 
psyliiuui (see above p. 209) its character of an austral group is revealed, even if 
it extends north to X. 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 truucaia, represented in Juan Fernandez by an endemic (.') variety, 
belongs to the large section Xoiwrbis and needs no further comments. 

The neotropical Ilcdyotis ihesiifolia is a recent addition to the flora and may 
have been accidentally introduced. 

Xcrtcra is an austral circumpolar genus allied to Coprosvia and best devel- 
oped in Xcw Zealand where 5 species occur, 4 of them endemic; the fifth is 
X. grauadfusis (depressa), claimed to be very widely spread. One species is en- 
demic in Tristan da Cunha. Recent observations tend to show that granaderisis 
includes taxonomically distinct forms; the Malaysian plant is not identical with 
the iXndean, and other forms will perhaps become distinguished after a critical 
revision. He this as it may, the genus is Antarcto-tertiary and, if we link Tristan 
da Cunha with Africa, tiicentric. 

CoprosDia is a parallel to I lalurrJiagis 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 — lloolccri 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 o{ pyrifolia, and to the believers in transoceanic 
migration combined with evolution of local endemics wherever (T^/r^j-;;/^ 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 Hookeri becomes somewhat embarrassing. With an extension north 
and east of an Antarctic borderland Coprosnia 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 (^j. 20), "von den Urwaldern Perus und Co- 
lombias". 

Of the more than 500 species described under Galitun 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. Galiiun is a boreal genus with a strong represen- 
tation along the Andes, a not uncommon case. G. masafuera?inm 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 
Brighamia 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 (i), Australia (7), 
New Zealand (7), Lord Howe I. (2) and St. Helena (3). Species are few in America 
(North Amer. i, South Amer. 8); a single species, W. gracilis, 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, i is from tropical Africa, E. Indies and Brazil, 4 Asiatic 
and I from S.E. Europe, all according to the old synopsis in Natiirl. 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 W. Masaftierae 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 Lagowphora 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, i in Fiji 
and 3 in Hawaii, the four Pacific ones connected with L. Bil/ardieri Cdiss. (Austral.), 
whereas the Magellanian species point toward L. puiuila Cheesein. 2iX\d. petiolata 
Hook. fil. (New Zealand). 

In Iiyii:;tyo}i we meet again with a large boreal genus strongly represented 
along the Andes and ranging south to Patagonia, P^iegia and P'alkland; 135 species 
are reported from Kurasia, inch 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), E. rapensis 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 
I lawaii the genus is represented by l^ctra)nolopiu7)i, which is very close to Erigeroit\ 
\'lKKii.\ri'KR [22g. 182) suggested that the Juan P^rnandez species came near the 
Hawaiian /:. lepidotus Less., which is now referred to Tetraniolopmnt, and that we 
ought to look for relatives among the Andine and Mexican forms. Among our 
island species li. nipicola differs much in habit from the rest, but all belong together 
and most likely represent a special branch of the Andine Erigeroii flora which, in 
its turn, comes from a boreal stock. 

To what I said above about GnaphaEiuj>i spicifornie nothing can be added 
at present. Chile is well provided with poorly limited species related to G. pur- 
pur ciiui L.; most of them are badly known. Their boreal parentage can hardly 
be doubted. 

Ahrotajiclla resembles LagejiopJiora in its distribution, but is absent from Oce- 
ania; tiie majority inhabits New Zealand with its subantarctic islands (9 sp.), i is 
found in X'ictoria, 2 in Tasmania, i in New Guinea and 5 in S. America (W. 
Patagonia to h\iegia and P'alkland); in addition, one is found on Rodriguez I. 
Tiie genus is, as it were, tricentric and its Antarcto-tertiary character indisputable. 

The concentration of isolated arborescent Compositae in the Pacific was em- 
|)hasized by Bkn'IIIAM; tiie main grouj)S of the family are represented among them^ 
and the accunuilation of endemic genera in Hawaii and Juan P^rnandez has led 
to nuich discussion. Tiic 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 (ill'l'V [121) does not seem [)ossible. Speculations (comp. 
f. inst. Sl'/i (111:1.1, 2r()) led back to Antarctica, but not until BracJiioioslyluin was 
discovered in .\ew (iuinea and found to be nearly related to Robinsoiiia o'i ]\x^Vi 
l^>rnandez were we able to stand on tolerably firm ground. 

With regard to Geiitaurodoidroji, to which J//';/*//^'^^'? seems to be related, the 
situation is different. Coitaurca and all the genera of Centaureinae belong to the 
Old World with the exce})tion of a single species in N. America and a few in 
the Andes, south to Centr. Cliile. Tlie grou{) ranges over luirope with a strong- 
hold in the Mediterranean, the Orient, P'.thiopia and through Centr. Asia to Japan. 
The Chilean species of Centaurca belong to the Plectocephali, but Ceiitauroden- 
droji differs not only from this section but from all in sex distribution (24J). It 



DERIVATION OF THE FLORA AND FAUNA 269 

serves no purpose to say that Centaurodendron 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 Centaurea, a relic from an epoch previous to the 
final uplift of the Andes, 

The four Cichoriaceous genera, to which Z//<3'w;/d7^^;7V 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 Fitchia, but the differences 
are too profound to allow us to visualize an Antarctic-Pacific ancestry of the Den- 
droseris 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. 

b. Species also found on the continent (16): Danthonia collina, Koeleria micra- 
thera (.f"), 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 Berterii. 

b. 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. — i sp. 

II. Neotropical- Andean element. — 54 sp. (36.7 %). 

I. Endemic genera or species of neotropical parentage; non-endemic species 
South American. — 31 sp. 

a. Belonging to endemic genera (4): Juania australis, Ochagavia elegans, Notho- 
myrcia fernandeziana, Selkirkia Berteroi. 



270 C. SKOTTSBKRG 

b. luidcmic species of non-endemic genera, some of wider distribution (14): 
Chaetotropis imberbis. Chusquea fernandeziana, Hesperogreigia Berteroi, Urtica 
Masafiierae, Dysopsis hirsuta. Colletia spartioidcs, Azara fernandeziana, Ugni Sel- 
kirkii, Myrceugenia Schulzei, Rhaphithamnus venustus, Solanum fernandezianum, 
robinsonianum and masafiieranum, Xicotiana cordifolia. 

c. Sj)ecies also found on the continent (13): Stipa neesiana and laevissima, 
Piptochaetium bicolor, Chaetotropis chilensis. Cyperus eragrostis and reflexus, Eleo- 
charis fuscopurj)urea, Peperomia fernandeziana, Parietaria humifusa, Myrteola num- 
mularia, Minnilus ghibratus, Phuitago truncata, Hedyotis thesiifolia. 

2. luidemic 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. Ik'longing 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 j)ruinata, IMioenicoseris j^nnata, berteriana and regia, Hesperoseris gigantea. 

h. Ik'longing to endemic sections (7): Urtica fernandeziana, Chenopodium 
Sanctae Clarae, crusoeanum and nesodendron, Eryngium bupleuroides, inaccessum 
and sarc()j)h\'llum. 

III. Arcto-tertiary element. — 23 sp. (15.6%). 

Cicncra essentially boreal but extending south along the Andes or reappearing in S. 

America. 

a. luidemic 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. 

b. Species also fountl on the continent (lo): Trisetum chromostachyum, Carex 
Banksii, Paronychia chilensis, Cardamine chenopodiifolia and flaccida, Rubus geo- 
ides, Callitriche Lechleri, P^mpetrum rubrum, Calystegia tuguriorum(.?), Gnaphalium 
s{)ici forme. 

IV. Palaeotropic element. — 2 sp. (1.4%). 

Plndemic genus: Cuminia fernandezia and eriantha. 

V. Austral element of wide-ranging seaside species. — 6 sj). (4,1 %). 

Scirpus nodosus and ccrnuus, Salicornia fruticosa (peruviana), Tetragonia ex- 
[)ansa. Dichondra repens. Lobelia alata. 

I'Acn 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 cxceptif)ns, aj)i)ly to I. i, 22.4% of the total or 53.2% of I. 
Luzula and Jiiucus were [)laced here because the family was regarded as Antarctic, 
but even if this be true it is po.ssible that the sections including our species are 



DERIVATION OF THE FLORA AND FAUNA 27I 

of boreal origin and has spread south. Pernettya 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 Nicotiana according to Merrill 
(jo<5. 310) who thinks that its actual distribution was attained in the Tertiary pe- 
riod by way of Antarctica; however, N. cordifolia cannot be removed from II. i. b 
and find a better place with I. i.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 Peperomm berteroana 
calls for attention. As I have shown (^^5) it is so close to P. tristanejish that 
their common origin cannot be doubted and that little prevents us from regarding 
them as forms of one 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. i only Juaiiia 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. Bentiiam looked upon the arborescent Pacific 
Compositae as relics of an old Polynesian flora but did not refer directly to Ant- 
arctica as source; Guppy {^121] quoting Bentham believed that, with the Hawaiian 
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 [2/f.8), 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 [^218) 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 
{2ig) 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, I 
did not venture to include the Cichoriaceous genera, but I cannot assure that 
Setchell was wrong. 



272 



C. SKOTTSBKRG 



Before quotin*]^ some other authors who have paid special attention to the 
Antarctic i)roblems 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, advancinj^ especially along the rising Andes and in some cases 
extending north of the Iu]uator. 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, r\iegia 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. I^^ather 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 Acae?ia, Margyricarptis, Es- 
callojiia, GuuJicra, /)r/;^n's and PJirygilanthusdiX^ represented; in 202. 30 he mentions 
Ayaucaria ajigusiifolia, Podocarpus Lmnberti, Drimys Wmteri, Acae7ia fuscesceiis, 
h'tichsia regia, Ciumicra manicata, and Griselhija riiscifolia, elements which, in his 
opinion, constitute the last remnants of the old flora that inhabited the southern 
lands united until early Tertiary times. 

IliLL (/iV) 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 
rea( lied the south, evolve along special lines in lands comparatively isolated from the 
land masses in the north, or did certain grouj)S first ap])ear in the South Hemisphere 
in an ancient Antarctic Continent and become dis})ersed northward into our j)resent- 
(lay New Zealand, Australia, South America and South Africa? 

With regard to Juan h'ernandez 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 prol)al)ly 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 ei)0ch towards Tasmania and Australia, and so to Asia, and towards New Zea- 
land, and the distribution of certain j)resent-day ])lants in the Australasian region lends 
consideral)]e suj)port 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, Jovellana, Fuchsia, 
Ourisia, Pelargonium, Caltha, Lilaeopsis, Gunnera, Hebe, Fernettya, Azorella, Drapeies 
and other Thymelaeaceae, Noihofagus, 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 w^hich 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, w^hen 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 (4.J) interests us here. My Antarcto-tertiary 
element lists Drimys 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 Jovellana, all as a rule considered to 
be Antarctic; here Camp has two alternatives, one track leading from the north 
as in Drimys, 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 T^^e Nat. Hist, of Juan Fernandez and Easter Isl. Vol. I 



74 



C. SKOTTSBERG 



majority of our basic angiosi)erm 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. 180-181), 

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 hind mass certainly by the mid-Mesozoic. 

This would make the Jurassic the great period of ascendency of the angiosperms. 

C'ami'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- 
Xew Zealand, and the actual distribution of this family testifies to a southern origin. 

Recently SlKHBlNS (^57) took up the question of the history of the Ranales. 
He points out (p. 8) that this order » 

includes a high j)roj)ortion of sj)ecies which on the basis of all characteristics must be 
])laccd not only in monotypic genera but even in monogeneric or digeneric families. 
Thcv 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 (lod) 
and perhaps strengthened by the sensational discovery there of a great number of 
Xothofagus 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 com[)els 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, Xothofa^i^iis 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 
s{)read soutii during the Tertiary just as, in pre-Tertiary times, tropical families 
had extended to the j)rcsent Arctic. GoRDuX (77?), 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 :i genus like Xothofagus originated in \. America and that two tracks 
lead sf)uth, one to subantarctic America, one across the Ikring land to Australia 
and Xew 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 (pj) 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 Ll^ON Croizat (yi), who throws all other 
speculative authors in the shade. His ideas were criticised by Sparre (2j6), 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 Pringlea and Lyallia 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 l>>rnandez, endemic or not, are mentioned by Croizat ; 
several of them are, also in my opinion, of Antarctic origin. 

Onoholus. Instead of allowing it to enter S. America through the "Magellanian 
gate ' Ckoiza r makes a circuit along his Pacific baseline from New Zealand to F'uegia. 

Juauia. I ha\ e already discussed the taxonomic position of this genus which 
Ckoi/ai' derives from a Mascarene centre in company with related Andine genera. 
Ik nillNSON linked it with Ceruxyloii to form a tribe Ceroxyleae; Croizat remarks 
that 

this mav be better taxonomy, we do not know, but does not change substances in the 
least, 'i'he ingredients that make up the Morenieae, Iriarteae and Ceroxyleae are all from 
the same kitchen. In one case Hyophorbe speaks, in another so does the face of dispersal. 
Kvervthing in the end rhymes to the very same. With or without Hyophorbe, seasoned 
with this or that sauce, the stew does not change . . . classification of such a genus as 
this is bound forever to remain the j)laything of opinion (pp. 456-457), 

a statement quite characteristic of this author's method of approaching a phyto- 
geogra[)hical problem. 

Pipcyo))iia has its stronghold in tropical America where a number of subgenera 
are found. To CROIZAT the genus is Pacific and Juan F'ernandez one of the starting- 
points of its dispersal toward the heart of S. America, the four island species leading 
straight to north Chile, lie based this opinion on YUNCKER who wants to bring 
them to subgen. SphacrocarpidiuDi^ 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. ty'istanciisis, which, unexpectedly it is true, brings the Antarctic 
within sight. In a footnote CkOlZA T admits that I may be right in my opinion of 
the taxonomic {)osition of P. bcrtcroana\ none the less he finds the contact Juan 
hYM'nandez- Tristan not at all j)eculiar. EDipetruni is another example, both "a 
mere asj)cct f)f the standard dispersal between Africa and South America" (p. 102). 
Hut tiiis "standard dispersal", as discussed by Engler, refers to a number of 
tr()j)ical genera and families bearing witness of a transatlantic bridge. In Croizat's 
opinion the track Mascarenes-Africa-America was a very important route. 

I'alms and Pepcromias are pantropical and range south to New Zealand and 
central Chile, and there are man\' endemic palm genera on the Indian and Pacific 
islands south of the Ivpiator, whence follows that the field is open for speculation 
and that the possibilit)- of a ])rimary 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, Kai)ateaceae etc. — and in an earlier chapter he asserts that 
Malesherbiaceae also had their starting j)oint in Africa — says that 

to credit grouj)s of the kind with "American" origin steps must be taken as a prelim- 
inary which no one trained in |)rccise thinking will encourage. . . . Rhipsalis is doubtless 
a (iondwanic genus because it is rooted within the triangle Ceylon-Madagascar-Masca- 
renes. It well may have reached ]')razil and the West Indies from West Africa. . . . the 
baseline of the Cactaceae scpiarely 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 Afroantarctic triangle (pp. 364, 365). 

Much could be said to this, but I shall confine myself to remarking that the 
claims of Rhipsalis 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 uniformly 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. 

Dysopsis. 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 Anlarctic 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 oi Penietiya 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 Empetrum 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 Cuminia (for which Croizat uses the fancy name Johowia), whether 
or not to be linked with the Old World and Hawaiian Prasioideae, is embarrassing: 



278 C. SKOTTSBERG 

my discussion with F. H. H. l^Rowx [jji] made Croizat remark that "the debate 
proved futile, as could be anticipated". To attempt a solution and to discuss possible 
routes of mi<i;ration is called "obvious nonsense". 

Of the presumable history of liupJirasia enough has been said above. In Croi- 
ZAl's opinion there cannot be the cjuestion 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. 

I'rom the lu'pothetical "main angiosi)ermous center" south of Madagascar, 
where Plantao^o-Pa/afopsyl/iii/fi 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 P^rnandez. 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. 

Copros))ia is one of the standard Antarctic dispersal types but lacking in Africa 
and in continental America. P>om 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 disj)ersal is all westward from the Americas (Juan Fernandez to New Zealand, Aus- 
tralia, New (iuinea, Malaysia) ultimately veering again eastward to reach Hawaii (p. 101). 

To make Juan P^ernandez the source of Coprosnia 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 F'ernandez. 

1 am not going to enter upon a discussion of the origin of a world-wide family 
like Com{)ositae; I shall only quote the following sentence: "The perfect circum- 
[)olar distribution of I'araxacuui viagellanicuin ... 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 
(/■r) who gives us a palaeontologist's view on angiosperm evolution. 

The cradle of angiosperms stood in the tro})ical 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. P\)ssil 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 (i) 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 (dp), 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 (dy. 174) that "no 
other plant family of its size and diversity is quite so conspicuously Antarctic in 
origin as this one". 

Trichommies-VandenboscJiia is pantropical; as 2 of the 3 insular species are 
decidedly neotropical in their affinities, also T. phzlippianum 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. 

Hymeiiophyllum-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 o{fuciforme. H. pectinatum stands, COPELANI) says, apart from the other 
species, but I can find no good reason to call in Antarctic, while the distribution 
of Sphaerocionium ferrugineuni and its near relatives testifies to their Antarctic origin 
in spite of their affinity to ciliatiim (comp. above p. 219). Hymenophylluni ^. str. 



28o C. SKOTTSBERG 

as limited by Coi'Ki.AM) 1938 (66) does not \x\q\w<\^ falklandicum [Mecodium accord- 
ing to CcM'KLAXD p. 94), though it is closely akin to peitatiim, and H. riigosiim 
"is to //. tuuhridi::€US€ what H . falklandiciim is to H. peltatuin' (62. 13). According 
to CliRisr (-r^*. 146), pi'ltattou and tiinhridgense 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 L2urope. The conclusion would be that also 
falkUvidicuni and rNgosiiffi should be brought to the Antarctic group. 

IlymoiopJivlluni-Mey'nigiuDi secundum, plicatiiui and tortuosum are subantarctic- 
American in their present distribution; JMcrijigiuni is austral and extends north to 
tropical Asia, and its origin most likely Antarctic. 

'Iliyrsoptcris. 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). 
llll.l. [1J4. 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 COPELANI) [6y . 175) writes: 

As juan l-'ernandez 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. Kuropean fossils have been referred to this genus, but Juan Fernandez was not 
colonized from FAirope. 

I.op/iosor/a is exclusively neotropical at present, but if we are to believe, as 
C<)i'i:i,AM) thinks, that the Cyatheaceae derive from Antarctica, it is unlikely that 
Lopliosaria would be an exception. 

The Antarctic origin of Dicksonia 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-Xew Caledonia-Fiji (-Samoa)-Malaysia 14 species are 
found, along the Andean path 9, and that the species of Juan F^ernandez are not 
linked with the neotropical but with southwest Pacific species. 

'Fhc present distribution of the genus Cystopteris suggests a boreal origin. 

In its wide as well as in its more restricted sense Dryoptcris is world-wide, 
and C()i'i;i and's remark (^7. 181) that he has little doubt that the group as a 
whole is of southern origin (see also 6g. 122) is perhaps little more than a guess. 
I think we can take it ior granted that hundreds of species have evolved in the 
neotropical region, where the single Juan FY-rnandez species D. inaequalij'olia has 
its relatives. 

rolystiiliuui . Mistern .Asia is the centre of greatest concentration ; the section 
occurring there is, in C( )ii;i,AM)'s oj)inion, the least primitive, and we have a number 
ot southern hi- or tricentric frjrms, among them P. vcsiiiuvi coll. The endemic P. 
bertiricDiuui is very close to the circumj)()lar adicDitiforme. To quote CoPKLAND: "The 
case for ;ui Antarctic origin of Polysticlnou is so clear that the evidence has long 
been familiar even t(j those hesitant to draw the obvious conclusion" [dj. 1 81); 
and, "Tile distribution . . . testifies clearly to its Antarctic origin" [6g. 109). 

Aythroptcris is the only fern genus found in Juan Fernandez which is absent 
from continental America, but otherwi.se typically austral. Its Antarctic origin is 



DERIVATION OF THE FLORA AND FAUNA 28l 

beyond doubt, and A. altescajidens has its nearest allies in Oceania; comp. above 

p. 222. 

Aspleniwn. 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" {6g. 167). Of the species found in Juan Fernandez, obliquum 
is austral-bicentric, stellaimn close to neotropical species, macrosorum most likely 
of neotropical affinity; day^eoides comes very near alvarezerise from Tristan da 
Cunha and is not, as has been suggested, related to the boreal ruta muraria. 

Blechnum. According to Copeland (^9.157) 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 auriculatmn , chilense and cy- 
cadifolium belong to an Afro-american group, Schottii and perhaps also valdiviefise 
of austral-circumpolar type, and longicauda intimately related to the neotropical 
Spriicei\ even so it cannot blur the picture of an old Antarctic genus. 

Pellaea is an austral-tricentric genus extending north to Canada, but P. chi- 
lensis, endemic in Juan Fernandez, but dangerously close to a widespread American 
species, belongs to a group that, according to Copeland [6^. 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 'iox^otrugo- 
sula. This species is bicentric or, including Polypodium villosoviscidum of Tristan, 
tricentric; there is another variety on St. Helena. Thus when CoPELAND 6y. lyy 
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. 

Adimituin is most numerous in S. America, and A. chilense and related species 
tell us nothing of the earlier history of the genus; CoPELAND 6g. 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 dy. 178 he 
says that "Antarctica has played a major part in its history", but that the genus 
"may include elements surviving from other floras and from an age prior to the 
great dispersal from the south". I refer A. ckilefise 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. chileiisis is of neo- 
tropical character, perhaps also P. semiadnata, while berteroana 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 [6g.6o), "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 (<57. 184), "a plain Antarctic case", and G. Bil- 
lardieri "surrounds Antarctica more closely and completely than does any other 



282 C. SKOTTSBERG 

fern ' [dg. 21 1; see above p. 223); it was at that time supposed to be circum- 
polar, but is replaced in S. America by /\ DiagclUmicum. As mentioned above, 
I believe that if a t>enus Syiiaimiiia with PolypodiiDii Feuillei 2,s the typical species 
is acce{)ted, the endemic P. i>iier)iiedin)n finds its place with this, but I fail to see 
why Coi'KLANi) y6S) regards .V. Ju'uil/ii di^ "clearly Antarctic", whereas the nearly 
relatetl CofiiophUhiuDi is called "a northern genus". Of the remaining species 
P. Masafiicrae and P. (Xij)ho|)teris) trichouajioidcs are neotropical and P. (Pleo- 
peltis) lajiceolatio)! pantropical but not reported from Australia and not indicating 
an Antarctic origin. 

We have no good reason to look for an Antarctic ancestry of Elaphoglossmn, 
though C'()ii;i,.\M) thinks that it "may have come from the south" {dj. 185), and 
P. Li)idi)iii is a neotropical species. 

The species of Gleichoiia occurring in Juan Fernandez belong to Sticherus, 
a genus segregated by some modern authors; 11 species are scattered over the 
austral zone, and in Cotkland's opinion Gleicheniaceae are "obviously and en- 
tirely of Antarctic ancestry" (<5^. 26), or "entirely Antarctic at some stage of its 
histor}- ' [dy. 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 Ophioglossaceac are an ancient family, "scattered with remarkable uni- 
formity over the habitable globe" (CoPELANl) 6g. 12). In dy. 167 he paid special 
attention to Hotrychiion australe^. 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 
I'\iegia is not this, but the northern B. lunaria L. We have no reason to regard 
OphioglossiDn fcy}ia)idi-zia)ium as coming from the south. 

Lycopodiion was not discussed by CoPELANi). L. magellaiiiaiin is subantarctic- 
circumpolar, and part of the history of the genus may have been enacted in Ant- 
arctica, the more so as L. scariosuni^ 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. ICndemic sj)ecies (2): Hymenophyllum rugosum, Polystichum berterianum. 
h. .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, Hlechnum valdiviense and chilense, Hypolepis rugosula 
(w), Ilistiopteris incisa (w), Polypodium magellanicum (w), Gleichenia pedalis, 
quadripartita and cf. litoralis, Lycopodium magellanicum (w) and scariosum (w). 

2. S[)ecies 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. 

4. Species closely related to S. African ones. — 2 sp. 

a. Endemic: Blechnum cycadifolium. 

b. Also on the mainland: Blechnum auriculatum. 

5. Endemic species very nearly related to a neotropical species: Blechnum 
longicauda. — i sp. 

6. Endemic genus without affinities to living genera: Thyrsopteris elegans. 
— I sp. 

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. 

b. 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- 
inanoides, Elaphoglossum Lindenii. 



III. Arcto-teftiary element. — i 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 
«ven 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 ((5/. 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, CorELAND remarks, if the ferns as a group 
were of tropical origin the\- 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 da\', this fact is a strong indication of its Antarctic origin and that, if the pre- 
sent range is wholl}' 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 i)resent distribution, because tropical and northern species must outnumber the whole 
tlora of Antarctic America, or even of New Zealand {6y . 158). 

With this he wants so sa}% I suppose, that a vigorous family has invaded the 
tr()j)ics and also extended north under rapid evolution of genera and species con- 
cealing the j)rimary origin; he believes this evolution to have been so rapid that 
the j)eriod 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 (I.e.). 

We have no evidence that the Tropics as a whole were at any past time unfit for ferns; 
and the assumi)tion that this has been the chief place of evolution obviates the neces- 
sity of assuming and explaining migration in latitude. It is only w-hen we open our 
eyes to anomalies in present distribution that appeal to other places of evolution be- 
( omcs necessary [6y. 163). 

(ireat geographical disjunctions, of which the Juan Fernandez fern flora offers 
man\- 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 Alagellanian moss flora Cardot (48) distinguished an 
Antarctic element and he believed in a common origin of the floras of Magella- 
nia and New Zealand (j). 44). Hkkzoc; (i2g) found that Juan Fernandez has "eine 
last \-ollstandig austral antarktische Moosvegetation", but also possesses "ein paar 
tropisch annuitende Artcn: ThysaiioDiiiriuni RicJiardi, Porothanniium fasciculaiuvi, 
Pi)niatilla luaooslicta und ein kliacopiliDiiy The monotypical Chilean genus Lam- 
l^yof'hylluDi 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 
l'\iegia; characteristic genera, also re[)resented in Juan Fernandez, are Dkraiioloma, 
( lota, Dindyocryf^hacd, I .cpyycuhi}!, rtycliouinium, Weyniouthia, DisticJiophylluni^ 
r/in\^-(>/^/iyI/in//, liriof^us, J f]f(phn\i^ju»i, Sciaro))iiiu)i, perhaps also CatagoJiiopsis 
and Psihpilii))!. Ii<ms( [ikr (//,') gives many examples of austral-bicentric taxa: 
Lcpyrodo)itac(ai\ /\>Iy/r/i/i/i</i//^//!(s^ Wcymouthia, PtcrygopJiylluni, Sciaromiuin sect. 
Aloma, Hyp(ptn-yo;iu))i sect. Stoiobasis, I lypnodcndron, and species of Disticko- 
phylluui^ J ovinia, Macyoviityiuui and MielichJioferia. 



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; 1 1, belonging to large and widespread genera were 
left out (see list p. 235). The 8 species of Tham7imm, all endemic, were tentatively 
included in group IV. In Natiirl. 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, i 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, 3 N. 
America, 3 mediterr.-macaronesian, i England, i 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), Rhacomitrium 
symphyodontum (t), Zygodon intermedins 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. 

b. 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 C. SKOTTSBERG 

II. Neotropical element. — 26 sp. (21.7 %). 

a. ICndemic species (11): Campylopus subareodictyon, Leptodontium fernan- 
dezianum, Didymodon calymperidictyon and linearis, Bryum fernandezianum, Pin- 
natclla niacrosticta, Isopterygium fernandezianum, Rhaphidostegium Masafuerae 
and caespitosoides, Rigodiuni robustum and Looseri. 

Ik Also in S. America (15): Gymnostomum calcareum (widely dispersed), Cam- 
pyl()l)us areodictyon, Thysanomitrium Richardi, Mielichhoferia longiseta, Anacolia 
siibsessilis, Porothamnium fasciculatiim and arbusculans, Lamprophyllum splendi- 
dissimum, Rhacopilum fernandezianum, Stereodon Lechleri, Rhaphidostegium 
caespitosum, Rigodium toxarium, arborescens, hylocomioides and tamarix. 

III. Chilean element. — 24 sp. (20.0 %). 

(7. luidemic species (4): Fissidens pycnotylus, Ulota fernandeziana, Philonotis 
glabrata, Rhaphidostegium aberrans. 

/\ Also in Chile, mostly southern (17): Pleuridium Robinsonii, Hymenosto- 
mum kunzcanum, 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. 

(-. Cosmopolitan (3): Ceratodon purpureus, Rhacomitrium lanuginosum, Funa- 
ria hygrometrica. 

Xoie. — On p. 234 Campylopus introfJexus was placed here; it was transferred to 
I: I 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 j)antroj)i(al species which has spread both north and south. 

IV. Atlantic-Mediterranean element. — 3 sp. (2.5 %). 
Comp. ahove, p. 235. 

a. I'Jidcmic: h'issidcns crassicuspes. 

h. \ot endemic: Campylo[His polytrichoides, Trichostomum brachydontium. 

W'c have seen above that Hkrzog regarded the moss flora of Juan P^ernan- 
dcz 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 l\)r()tJiai)niiui}i arbusculans is, as far as I know, restricted 
to Chile and I'atagonia, but the genus is essentially tropical, Stereodon Lechlen 
extends from S. Chile to W. I'atagonia, but is the only species reported south 
of the lujuator; Rii^odiuDi 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 s[)ecies 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 (210) 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. Ditrichum affine, Trichosiomum brachydontium, Rhacomiirium symphyo- 
dontum, Bartramia patens, and Pterygophyllum denticulatum, listed for New Zealand 
by Brotherus (j4), 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 UOMlN (yd) lays 
stress upon that they are just as specialized geographically as the flowering 
plants, and 

einen uralten, heiitzutage in den gemassigten iind kalteren Gebieten sozusagen erstarr- 
ten, sich von ihren ganz speziellen Standorten nicht weiter ausbreitenden, wenig an- 
passungsfahigen Typus darstellen, welcher auf der nordlichen Hemisphare seinen Ent- 
wickelungsgang in weit zuriickliegenden Epochen durchgemacht iind 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 Schistochila, 
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 Riccardia, Symphyogyjia, Pla- 
giochila, Madotheca, Lepidozia, 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: HymenophytuDi, Acrobolbus, Tylunanthus, Jainesoniella, Saccogyna, Lepicolea 
and Marsupidiuni, to mention only genera also found in Juan Fernandez. Lepi- 
dolaeiia 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. — 61 sp. 



288 C. SKOTTSBERG 

a. luidemic (17): Riccardia adghitinata and leptostachya, Metzgeria multifor- 
mis, TylunanthussilvaticLis. bilobatiis and densiretis, Lophocolea papulosa, angulata 
and submuricata, Trichocolea opposita, Schistochila Skottsbergii, Balantiopsis hians 
and lancifolia, Lopholejcunca spinosa, Stre{)silejeunea squarrosula and macroloba, 
Cololejeiinea Skottsbergii. 

/;. 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, 
Lo[)hocolea rotundifolia, fernandeziensis, chilensis, attenuata, textilis and divergenti- 
ciliata, Chiloscyphus integrifolius and lobatus, Saccogyna squarristipula, Adelanthus 
sphalerus, Lepidozia bicuspidata, pseudozoopsis, fernandeziensis and Jacquemontii, 
Trichocolea verticillata, Schistochila berteroana and pachyla, Balantiopsis cancellata, 
chilensis and purpurata, Radula hastata, microloba, Mittenii and Dusenii, Madotheca 
chilensis and snbsquarrosa, FruUania Eckloni, chilensis, lobulata and stipatiloba, 
Hrachiolejeunea spruceana, Strepsilejeunea acuminata. 

II. Neotropical element. — 13 sp. (10.5 %). 

a. I'jidemic (3): Fossombronia fernandeziensis, Anthoceros Skottsbergii, Le- 
jeunea reticulata. 

Ik 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, 
Col lira bulbosa. 

III. Chilean element. — 27 sp. (21.8 %). 

Chilean species without tropical connection, nor suggesting Antarctic rela- 
tions. 

a. luulemic (5): Solenostoma obtusiflorum and rostratum, Plagiochila fusco- 
brunnea, Lejiidozia fragilhma and disticha. 

d. Also in Ciiile, especially in the south (18): Solenostoma crassulum, Ana- 
stre[)ta bifida, Plagiochila gayana, fasciata, hyadesiana, deformifolia, chiloensis, 
rectangulata, remotidens, pudetensis, homomalla, neesiana, riparia, squarrosa, 
robusta, data and Xotarisii, I Icrberta runcinata. 

c. Cosmop(j]itan (4): Plagiochasma rupestre, Reboulia hemisphaerica, Lunularia 
cruciata, Marciiantia p()l\-morj)ha. 

ICven if not the Juan r'ernandez Hepatics are almost entirely Antarctic as 
Hkr/jx; thinks, the Antarcto-tcrtiary element is proportionally larger than in any 
other grouj), and it is not imj)ossible that genera like Megaceros or Harpalejeunea, 
the island species of wiiich were referred to the neotropical element, are Antarctic, 
and that this is true also of JUizzaiiia, represented in I and II, but on the other 
hand we must admit that pantropical genera like Riccardia, Anthoceros, Mega- 
ceros, Hazzaiiia, Lepidozia, Radula, Madotheca, FruUania 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: ^/^^i-Zr^'//^, 
Adelanthus and Herberta, for instance. 

The discontinuous distribution of many southern genera and several species 
was commented upon by Miss Fulford (/oj. 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 [rm^ 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 

Angiosperms 42.2 36.7 15.6 

Pteridophytes 60.4 37.7 1.9 

Mosses 55.8 41.7 2.5 

Hepatics 67.7 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 in a 
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 



C. 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 diflerentiation 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 I^oreal-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 tenifierate flora. The exceptions from this rule are few; for instance, gen- 
era like Pachypliialc, Lemmopsis and Massalongia may be classified as northern, 
while I\vssocaulo}i, PscudocypJiellarici, Xephronia and Stercocaiilon 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 C'hilean-Juan I'Y'rnandez 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 
.\ew Zealand is no {)roof 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 v.ith their disjunct areas were also excluded: Bacidia 
delapscDis, I sjica dasypogoides and suhtorulosa, Caloplaca rubina, Buelliti htilophiloides 
'dndfcr)i(nidirjia?ia. 

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 iinpossible to decide if they have reached their 
austral-bicentric stations (south Chile, New Zealand) independently from the north, 
or it Antarctic routes arc 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. 

b. 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 (9): Arthonia subnebulosa and berberina, Enterostigma 
Skottsbergii, Coenogonium velutinum, Lecidea avium, Pertusaria hadrocarpa and 
Skottsbergii, Lecanora masafuerensis, Caloplaca orthoclada. 

b. Non-endemic (45): Arthopyrenia cinchonae, adnexa and planorbis, Pyrenula 
aspistea, mammillana and Kunthii^, Pyrenastrum chilense, Arthonia complanata, 
Graphis intricata and Dumastii, Dirinalimitata, Schismatommaaccedens, 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. 

b. 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 <^eographicum and obscuratum, Cladonia bacillaris, coccifera, furcate, 
gracilis, pyxidata, rtmbriata and pityrea, Pertusaria leioplaca, Lecanora coarctata, 
atra, polytropa and clirysoleuca, Placopsis gelida, Candelariella vitellina, Parmelia 
laevigata, revoluta, cetrata, saxatilis, conspersa, perlata, cetrarioides and caperata, 
Ramalina linearis and iisnea, 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 Nasna rufa. 

Aves. 

Lr)\Ni!KR(; j-;4 ; names in brackets used by Goodai.l-Johnson-Philippi (//o). 

Indigenous land-birds. 

Tunius ifKii^f/iiuiicus 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", L()XXP,KRG wrote I.e. 3, a colouring characteristic of T.falklmtdi- 
cus (Ouoy et (jain), comp. 5/. V. pi. XIII. In no this is called T. f. falklandii diV\d 
the island form is referred to inagellanicus. 

A}iacretiS fevjiajideziaiius (Phil.) (Spizitornis f.). Pandemic on Masatierra. Anae- 
retcs is a neotropical genus of 7 species (Ecuador to N. Argentina and Chile). 
A. fer?iirndc.zi(Uiiis is related to a Chilean species. 

Aphyasiura Diasafucrae Phil, et Landb. Endemic on Masafuera. A second 
species ranges from Centr. Chile to Patagonia and Fuegia. 

Ciuclodfs oustalcti Scott ssp. bacckstroe^nii Lonnb. Both islands, endemic but 
very near the typical species (Antofagasta-Chiloe). A genus of 13 species (Ecuad. 
to Argent., Patag., I'alkl. Is.). 

liiislcpluvnis fiyjuvidcusis (King as 'J'rochiliis) Gould (Thaumaste f.). Pandemic, 
forming an endemic genus according to no. 300: "Coloracion totalmente diferente 
de la de cualcjuier I'icaflor cjue habita el continente." These authors leave open 
the (juestion whether or not /:'. Icyboldi Gould of Masafuera is distinct but list it 
as 111. f. Icyboldi and the form from Masatierra as lit. f.fernandoisis. L()NNBERG 
p. 7 gave gofxl reasons for considering them as identical. The humming-bird 
seems to be extinct on Masafuera. 

l'Mstcpha)ius gaUritus Mol. (as Trochilus; Orthorhyncha sephanoides Lesson 
et Garnot i<S27 — but Molina's name must be about 50 years older; Sephanoides 
s., Hi)). Masatierra and on the mainland from Centr. Chile to P^uegia. Said to 
migrate to the coast and spend the winter there (/7^), 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; suinda ranges over South America from Venezuela to 
Fuegia. 

Cercneis 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 erythronotus 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 Chiloe. 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 exterfia 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 Gigl. et Salvin. Endemic to Santa 
Clara, Masatierra and Desventuradas. Typical cooki ow New Zealand, P. cooki orien- 
talis Murphy on the coast of Peru 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. leucoptera 
"in spite of the zoogeographical difficulties for such a theory" (p. 15); LoNNBERG's 
opinion is strengthened by the fact that, whereas defilippiana and leucoptera 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 i8th century and now common, d^nd Lop hortyx calif ornicus ShdiW et 
Nodd^ introduced 191 2 or 191 3. Several landbirds have been observed as acci- 
dental visitors, Crymophilus ftdicarius and Buteo obsoletus migrants from the north 
2,wd Belonopterus chilensis, Cathartes sp., Cygnus melanocoryphus, Haematopus ater 
and Circus maculosus from the opposite coast. Petrels, albatrosses, Cape pigeons 



294 



C. SKOTTSBKRG 



and penguins arc occasionally seen around the islands but do not breed 
there. 

Of the i; s{)ecies l)reedin<^ on the islands 3 are endemic; of the remaining 
12, 6 are re|)resented b\- endemic subs|)ecies 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 noteworth}-, as also the fact that the affinities of the landbirds are all with 
S. America. 

With regard to the actual distribution we can distinguish the following two 
groups. 

I. South American (especially Chilean) group. — 10 sj). 

a. luulcmic (6): Anaeretes fernandczianus, Aphrastura masafuerae, Euste- 
phanus fernandcnsis, Cinclodes oustaleti baeckstroemii, Cercneis sparverius fer- 
nandcnsis, lUitco erythronotus exsul. 

/>. Not endemic (4): Turdus magellanicus, luistephanus galeritus, Asio flam- 
meus, Pufhnus creatopus. 

II. South Pacific group. — 5 sp. 

a. l^ndemic (3): Pterodroma externa externa, cooki defilippiana and cooki 
masafuerae. 

i). Xot endemic (2): Fregetta grallaria, Pterodroma neglecta. 

The fust grouj) includes of more tropical birds jhiaeretcs, Cinclodes and Eu- 
stiphaiius, and of more temperate Turdus, Asio, CevcJuieis, Buteo and Puffiuus. 
Of the endemic species luistepliauus fernandcnsis is the most notable, in certain 
characters a unicpie t}'pe in the family Trochilidae. The second group is of par- 
ticular interest as including, beside the widespread frigate-bird, four species of 
Plcrodroina not breeding on the mainland, where, perhaps, a special race of 
/'. cooki breeds. The genus is essentially austral-circumpolar, as it were tricentric, 
w ith Tristan da Ounha representing the African sector. Cases like those of /^. neglecta 
and externa call for a common source and suggest that Pterodroma belongs to 
an Antarcto tertiary element which inhabited the coasts and islands of Antarctica 
n prjglacia! timjs. 

Oligochaeta. 

Mich Al.l.sKx (/<S7) regards all the earth-worms of Juan Fernandez as adven- 
titious. The single strictly South American Kerria saltensis was, he believes, in- 
troduced from Chile with the human traffic, and this is also true of the three species 
of Allolobof^hora, introduced to Chile from luirope, and by Friedericia galha. The 
occurrence ot I\uhydrihis rcrrucosus offers more interest. It was known from Great 
Ikitain, the Hebrides, S.W. Africa and h'uegia, 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 19 17 in the highland of Masafuera was described 
as a new species of the Australian genus Pliilaenion, Ph. skottsbergi L. Joh, [147). 



DERIVATION OF THE FLORA AND FAUNA 295 

Whether or not the two species described from Samoa and Madagascar, respec- 
tively, belong to PJiilaeDion remains to be settled. JOHANSSON expressed some 
doubts with regard to the position of the Juan Fernandez leech and after his death 
the question was taken up by NvBELlN who showed that it should form a separate 
genus, Nesophilaemon [i88). 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 Dicksonia forest, where Pterodroma cooki masafuerae makes 
its burrows, the only possible host existing here (see also 14.'/. 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 (5^). 

OrcJiestia chileJisis (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, Ligia litiginosa and Philoscia minjica, 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 Araneus 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 Keyserl. Chile. — Mf. 

+ Lephthyphantes Fernandezi 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. 

+ Leptorhopirum (.^) Platei F. Cambr. — Mt. Doubtfully referred to this Eu- 
ropean genus, from which it differs in certain characters: "I'epigyne est d'un type 
tout a fait different" (22.430). 

+ Tmeticus Defoei F. 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 

Mtta uigrohumeraiis F. Cambr.— Mt, endemic? (see I.e. 430). The genus is 
known from all continents. 

^Selkirkiclla alhoiruttata Berl. — Mt. The genus is related to the neotropical 
G)iolus, known from Perii, l^razil and Chile, and appears not to be restricted to 
Juan Fernandez; Hi:rl.\M) has seen a very closely related species from Valdivia 
[22.A12). 

Miivs)naucheuius scg)>untatus Simon. Patagonia and Fuegia. — Mt. A genus 
of 2 species, the second one from the Magellanian region. 

^ Misumoiops Sjoestedti Berl. — Mt. An American, especially X. American 
genus. 

+ (uiycjma Skottsbergi l^erl. — Mt. A S. American, especially Chilean genus 
of numerous species. 

(',. niaculaiipes Keyserl. Chile. — Mt. 

rOxysoma Delfini Simon. — Mt. A S. American genus. 

^ Plulisca or)iata Berl. — Mt. A subantarctic-magellanian genus extending north 
into Chile. 

^/V/. iugeus Herl.— Mt. 

■r Lycosa I'CDiajidezi F. Cambr. — Mt. The genus is cosmopolitan. 

lii'oplirys quilpiioisis Simon. Centr. Chile. — Mt. The genus is known from 
Centr. and S. America, ICuroj^e, 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 /.^//^///^//r//;;/ nothing can be said. The 
{)resence of a southern, eventually Antarctic element is indicated by MecysinaucJic- 
)iiiis, possibly also by riiilisca, but so far there is no sign of a bicentric group. 
\\ hethcr it can be distinguished in subantarctic America I cannot tell. 

Acarina. 

TR.'u;.\Kl)n [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 onl}' a small part of the acarofauna; this is evident 
already from the fact that not a single sj^ecies came from Masafuera or Santa Clara. 
Whether the cosmopolitan sj)ecies 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 Euiergvs 
similis Trag. belongs to a genus hitherto recorded only from New Zealand and 
that Phyllhermannia deiitata Trag. is related to a neozelandic species; the genus 
is also found elsewhere. 

Pseudoscorpionidea (jp). 

+ + Asterocherftes vittatus Beier. — Mt. The genus has its greatest resemblance 
to Thalassochernes Beier from New Zealand. 
+ Chelanops insularis Beier. — Mt. 

+ Ch. ktischeli Beier. — ^Mt. Related to a Chilean species. 
+ Geogarypus bucculentus Beier. — Mt. 
'V Parachernes kuscheli Beier. — Mt, Mf. 
+ + Protoivirthius fernandezianus 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 (I.e. 205). 

Myriapoda. 

The very small and incomplete collection — no specimens were brought from 
Masafuera or Santa Clara — was studied by Verhoeff {2^4). In order to get some 
information on the distribution of the genera I asked Dr. Otto Sciiubart 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 pusilhis Leach (Microbrachyiulus litoralis Verh.). Indigenous in 
western Europe, adventitious in N. America and Argentina. 

Brachydesmus supertis Latzel. A European species, adventitious in N. America 
and Argentina. 

+ Aulacode sinus 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 Be, Austral., 
Tasm., N. Zeal., N. Caled., Hawaii). 

Nopoiulus veiiustus Meinert (pulchellus Leach). Widely distributed in Europe, 
introduced to N. America and Chile. 

Cylindroiulus frisius oceanicus Verh. Typical frishis {C. O^veni Bollman) 
introduced to N. America, Argentina (also in forma oceanicus), S. Africa and St. 
Paul's I. 



298 C. SKOTTSBERG 

Chilopoda. 

+ Xesoj^fop/ii/iis laticpllis (Attems) Schubart (Geophilus, Verb.). Endemic. The 
^einis, wbicb has not been reported from S. America, includes after the latest revi- 
sion by Airr.MS (as subgenus of Geopliilus) i i species (i SAV. Austral., i N. Zeal, 
I X. Caled.. i An nam, 3 Jap. and 2 Eur.). 

r Xi'S(>o-i-()p/iiii(s biU-ckstrocini (Verb.) Schubart (Geophilus, Verb.). Endemic. 

Schizotoiia alaccr (Pocock) Silvestri. Chile, south to Fuegia, Argentina. A 
genus of 6 s[)ecies (3 Chile and Argent, to Patag., Eueg., i E. Austral., 1 N. Zeal., 
I Chatham Is.). 

Litliobio)>wyplia afyicaua Porat (Lamyctes insignis Pocock, insignisbaeckstroemi 
X'erh.). 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., \\. Ind., Austral., Tasm., N. Zeal., Chatham Is., N. 
Caled., Kermadec Is., Guam, Hawaii]. 

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 Eernandez. 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. 

CoUembola. 

Of the 8 species distinguished by SciloTT [216), the first ever collected in 
Juan PY'rnandez. 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 wdde range is due 
to the great age of CoUembola or a result of later dispersal is unknown. 

Thysanura [222, 2pj). 

 Is(>U'pis))ia luiucctois Silvestri. — Mt, Mf. The specific epithet refers to the 
intermediate position between IsolcpisDia and ll€tcrolcpis7>ia\ the species is com- 
pared with forms known from Africa and Australia. 

' h KuscJielocliilis Ocliai:;aviac W'ygodz. — Mt. A monotypical endemic genus 
related to Alloinacliilis and XesoniacJiilis from Australia, but not, as far as known, 
to an American genus. 

Among the Invertel)rates 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 
obhged to Dr. Olof Ahlberg, Stockholm (Thysanoptera), Dr. KjELL Ander, 
Linkoping (Orthoptera), Dr. Per Brinck, Lund (Coleoptera), Mr. Nils Bruce, 
Gardby (Coleoptera), Dr. Lars Brundin, Stockholm (Coleoptera), Mr. Felix Bryk, 
Stockholm (Lepidoptera), Dr. W. E. China, London (Hemiptera), Dr. K.-H. FoRSS- 
LUND, Stockholm (Trichoptera), Dr. G. J. Kerrich, London (Hymenoptera), Dr. 
K. Princis, Lund (Orthoptera), Mr. Bo Tjeder, Falun (Neuroptera) and Dr. 
B. P. UVAR(3V, London (Orthoptera). 

JOHOW (ijo) 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 19 16-17 
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 195 i 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 (^.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. Kusciiel's large material has been worked up 
by specialists, and I can only refer to what has been published (142, 208, 2g2, 
2gj, jog, 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 [22j). 

+ Euborellia annulipes (Lucas). — Mt, SC, Mf. The genus S. Amer., E. Afr., 
Orient, Ind., Ceylon, Tasm. 

Anisolabis Bormansi Scudd. Galapagos Is., Easter I.— Mt. A large genus of 
world-wide distribution. 

Saltatoria [j8, 22j). 

^Hoplospyriuin Skottsbergi Chopard.— Mt. An American genus, the species 
related to species from N. America and Chile. 

Trimeroiropis ochraceipeimis Blanch. Chile.— Mt. The genus is American. 



300 



C. SKOTTSBERG 



Corrodentia. 

Isoptera. 

- Kiiloti-nfics i^^racilii^^Jicitus luiierson. — Mt. The only Termite known from Juan 
Fernandez. "The wing venation is close to that of Kalotermes broiini Froggatt 
from New Zealand" [8j. 393). 

Mallophaga [266). 

Puffimis creatopus and Ptcrodroma neglecta and exterua are infested with the 
same mallophagous parasites found on these and related birds in other regions. A 
new Halipcurus is mentioned but not described. 

Thysanoptera (7). 

Aeolothrips fasciatus L. Boreal. — Mt. 

-X- Physotlirips Skottsbcrgi Ahlb. — Mt. The genus is distributed over N. Amer- 
ica, luirope, W. Asia and Australia. 

-SencotJirips vieptiis Ahlb. — Mt. Perhaps nearest to a Californian species. 
The genus is otherwise confined to F^urope, where it is widely spread. 

rinips tabaci Lindem. X. America. — Mt. The genus known from N. America, 
luirasia, \. Africa and Australia. 

The two non-endemic species may have been introduced accidentally. Both 
were found in the spathe of Zantcdcschja acthiopica, cultivated and naturalized. 

Neuroptera (zcVf, g2, I2j). 

+ + Couclioptcrella kusclieli Handsch. — Mt. 

+ 6'. Diacidata Handsch. — Mt. 

(layoinyia falcata (Blanch.). Chile, Argentina. ^ — Mt, Mf. A small S. American 
genus. 

' I Ic))U'yobiiis Siocstedti Xavas (M. fumosus l^>sb. -Peters., H. nigrinus F"sb.- 
Peters.). — Mt. An almost world-wide genus, absent from the S. hemisphere except 
for the Andean region (Colomb., Perii, l^oliv., Ikaz., Argent., Chile). 

-i //. Skottsbn-gi Xav.-is.— Mt, Mf. 

Trichoptera [21^. 

+ Australoinyia inastUierra Schmid. Mt. The genus is known from Chile^ 
Patagonia and I^'alkland Is. 

■\-A. luasafuera .Schmid. — Mf. 

I'ergcr Por/iri Xav. Centr. Ciiile. — Mt. A Chilean genus. 

Lepidoptera (tJ). 

A great number of genera and species will have to be added when Dr. 
Klsciikl's material has been determined. 



DERIVATION OF THE FLORA AND FAUNA 3OI 

Tineomorpha. 

Gelechidae. 

+ + Apotheioeca synaphrista Meyr. — Mt. The genus is closely allied to the large 
and widespread Gelechia (N. and S. Amer., Galap., 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 Schifif. — Mf. Widely spread, domestic. 
Tineidae. 

Monopis crocicapitella 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. 

+ "^ Fernandocrambiis Baeckstj'oemi Kwx'w . — Mt, Mf. The genus nearly related 
to Crambus. 

+ F. brunneus Auriv. — Mt. 

+ F. fuscus Auriv. — Mt. 

+ +yuania anmdata Auriv. — Mt. Similar in some ways to Ptochosiola Meyr. 
(S. Afr., Australia). 

Nomophila 7iociuella Schiff. Cosmopol. — Mt, Mf; adventitious. 

■\-Pionea fiunipefinis (Warren) Hamps. — Mt. A world-wide genus. 

Scoparia Ragonoti Butl. — Mt, Mf. Chile. A very widespread genus. 

Geometfina. 

Geometridae. 

^ Eupithecia halosydne Prout. — Mt. A widespread genus, but not found in 
Australasia. 

+ E. (.?) znepta Prout.— Mt. 

-\-E. physocleora Prout. — Mt. 

■\-Lobophora insulai'is Auriv. — Mt. An essentially Palaearctic genus. 
Tortricidae. 

+ Crocidosema insulana Auriv. — Mt. A S. American genus. 

■vEulia griseiceps Auriv. — Mt. The genus Holarctic, also in Hawaii; few else- 
where. 

■\-E. Robinsoni 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 Guen. Amer. (south to Chile); Macaronesia. — Mt. 



30. 



C. SKOTTSBERG 



-V ~ Hoplotaysia inai^iia Auriv. — Mt. Related to Copitarsia. 

Li'ucauia inipuiicta (iucii. Chile.— ^It. A bipolar genus (Palaearct., N. Zeal.). 

-^ Lvcopltotia luuiks/roojii Auriv. — Mt. The genus widespread (Arct., Amer., 
luir., S. Afr., Madag., X. Zeal.). 

L. DitssiiDH Gucn. Chile, Patag. — Mt. 

Kac/nplusiii uu (iucn. I'atag.. Urug., Argent., Chile.— Mt. 

Sv>ii:-yapha i^-auiDioidts Hlanch. Chile. — Mt, Mf. A Palaearctic genus; Mex., 
S. Anicr. 

Rhopalocera. 

Pvrtv/ii/s cnrvi- Iluebn. Widespread in S. America and probably introduced 
in Juan bY'rnande/.. — Mt. 

Diptera [84, 142). 

Acroctyidac [208^. 

-V Opcodes knsclicli Sabr. — Mt. A temperate genus, recorded from all con- 
tinents, but only 2 species known from the mainland of S. America. 
AiitlioDiyzidae [12S]. 

Anthony zii cursor (Kieffer). Cosmopol., also S. Chile. — Mt. 
Calliplioridac {2-;-;) . 

L 'ally}itropyga huuteralis (Walker) Souza L. et Alb. (C. Selkirk! Enderl), Chile 
C<)ncej)ci6n). — Mt. Ml". 

raraludlia fuh'icrura (Desvoidy) Aub. et Baxt.— Mt, SC, Mf. 

Sarcojicsia cliloroi^astcr (Wiedem.) Arrib. Chile, I{^aster I. — SC, Mf. 

Sarc())ies}onii)na hicolor Souza L. et Alb. Chile (Santiago). — Mt, SC. A mono- 
t\-pical genus. 

Cecidoniyidae [84, 106). 

+ ^- Tsadaria pallida iMulerl. — Mt, Mf. Related to Canipyloniyza Meigen. 
(. hloropuiac [2()g). 

Ilippilates ausiralis Sabrosky [//. (Cadrema) nietallicus P3nderl. non Beck. 
[^ II. flai'ipcs (Loew) Sabr.)]. I^xuad., Peru, Argent., Chile. — Mt, probably ad- 
ventitious. 

L uliiidae. 

Ciilcx intcrfor Dyar. — Mt. Mf. 
J) (die hop odidac [126). 

^ Ilyilrophorus hu.ululi llarmston. — Mt. 

//. polioi^asUr (I'hil.) llarmston. Chile. — Mt, SC. 
rSy)npyc?ins fdiiandtrjrnsis llarmston. Near a Chilean sp. — Mt, Mf. 

liphydridac [28(/). 

Diniciocnia cars/a Iv.d. Wulp) Wirth. Argent., Uruguay.— Mt, SC. 
-V Discoceriiia fu))iipc)nus Wirth. — Mt. Near a Chilean sp. 
Ilyadina certa Crcsson. Chile. — Mt, Mf. 
Ilydrellia vulgaris Cresson. (iuatem., l^oliv., Chile. — Mt. 



DERIVATION OF THE FLORA AND FAUNA 303 

+ Scaiella a7igustipennis VVirth. — Mf. An almost Cosmopolitan genus. 

+ 5. argeniifacies Wirth. — Mt. 

+ S. brachyptera Wirth. — Mt. 

+ 5. decemguitata Wirth. — Mt, SC, Mf. 

^S. discalis Wirth. — Mt. 

+ 5. fernandezensis Wirth. — Mt. 

+ 5. kuscheli Wirth.— Mt, Mf. 

+ S. lutea Wirth.— Mt. 

■vS. niarginalis Wirth. — Mt. 

+ 5. niasatierrensis Wirth. — Mt. 

■vS. minima Wirth. — Mt. 

+ 5. 7ia7iopiera Wirth.— Mt. 

+ 5. pallida Wirth.— Mt. 

+ S. pilimana Wirth.— Mf. 

+ S. stenoptera Wirth. — Mt. 

+ S. vittata Wirth. — Mt. 

+ Scatophila fei'naftdesiana Wirth. — SC. 

vS. medifemur Wirth. Chile (Coquimbo). — Mt, SC. 

Heleidae [288). 

+ Dasyhelea australis Wirth. — Mt, Mf. Near a Chilean sp., genus cosmop. 

+ Forcipo7Jiyia te7ttiisqica77iipes Wirth. — Mt. A widespread genus (N. and S. 
Amer., Eur., Afr., Austral.); one species common to Paraguay and Australia. 

+ F. sa7ictaeclarae Wirth. — Mt, SC. 

Helo77iyzidae (128). 

Blaesochaetophora picticoimis (Bigot) Henn. S. Chile. — Mt. 

Prosopa7ttru7n flavifro7is Tonn. et Mall. (Cnemospathis Baeckstroemi et 
Schoenemanni Enderl.). Chile, S. Africa, New Zealand. — Mt, Mf. 

Lo7ichaeidae (128). 

Lo7ichaea patago7iica Malloch. Chile. — Mt. 
Mnscidae (128). 

Austrocoe7tosia ig7wbilis (Stein) Hennig. Chile. — Mt. 

Ci'aspedockaeta Ii77ibi7iervis (Macq.) Hennig. S. Chile. — Mt, Mf. 

Delia platiira v. sa7icti-jacobi (Bigot) Hennig. Chile. — Mt, Mf. 

Euryo77i7na peregri7iU77t (Meigen) Hennig. Peru, Chile. — Mf. 

Fa7mia a7tthraci7ia (Walker) Hennig. Chile. — Mt. 

F. ca7ialicularis (L.) Cosmopol., also in Chile. — Mt, Mf. 

F. pu7ictive7ttris Malloch. S. Chile. — Mt. 

Fucellia i7tte7n7iedia Lundbeck (Egeria masatierrana et masafuerana Enderl.), 
Eur., Oceania. — Mt, Mf. 

Hydrotaea cya7ieive7itris Macq. Chile. — Mt, Mf. 

Li77i7iophora patag07tica Malloch. S. Chile, Patag. — Mt. 

■\- NotoschoeTiomyza kuscheli Hennig. — Mt, Mf. 

Ophyra caerulea Macq. Centr. Chile to Fueg. — Mt. 



304 C. SKOTTSBERG 

-\-ScliO€)W})iyzJ}ia eiudejii liennig. — Mf. 

+ Sylliifniopliora lispofi'nfia Hennig. — Mt, Mf. 

MycetophiUdac [100]. 

-. lixicJiia furcilla h'reeni. — Mf. Near a Chilean species; world-wide genus. 
- Leia tiiallcolns Freem. — Mt. Allied to a species reported from Bolivia, 
Peru and Brazil; the genus world-wide. 

Macrocera fujicrca Freem. Chile.^ — Mt. A world-wide genus. 

-r Mvcetophila a)iij;ustifuyca P^ndcrl. — Mt, Mf. The genus world-wide. 

M. co)iiftra Freem. Chile. — Mf. 

J/, corjiuta Freem. Chile. — Mt. 

iM. fliU'oluuata hVeem. Chile.— Mt, Mf. 

M. (.') iusccta iM-eem. Chile.— Mt. 

M. spiiiosa Freem. Chile. — Mf. 

+ J/. subfiiDWsa Freem. — Mt. 

k- Paraleia iiephrodops (r^nderl. s.n. Selkirkius) Freem. — Mt. 

/'. Jiubilipcuuis Walker. Chile. — Mf. The genus neotrop., Austral., Tasm. 

SciopJiila ocJircata Phil. Chile. — Mt. A w^orld-wide genus. 

Piophilidae [12S). 

Piophjla case? (L.). Cosmopol., domestic. — Mt, Mf. 
P. foi'colata Meigen. Cosmopol. — Mt. 

Phoridae [S4). 

-f Lioyella juajifcDiajidczica Enderl. — Mt, Mf. A F^uropean genus. 
P/iryjuidae (i2cS'). 

Plirync fuscipcuuis Macq. S. Chile.— Mt, Mf. 

l^latypizidac. 

Microsaiiia pallipes Meigen. Cosmopol. — Mf. 
Psycliodidat- (cS'y, 212). 

PsycJioda ci}ierea l^anks. Cosmopol., also Chile. — Mt. A widespread genus. 
\ l\ t>nuat}eyyc7isis Satchell. — Mt, Mf. Possibly = the following. 
vP. ))U)nitissima ICnderl. — Mt. 
/'. sci>tyi)ii Tonnoir. Widespread in temperate regions. — Mt, Mf. 

Say cop hagidac ( i* -f -f ) . 

Jfypopy^Q^ia nay'ia (Walker) Townsend. Chile.— Mt, SC, Mf. 
Scatopsidac [S^, roo). 

■^ ^ Masatieyya fcyyugbiea luiderl. — Mt. Related to the European Rhaeboza 
linderl. I^'kkemax does not mention Masaiicyya. 

Scatopse fuscipes Meigen. — Mt, Mf. A world-wide genus. 

S. iioiata (L.). Cosmopol., introduced with the traffic. — Mt. 

Sciayidae [ioo\ 

+ Brcidysia fnsca I^Veem. — Mt, Mf. A world-wide genus. 



DERIVATION OF THE FLORA AND FAUNA 305 

+ B. media Freem, — Mt, Mf. 

+ Merianina kusckeli Freem. — Mt, Mf. Another species in Brazil. 

+ Psiiosciara nitens Freem. — Mt. 

Sunuliidae (2g4). 

+ Gigantodax kusckeli 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 Leptocera by RICHARDS who, 
however, does not list the two Gyretria species described by Enderlein, but 
not found in Dr. KuscilEL'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. 

+ Z. cultellipennis (Enderl. ut Skottsbergia) Richards. — Mt. 

L. darivini Richards. Chile, Argent. — Mt, Mf. 

L. divergeiis Duda. Peru, Boliv., Chile, Argent. — Mt, SC. 

+ Z. duplicata Richards. — Mt. 

+ Z. ellipsipemiis Richards. — Mt. 

L. flavipes (Meigen) Richards. Eur., N. Afr.— Mt. 

L. mediospinosa Duda. Cosmopol. — Mt. 

L. pectinifera (Villen.) Richards. Eur., Falkl. Is.— Mt. 

L. pulchripes Duda. Argent., Parag., Urug. — Mt. 

+ +Phthitia alexandri Richards. — Mt. The genus must be very u^diV Leptocera. 

+ Pk. selkirki (Enderl. ut Pterodrepana) Richards. — Mt. 

+ /%. 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. 

+ J/. Lundbladi (Enderl. ut Carposcalis) Fluke. — Mt. 

Eristalis tenax (L.). Cosmopol, also in Chile. — Mt, Mf. 

Sterphus aurifrons Shannon. Chile. — Mt. 

Tachinidae (70). 

Lrtcamyia chilensis Aldrich. Chile. — Mt, SC, Mf. A S. American genus. 
+ Pkanfasiosiphona kusckeli Cortes. — Mt. A Centr. American genus. 

Tendipedidae (288). 

Anatopynia vittigera Edw. S. Chile, Patag. — Mf. A world-wide genus. 

-^Clunio fuscipenitis 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 



3o6 C. SKOTTSBKRG 

//. pnitontni (Goett^eb.) Coe. luigland. Chile, Patag. — Mt. 
//. pusilliis (luiton) Coc. luiglaiul, Kerguelen. — Mt. 

-^ l\)tio>io)uus acutus W'irth.— Mf. A genus of numerous species in southern 
S. America, few elsewhere (luir., \. Amer.). 

rP. t/isiish'/us W'irth.— Mt. Mf. 
P. kirfi-ri ((iarrett) lulwards. l^rit. Columb., Chile, Eur.— Mf. 

rP. kuscheli Wirtii.— Mf. 
/'. )iii^rinNs lulwartls. — S. Chile. — Mf. 

+ /'. Si'lkirki W'irth.— Mt. Mf. 

'Pit?iytars/(S ffar/pis (Meigen) Townes. \. Amer., S. Amer., also Chile, 
luir.— Mf. 

'J'i/'iilii/iu- (7). 

l-lrioptcra pilipcs (Fabricius). Cosmoj)ol.— Mt, SC, Mf. 

+ Lii/io?iiij (I)icranomyia) affabilis Alex. — Mt, Mf. A very large and wide- 
ranging genus. 

■tL. a))ipliio)iis Alex. — Mt. 

-f A. axierasta Alex.— Mt. 

+ A. Jiarpax Alex. — Mt. 

+ A. kuscluliaiia Alex. — Mt. 

-tL. iiiasafuerae Alex. — Mf. 

+ A. pidcsti'is Alex.— Mf. 

+ A. selhiiki Alex.— :\lt. Mf. 

+ A. siuardoi Alex. — Mt. 

L. tyilubeyculata Alex. S. Chile, Patag. — Mt, SC (an endemic variety). 

+ A. I'oiatrix Alex. — Mt. 

+ A. luucris Alex.— Mf. 

+ A. yioiqucinia Alex. — Mt. 

-V MolophilHs ainpliacajitlius Alex. — Mt. A world-wide genus well developed 
on the mainland. 

+ J/. ivitiinonis Alex. — Mt, Mf. 

+ J/. apprcssospiiius Alex. — Mt. 

+ J/. arcifirus Alex. — -Mf. 

■V M . civiopus Alex. — Mt. 

+ J/. iiffocajiiis /\lcx. — Mf. 

■\M. (iistifurcus Alex.- Mt. 

+ J/. filioliis Alex.— Mt. 

+ J/. films Alex.— Mt. 

+ J/. uiasafucrac Alex. — Mf. 

+ J/. )}U(ltifi(ius Alex. — Mt. 

+ J/. fNpfiuii/s Alex. — Mt. 

+ J/. pcclijiij'crus Alex. — Mt. 

+ J/. rcctispiiius Alex. — Mt. 

-f J/, sclkirkiainis (Knderl. ut Archimolophilus) Alex.— Mt. . 

-fJ/. .sVj'.r Alex.— Mf. 



DERIVATION OF THE FLORA AND FAUNA , 307 

+ M. tridens Alex.— Mt. 
■vM. variants Alex.— Mf. 
-{■M. yunqiiensis Alex. — Mt. 

+ Skan?to?toinyia kuscheli Alex. — Mf. An American genus, well represented 
also in Chile and extending north to Canada. 
■\-Sh. masatierrae Alex. — Mt. 
■\-Sh. selkirkiana Alex. — Mt. 
+ Tipula baecksU'oemi Alex. — Mt. The genus world-wide. 

Coleoptera. 

As yet little has been published about the beetles collected by Dr. KUSCIIEL; 
to judge from what is known the number of island species no doubt will be 
multiplied. 

Anobiidae [igs\ 

Anobiuni punctatMin De Geer. Cosmopol, introduced. — Mt. Beside the typical 
species an endemic ssp. described by PiC (Mt). 

A. striatum 01. Cosmopol., introduced. — Mt. 

+ Calymmaderus atronotatus Pic. — Mt, Mf. Near a Chilean species. Numerous 
species in N. and S. America. 

+ ^ Masatierrum inipressipeime 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. pubescois Md,x. piceitarsis Pic (fumosus var., Pic). — Mt. The variety endemic. 

A nth ribiidae [133] . 

+ + Opisolia lenis Jordan. — Mt. Related to Eucyclotropis Jordan (Centr. and 
S. Amer.). 

Bostrychidae [166). 

Neoterius pulvinatus Blanch. Chile. — Mt. A small genus reported from Peru 
and Chile. 

Prostepha7ius sulcicollis Fairm. et Germ. Chile. — Mt. 

Carabidae (5, 260). 

Bembidimn inconstaris Solier. Chile. — Mt. A world-wide genus. 

B. 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. kuscheli Straneo.— Mt. 

+ J/. ovalipennis Straneo. — Mf. 



jo8 C. SKOTTSBERG 

+ Plerostichus kusclicli Straneo. — Mt. A world-wide genus. 

+ /V. selkirki Aiulrcwes.— Mt. SC. 

+ /V. skottshfyi:;i Andrewes. — Mt. 

+ /V. H'alkiri Andrewes. — Mt. 

+ + Iraclnsivus hasalis Straneo. — Mt. The genus presumably endemic [260. 

■3«). 

■T '/'. bicolor Straneo. — Mt. 
+ r. emdi'iii Straneo. — Mt. 
-f /". kuscluli Straneo.— Mt. 
+ 7". (>i'ij/ipiN}iis Straneo.— Mt. 
f '/'. pallipcs (ierm.— Mt, Mf. 
+ /'. pH)ntii:;ey Andrewes. — Mf. 
+ 7. sericeus Andrewes. — Mt. 

+ Trechisihus bacckstfoe)}ii (Andr.) Straneo. — Mf. An American genus. 
7". fe»ioralis Germ. Chile. — Mt (end. ssp.), SC. 
+ 7". kusclicli Jean n el. — Mt. 
r / 'ariopalpus crusoci Keed. — Mt, SC. 

( Inysoinclidac [2S2, iS]. 

^ vMiJiotula fcffiivnicziiVia l^echyne. — Mt. The genus is related to Hyp7io- 
pJiila (W. ICur. -Japan). 

\-M. kuschcU Hechyne.— Mt. 

fj/. iiitois W'eise. — Mt. 

Cioidac [166). 

 ( is hi))iaculatiis Germ.— Mt. A world-wide genus. 
+ C fcy}ia}idczia)ins Lesne. — Mt. 
r C '. nifus (jerm. — Mt. 

i Icridac [21^;). 

Xccrohia rufipcs De (ieer. Cosmopol. — Mt, introduced. 

C Occific/Iidnc [2iS2). 

liriopis opposita (nicr. Chile. — ^It. The genus ranges from Vancouver I. to 
Patag. and I'"uegia. 

C(dydiidac [2oiA. 

^ /[ycHonicrodcs uiasd/ucroisis Poj)e. — Mf. lieside the two Juan Fernandez 
species there is a third in New Zealand. 

'r I\ ni(isa/icyrc>!S/s I'ope. — Mt. 

V Pyotoincrus iiisularis (ironvelle. — Mt. The genus N. and S. Amer., E. Ind., 
Japan, .Australia, \. Zeal, (numerous), .Samoa. 

4 7'. j^cnudiiii Pope. — Mf. 

C yyptophagidac {j;6]. 

Cyyptopliai::us atomayioidcs (iron v. (Selkirki l^ruce'). Chile.— Mt. A world- 
' Letter 19.9. 1954. 



DERIVATION OF THE FLORA AND FAUNA 309 

wide genus, the subg. Mjiionomus, 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. 

+ Loberosche?na convexum Bruce. — Mt. Beside the island species 3 in Bo- 
livia and 2 in Chile. 

+ Z. discoideum Bruce. — Mt. 

Curculionidae [12). Dr. Kuschel, who specializes in this family, estimates the 
number of species collected by him to exceed one hundred. 

+ + Arwletkrus gracilis Auriv. — Mt. 

+ -v Apteronanus dendroseridis Auriv. — Mt. 

+ A. (.^) gunnerae Auriv. — Mt. 

Arajnigus Fulleri 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 (KuscilEL in litt.). 

+ Cyphometopus masafuerae Auriv. — 'Mf. 3 species in Chile, where the island 
species most likely also occurs (Kuschel in litt.). 

■^ -^Juanobia ruficeps Auriv. — Mt. 

+ ^Juanorhinus Robinsoni Auriv. — SC. 

Otiorrhynchus rugosostriatus Goeze. W. and S. Europe. — Mt, accidentally 
introduced via Chile. 

+ -v Pachy stylus dimidiaius Wollaston. — Mt. 

+ P. nitidus Auriv. — Mt. 

+ Pachy tragus crassirostris Wollaston. — Mt. 

+ Pentarthrujn affijte Wollaston. — Mt. A widespread genus, found on many 
oceanic islands. 

+ P. nigropiceum (Phil.) Auriv. — Mt. 

+ P. nitidum Wollaston. — Mt. 

+ /^. rufoclavatum Auriv. — Mt. Close to P. ^//V^/^' Broun from New Zealand. 

+ +Platynanus arenarius Auriv. — Mt. 

+ P. Baeckstroemi Auriv. — Mt. 

+ P. hirsutissimus Auriv. — Mt. 

+ P. quadratifer Auriv. — Mt. 

+ P. sericatus Auriv. — Mt. 

+ /^ Skottsbergi Auriv.— Mt. 

+ Strongy/optei'us 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 (2^p, ii?)- 

Afiisomeria bisiriata Brulle. Chile. — Mt. The genus known from Tristan da Cunha. 



3IO C. SKOTTSBERG 

^Lancctcs Baeckstrocnii Zimmerm. — Mf. Tlie genus austral bicentric. 
RliaJitHs sigjiatiis ssp. knsclieli Guignot. — Mt, Mf. An endemic variety of 
a Chilean species. 

Latlnidiidae [ig6). 

r Co)ii)ioi)ius curtipe}uiis I'ic. — Mt. Near C. dijuidiaius Belon (Boliv., Chile). 
An essentially luiropcan genus. 

C. suhfasciatus Reitt. Chile.— Mt. 

Milasidae [joS). 

^ rscudodiiurctiis Sclk'nki Flet. — Mt. An Argentine genus. 

Mycetophaoidae [ic)6). 

Mvit'/op/iajrus c/iili?isis Phil. Chile. — Mt. The genus in Kur., Asia, Afr. and Amer. 

Xitidididae [nj6, loj). 

rC)iips acuta Ciillogly.— Mt. A Chilean genus. 

4-r. atrata (iillogly.— :\Tt. 

t L '. dii'vrsa Pic— Mt. 

+ 6". fi'Diandczia (jillogly. — ?^It. 

•C. niucrojiis (iillogl)-. — Mt. 

Scarahaiidae (J'//). 

Aphodius iiranariiis L. Cosniopol., domestic. — Mt. 

Plfuropliarus cacsus Creutz. Reported from X. Amer., Chile, Eur., Orient, 
.Madagascar. — Mt, introduced. 

Scolytidac \^^o^)). 

(iiiathotriclius tortliyloidcs Schedl. Chile. — Mt. The genus in N. and S. America. 

IVilocotribus ivillei Schedl. Peru and Chile. — Mt. 
Sti ip //] 'li)iida e ( J* -r ) . 

 Atluta Kob'nisoui l^ernhauer. — Mt. Near a Brazilian species. A cosmopoli- 
tan genus of about 2000 sj)ecies. 

 liltusis sonirufa h'airni. et (ierm. — Mt. A genus of about 1 50 species, S. 
Amer. (also Chile), .Austral., N. Zealand. 

\Mid(>)i irusdi-aiius Bcrnhauer. — Mt. A cosmoj)olitan genus of about 500 
sj)ccics. 

r Ocyusa l^acckstrocDu Bcrnhauer. — ?^It. About 40, mainly Palaearctic. 
IViilo)itlius nitidiponiis Solier. Chile. — Mt. A cosmopolitan genus of about 
(Soo species. 

I 7'rooop/ti(>fns Sko/Zs/xr^i;-/ Bernhauer. — Mt. A cosmopolitan genus of about 
350 -^P- 
I I'DuiocJiilidac ( / 9"^ ). 

+ r/ianodi-s/a crihraria (Blancli.). Includes Ph. auirulata Reitt.— Mt, SC. A 
Chilean genus. 

+ /'//. robust a Pic. — Mt. 

+ /V/. rarici^ata Germ.-Mt. 



DERIVATION OF THE FLORA AND FAUNA 3II 

Tenebrionidae [lO/f., ig6). 

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 gracilipes Phil. Chile.— Mt. Numerous species in Chile. 



Hymenoptera. 

Aphelinidae (211). 

Aphelinus jucmidus Gahan. N. America. — Mt, Mf. 

Bethylidae [igi). 

+ Cephalonomia skottsbergi Brues. — Mt. 

+ +Lepidosternopsis kuscheliana Oglobin. — Mt. 

+ Perisieroia maculicornis Oglobin. — SC. A widely distributed genus of about 
25 species. 

+ P. sanctae clarae Oglobin. — SC. 

Braconidae [i8y). 

+ Apa?iteles evadite Nixon. — Mt, Mf. A wide-spread genus. 

^■A. morroensis Nixon. — SC. 

Aphaereta minuta (Ns.). — Mt. 

+ Opms kuscheli Nixon. — Mt, Mf. 

^0. scabriventris Nixon. — Mt. 

Dryijiidae (igi). 

■^Haplogonatopns 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). 

+ +Kus chelae her ius acrasia De Santis. — Mt, Mf. 

■^ Pseudelachertus semijiavus De Santis. — Mt. The genus otherwise Aus- 
tralian. 

Encyrtidae [211). 

+ Hemencyrtus kuscheli De Santis. — Mt, Mf. A neotropical genus. 

Eniedontidae (211). 

+ Achrysochris bicarinata De Santis. — Mt, Mf. 

Eupai^acrias phytomyzae (Brethes) De Santis. Chile, Argentina. — Mt, introduced. 
+ Omphalomorphella elachertiformis De Santis. Said to come near an Aus- 
tralian sp.— Mt, Mf. 

Eulophidae (211). 

■V Diaulomyia calvaria De Santis. — Mt, Mf. Allied sp. in Australia. 



312 C. SKOTTSBERG 

Formicidae [2SJ). 

Ponera trigoua Mayr var. opacior Forel. \. Amer., W. Ind., Chile, Argent. 
— Mt. The typical sfjecies in Ikazil. Probably spread with the traffic. 

Proiolepis obscura Mayr ssp. raga Forel. Melanesia. — Mt. The typical spe- 
cies Java, Australia, Hawaii, another variety N. Guinea and Melanesia. 

Tet}a))ionu))i guiiueiise (Fabricius). An African ant, now widely spread with 
the human traffic. — Mt. 

IcJniciDnonidae {2o6\ 

Rnicospilus purgatus Say. Temperate N. and S. America. — Mt, Mf. An al- 
most world-wide genus (Amer., luir., Afr., Austral., N. Zeal., Hawaii). 

-\- llcniiteles Bacckstroe})ii Roman. — Mt. An almost world-wide genus, less 
rich in the tropics. 

+ //. luasafuerae Roman. — Mf. 

+ flo/ocn'UDia {^) juaniaua Roman. — Mt. The genus is known from Europe. 

Mettlia (Paniscus) gerliugi Schrottky. Chile. — Mt. Range of genus very 
wide, including S. Amer., Falkl. Is., Rodriguez, Austral., N. Zeal. 

Stilpiius gagatcs Grav. var. Robi)iso7ii Roman. — Mf. The typical species in 
luirope, the genus also in N. America and Greenland. 

Tyiptognathus aequiciiicius Spin. Chile. — Mt. 

MyDiaridae [igo). 

Aiiagnis incaniatus Hal. Palaearctic. — Mf, undoubtedly introduced. 

+ + CroimoDiytiiar fcrnandczi Oglobin. — Mt. 

■V C. jniperfcctus Oglobin. — ]\It. 

+ + Xcsopolynema caudatuui Oglobin. — Mt. 

P()ly}u))ia fiiscipes Hal. Palaearctic. — Mf, supposed to have been accidentally 
introduced. 

+ V ScolopsoptcroJi kiiscJieli Oglobin. — Mt. 

Rhynchota. 

Heteroptera. 

A)itli()coyidac {2 1\ 

-^ Hucliivianiclla dcria Ik-rgroth.— Mt. Related to B. continua B. White from 
Madeira; other species reported from Tasmania and Hawaii. 

Lyctocoris catupcslris Vi\h\. Cosmopolitan, probably adventitious. — Mt. 

Lygacidac {2/, j6i). 

' V Micrymcuus kuschcli Kormilev.— Mt. Most nearly related to Metagerra 
H. White from New Zealand (Kokmilfa). 

+ J/. seclusus Ik-rgroth. — Mt. 

vXysius Bacckstrooni Hergroth.— Mt, Mf. An almost cosmopolitan genus 
witii numerous species in Xew Zealand and Melanesia, east to Samoa; greatest 
concentration in Hawaii. X. Pcrrcks/rorifii is closer to X. Ilutioni V>. White from 
Xew Zealand than to any American species (KoKMlLKv). 

+ V Robnisowclioris tingitoidcs Kormilev. — Mt. P^orms a separate tribe. 



DERIVATION OF THE FLORA AND FAUNA 313 

Miridae (50). 

+ Derophthabna ferna7ideziana Carv. — Mt. A neotropical genus (Braz., Argent., 
Urug.). 

+ +Kusche liana masatierrensis Carv. — Mt. 

Nabididae [21). 

Nabis (Reduviolus) pujictipennis Blanch. Chile. — Mt, Mf. A world-wide genus. 
Reduviidae (2g2, jio). 

Empicoris (Ploeariodes) rubrornaculaius (Blackb.). Almost cosmopolitan. — Mt, 
probably adventitious. 

+ Metapterus additius Wygodz. — Mt. A wide-ranging genus (Amer., also 
Chile, S. Eur., N. Afr., W. Asia). 

■vM. kuscheli Wygodz.— Mt. 

^M. fnasatierrensis Wygodz. — Mt. 

Ploiaria chilensis (Phil.) Kuschel. — Mt, Mf. Almost cosmopolitan; also in 
Chile and probably adventitious in Juan Fernandez. 

Homoptera. 

Apkididae (communicated by Dr. Kuschel): 4 introduced species on garden plants. 
Cicadellidae (57). 

+ +Evansiella 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. 

-f-A^. minos Fennah. — Mt, Mf. 

+ N. oreas Fennah. — Mt. 

+ N. calypso Fennah. — Mt, Mf. 

+ N. philoctetes Fennah.— Mt, Mf. 

+ N. vulca7i Fennah. — Mt. 

^Delphacodes kuscheli Fennah. — Mt. A widely distributed genus. 

■vD. (Sogata) selkirki (Muir) Fennah. — Mt. 

Jassidae [21). 

-f + Alloproctus amandatus Bergroth. — Mf. 

Some zoogeographical 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. 

A^^^r^/>/^r^.— Endemism strong, 4 species of 5, 2 forming an endemic genus, 
the fifth an American species. Relations presumably Andean. 

Lepidoptera.—0{ the 26 indigenous species 18 (69 %) are endemic, and there 
are four endemic genera. Of these, Apothetoeca and FernaTidocrambus are related 
to world-wide genera also represented in S. America, Hoplotarsia to an American 



314 C. SKOTTSBERG 

genus; the systematic position of JuaJiia has not been stated. Together they 
include 6 species. Of tlie 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. Kulia (3 sp.) and Lohopliora 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. Kusc;ilEL'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, S. American or 
more wide-ranging genera with six or more species each, MolopJiilus with 19 
(all end.), Scatella with 16 (all end.), Lii)wiiia with 13 (12 end.), Leptocera 
with 10 (3 end.), MycctopJiila with 7 (2 end.) and Podoiwmus with 6 (4 end.), 
together 71 species of which 56 (79%) are endemic. 

The Xeotropical-Chilean character of the fauna is obvious. This is what we 
ex[)ect (juite ai)art 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 w^ould be Prosopantruni flavipes (austral- 
tricentric] and Inicellia jjitcDJicdia, said to be distributed over "Oceania". There 
are some striking cases of disjunction, suggesting bipolarity {IJoyella, Hydrobacniis, 
Podouoifius Kiejferi], but the distances will perhaps be lessened when the dis- 
tribution becomes better known. 

CoUoptoa. — I want to emphasize that of Dr. Klschel'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 s|)ccies hitherto reported from Juan Fernandez is 
103, belonging to 19 families, [)erhaj)s little more than V'.i of the species found 
there. Eleven species are anthropochorous. Of the remaining 92 no less than 74 are 
endemic — 80 "o, only 20% having been found elsewhere. Future research will 
alter these figures, I suj)p()se, 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 
j)rove to be endemic; of 22 indigenous species enumerated by AURIVILLIUS 21 
were described as new. 



DERIVATION OF THE FLORA AND FAUNA 315 

So far 49 indigenous genera are cited, of which lO are endemic; five of 
these are CurcuHonids. 

The non-endemic species are, with one exception — Xyletomerus piibescejis, 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, i in New Zealand and none 
elsewhere, as well as Pyawmerus, Stroiigylopterus, 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. — The 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 [']6.6%] 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 S. American forms. Haplogonatus is essentially south- 
ern, Prenolepis obscura is a southern, mainly Pacific ant. ApJielms jucu7idus and 
Stilpnus gagates are said to be boreal, but in these as in other similar cases 
the possibility of accidental introduction must be considered. 

Hemiptera. — The 21 indigenous species — there are 7 adventitious ones — are 
by no means a fair representation of the Rhynchota 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, Micry- 
menus, is most nearly allied to a genus in New Zealand. Buchaiianiella is quoted for 
Madeira, Tasmania and Hawaii, but not from America, Nysius, 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 NlLS 
Odhner who put his unique knowledge of this group at my disposal. The 
additions and changes in his earlier list (i8g) 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. 

E7idodontidae . 

-f +A?nphidoxa helicophantoides Pfeifif. — Mt. The genus (only 2 species known) 
is related to Stephanoda. 

+ A. marmorella Pfeiff. — Mt. 

+ Charopa (Endodonta, s. lat.) involuta Odhner.— Mt. Numerous species, 
Polyn., N. Guin., Austral., N. Zeal. 

-f C. occulta Odhner. — Mt. 

-f C. skottsbergi Odhner. — Mf. 



3l6 C. SKOTTSBERG 

^Pioictuni conicion Odhner. — Mt. A genus of a rather small number of spe- 
cies, in various parts of the world. 

^P. depressu))! Odhner. — Mt. 

^- Radiodiscus inasafucrae ((Odhner) Pilsbry. — Mf. An American genus of 
few species, found in X. America (Arizona) and in S. America, south to Patagonia. 

■^StepJuDwda iirctispira PfeitT. — Mt. About 30 species in western S. America 
(Chile, south to Fuegia). 

+ .V. ceratoides Pfeiff.— Mt. 

+ .V. quadrata b'erussac. — Mt. 

+ 5. sclkirki \\. A. Smith.— Mt. 

-S. tcssellata Muehlf.— Mt, Mf. 

HilicJdae. 

Helix aspcrsa Muell. Cosmopol., introduced to Chile. — Mt. 
Li))iacida€. 

Agrioliiuax agrestis L. Very widespread, introduced to Chile. — Mt, Mf. 

Liuiax arhoruui Buch. -Chant. As the former. — Mt, Mf. 

Mi/ax gagatcs Draparn. As the former. — Mt, Mf. 

Sitcciiicidae (subfam. Succineinae). 

+ Succiuea cuuiingi Reeve. — Mt. Succhiea is taken in its old sense; it has 
been split up, and the Juan Fernandez species belong to a group which must 
bear a diflerent name. It is reported from N. America, Panama, Galap. Is., St. 
Helena. S. Africa, Hawaii and Tahiti. 

+ 6. fcr)ia7idi Reeve — Mt. 

.V. fragilis King. (syn. S. texta Odhner). Hawaii. — Mt. 

+ .V. gayajia (D'Orbigny) (Odhner. — Mt. 

+ .V. Diasafucrae Odhner. — Mf. 

+ .S". pifigujs (Pfeiff.) Reeve. — Mf. 

+ .S". seuiiglohosa Pfeiff.— Mt, SC. 

Tor)iatclli)iidae. 

^ + I''ir?ia?idfzia buliniojdes Pfeiff. (inch consimilis Reeve). — Mt. 
+ /'\ couifcra Reeve. — Mt. 

.' + /'". cyli)idrella Odhner. — Mt. Possibly identical with Torvatellina (F21asma- 
tina) tunita Anton, credited to Oi)aru I.* 
+ /'". diapluDia King. — Mt. 
+ /'\ expivisa Pilsbrw — Mt. 
+ /'\ inorjiata Pilsbry. — Mt. 
^ F. lo)iga Pilsbry. — Mt. 
+ /'". pliilippiiUia Pilsbrw — Mt. 
+ /'". splendid a Anton. — .Mt. 
f/'\ tryojii I'ilsbry. — Mt. 
+ /'\ /('ils()?ii I'ilsJMy.— Mt. 

' Rapa seems to be tlie 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. (inch Hawaii), Melan. (Kermadec 
Is., N. Caled.) to Austral, and N. Zeal. 

T. bilamellata Anton. Recorded from Oparu I. 

+ T. callosa Odhner. — Mt. 

+ T. conica Anton. — Mt. 

+ T. plicosa Odhner. — Mt. 

T. reclusiaia Petit. — Mf. According to Odhner probably identical with T. iiir- 
rita Anton. 

+ T. trochiformis (Beck) Pfeiff. — Mt. 

T. 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. 

Zonitidae. 

Hyalinia alliaria Miller. A widely distributed, anthropochorous species. — Mt. 
H. 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 oi 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. — 8 sp. 
Amphidoxa (2), Radiodiscus (i), Stephanoda (5). 

II. Pacific element. — ^23 sp. 

Charopa (3), Fernandezia (11), TornatelHna (8), Tornatellinops (i). 

III. Austral (or more widespread) element. — 9 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 [2^8) 
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 



3l8 C. SKOTTSBERG 

j)ossibility that, at some period of its existence, it has formed part of the con- 
tinent. This is e.g. the case witli 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 perhaj:)s thousands of metres is required to bring it into contact 
with a continent. Mavr (/J^), 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 ()ceanograj)hical 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 [)er{)lexing 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 P>rnandez, but even so it seems worth 
while to (juote 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. 

H.MI.F.V Willis (2S4) 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. lie 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 
i)een as a whole to deepen the basin in conse([uence of the expansion of the under- 
lying ro( ks. 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 j)criod 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 
})eriods of orogenic activity alternated with those of (piiescence. 



DERIVATION OF THE FLORA AND FAUNA 3I9 

Willis did not question the permanence of the basin; what interests us here 
more particularly is his belief in the instability of the marginal regions. 

In H. E. 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 (116), 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 foundadon 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. P'or 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 



SKOTTSBERG 



Field observations show that those islands have had histories which present marked 
diftercnces from those of the continents and their marginal island arcs. 'J'hus, they 
appear to be comjiosed almost entirely of volcanic material, mainly basic, together with 
"coral reef" formations, whereas granitoid intriisives and acid lavas are lacking, to- 
gether with the sediments invarial)ly 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 ])eculiar topograj)hy of that portion of the Pacific 
floor which sej)arates the great island arcs and lands of "continental'' character from 
the more central grouj)s. Thus, on the American side of the Pacific, the "continental" 
lands are separated from the groups — such as Hawaii, the Marcjuesas, the Society Is- 
lands, and Juan I'crnandez — by a series of deep discontinuous ocean trenches, prac- 
tically collinear (j). 203). 

The question is whether these trenches are of quite the same nature and 
date from the same j^eriod 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 copj)er, 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, j)ortions of a continental margin which later 
be< amc involved in a j)Owerful movement within the marginal Pacific, resulting in the 
gradual submergence of these outer ))ortions, the present Hebrides, Tonga, and so on, 
being built upon su( h 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 j)C)int of interest is the association of great ocean deeps with youthful 
volcanic zones, and inasnuu h as the trough and crest of an earth undulation are parts 
of the same strut ture, it is a legitimate inference that the great Pacific deeps or trenches 
are relatively youthful structures (p. 203). 

Andkkw.s 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 |)resent. 

2. The great bordering island arcs of the Pacific — such as the Aleutians, Japan, 
the IMiilippines. the Netherlands Fast 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 (1J2). About 160 flat-topped peaks, 
presumably truncated volcanic islands, rise from 9000 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 loooo 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 (i2j). He finds (p. 7) that there is 

21 -557857 The Nat. Hist, of Juan Fernandez and Easter Isl. Vol. I 



02 2 t'. 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. 'J'here 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 
boundarv of the Pacitic l>asin. 

This is in conforniit}- with what I have quoted above from other sources. 
However, 

certain areas of the I'acific 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 niaj), (il TKXHKRC] goes a good way beyond the Juan Fer- 
nandez-San Anibrosio rise, but the blaster 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 
"granitic la\-ers of the continents continue far out under the bottom . . . probably 
at least some continental rocks underlie its bottom throughout its area". 

Turning to tiie speculations of biologists I shall quote some representatives 
from the two op|)osite camps. Arldt [6') did not draw his conclusions merely 
from facts of distribution but compiled a wealth of geological, palaeontological, 
bathymetrical dates and so on, and constructeci 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 F^ogene and 
left the west coast of South America in the same position as to-day. F>om what 
he sa)-s 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. Camphkll was for a long time a supporter of the land-bridge 
theorv. 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 ui)lift of the great Cordilleras (</./). Later, 
when discussing the Australasian element in the Hawaiian flora he expresses him- 
selt very positively: "We are justified in assuming the former existence of land- 
masses of consideral)le size, connecting more or less directly both Australia and 
New Zealand with Hawaii' (./-r. 22i); and when, for the third time, he took up 
the history of the Hawaiian flora, he expressed himself as follows (46. l8l): 

We may assume that the Hawaiian Arc hij)elag(), as it now exists, is but a remnant of 
a much larger landmass which has been in subsidence for a long period, and that extensive 
subsidence has also o( ( urrcd throughout Polynesia, and to a lesser degree in Australasia. 
One argument tor this assumjition is the great development of coral reefs in the Pacific, 
espec iaily in Polvuesia and northeastern Australia. The existence of active coral reefs 
involves continuous subsidcMKc 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 sul)mergcd 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 vvas converted to wegenerianism 
in the form modified by Du ToiT, thinking that "this theory would best explain 
most of the problems in the geographical distribution of the floras of the Southern 
Hemisphere" (^7.70) — he seems to have forgotten how badly Hawaii fits into 
the picture. 

GuiLLAUMiN, in his paper on the floristic regions of the Pacific {iiS), devotes 
a chapter to its geotectonic history. 

Un vaste continent parallele a I'equateur a du relier les regions australiens, canaque, 
et neozelandaise a I'Amerique du Sud tandis que la region malayo-polynesienne et le 
domain hawaiien, formant peut-etre un continent, en etaient separes. 

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, Kermadcc, Tonga 
etc., persisted longer and is perhaps better founded, but when GuiLLAUMiN thinks 
that his hypotheses "ne sont pas en contradiction avec les donnees geologiques 
ni avec ce qu'on salt 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 (121) 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 
(/05. 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 vvas "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 (pc?. 164 etc.); though the flora is "plutot denature 
continentale" it is not necessary to count with terrestrial connections to explain 
its characteristics. Among zoologists, Mayr (779) and Zimmerman (2^8) 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 (i8j. 247) : 



324 C. SKOTTSBKRG 

Kvideiue with regard to the nature and distribution of land snails is of the utmost 
imi)ortance, for it is from the nature of the land-snail fauna in Polynesia that Mr. Pilshry 
has rea( hed the conclusion that the whole of Polynesia, with Hawaii, was once a great 
continental land mass. l\utida- — an ancient generalized type of land snail — is wide- 
spread in Polynesia and Melanesia and not elsewhere ; whereas many groups (e.g. He- 
li( idae and Arionidae) found widely sj)read 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. 

Hut how early or how late.^ To explain the absence of modern land molluscs 
we may have to j^o back to late Tertiar)' times only, and we should need other 
proofs of a distribution of land and sea sufficiently different from the present one; 
perhaps the\- will be found. Ikit 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. SwEZKV (^^2), who for his own part believed 
that the entire insect fauna of Mawaii owed its presence to accidental migration, 
(juotes two of the authors of "P^auna Mawaiiensis" who did, in his opinion with- 
out an\- reason, build bridges where they^ found that they needed them, Meyrick 
and Lord \\'ALSiN(;iiAM. Mkvrkk, an authority on Microlepidoptera, when stating 
that among the endemic Ilaw^aiian genera three were of south Pacific affinity, pos- 
tulated "the former existence of a considerable land area (now submerged) be- 
Iwcen 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- 
j)()rancous. Lord \\'.\i,siN(;iiAM 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 — e(iuall\' be)'()n(l the possibility of exact j)roof 

It is cas\- to understand that, in all these discussions and speculations, Hawaii 
in its isolated j)osition 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 PL. H. l^RVAN's recent 
contribution, "'Phe Hawaiian Chain" [40), ecpially instructive as popularly written. 
Hkn AN 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, nor 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 a|)peared some time during Tertiary, the forma- 
tion proceetling 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. PRVAN, 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 San- 
tahun, 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 [2J1) 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) warden in der Ver- 
schiebungstheorie als von den Kontinentalschollen abgeloste Randketten betrachtet, die 
bei der allgemeinen, vorwiegend westlich gerichteten Bewegung der Erdkruste liber den 
Kern allmahlich nach Osten zuriickgeblieben sind. Ihre Heimat ware hiernach, ohne auf 



326 



C. SKOTTSBERG 



F^inzelheiten einzugehen, aiif der asiatischen Seile des Ozeans zu siichen, der sie jedenfalls 
in den betrachtcten geologischen Zeitcn erheblich niiher als heute gelegen hal)en miis- 
sen. Die biologischen Verhaltnisse scheinen dies zu l)estiitigen. So haben nach G rise- 
bach iind I) rude die Hawaiinseln eine Flora, die am niichsten verwandt nicht mit 
Nordamerika ist, das ihnen doch am niichsten liegt, und von dem heute Lult- und 
Meeresstromung herkommen, sondern mit der alten Welt. Die Insel Juan Fernandez 
zeigt nach Skottsl)erg gar kcine X'erwandtschaft mit der doch so nahen Kiiste von 
('hile, sondern mit Feuerhmd, Antarktika, Xeuseehind und den anderen pazifischen 
Insehi. Doch sei hcrvorgehoben, dass die biologischen Verhaltnisse auf Inseln allge- 
mein schwerer zu dcuten sintl als diejenigen auf grosseren Landraumen. 

In the 3r(l edition j). 59 we read after "Jnseln": "Dies passt vorziiglich zu unserer 
Vorstelluni^, class Siidanicrika, nach Westen wandernd, sich ihr erst in letzter 
Zcit so wcit i^enahert bat, dass der Morenunterschied auffallend wird." In the 
following editions this sentence was excluded. 

Certainly I never said anything like that and I fail to see where Wegp:ner 
got bis strange ideas; just as many other writers I have pointed out that the Andean- 
Chilean element is stronger than any other. F.vcn to a firm believer in the festoon 
theory the Juan FYTnandez and Desventuradas Islands ought to ofifer insuperable 
difficulties. \\'K(;i:m:r 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 decj) 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 tlie 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- 
j^lacement theory. 

\\'K(;i:M.k's theories were taken up by Dr Toil' and presented in a modi- 
fied form ((S7); I shall (juote his attitude toward the festoons. 

As Wj'.cini.r has observed, they are all comj)arable in size, regular, linked to- 
gether en (,( helon and convex to the Pacific; each shuts off a large ])ortion of sea and 
tronts an ex eani< deep, while the concave side bears a row of volcanoes. To Suess we 
owe the (onception of the deve]oi)ment of successive arcuate asymmetrical fold-waves mi- 
grating outwards from the more stable "Amphitheatre of Irkutsk", which led to ])rogressive 
e\i)ansion of .Asia towards the Tac ific. While the hypothesis has since had to be ap- 
jircc iatcl\ modihef], its fundamental ideas have been brilliantly confirmed by subsequent 
investigations... Significant are the (x^eanic fossae that immediately front the convex 
sides of the arcs — foredeeps subsiding in nih'ance of the outward-moving geoantoclines 
and incidentally tracts ot marked coastal instability (pp. 186-187). 

How tar (lid this outward movement, these advance-folds proceed? Does Du 
Toil allow all the Pacific island chains to be linked up here? When the great 
W'NW swing of Asia is rcj)laced by an expansion toward the Pacific, the system 
of rifts in the ocean floor, oxer 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 ri|>ples 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 ToiT 
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 ToiT 
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 Toit'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 

Xumerous 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 symi)osium, arranged in 1950 (dj) offers an opportunity to get ac- 
quainted with their attitude. 

J. II. T. L'MlKiKOXK, TJic case for the cvust-substratiiui theory, pp. 67-71. 

Si Kss' 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, a})[)roximately I300°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. 
U.Ml'.fiKovE asks if not the thick blanket of sediments would have been squeezed 
and [)iled 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 
recjuires, the Atlantic originated in comparatively recent times, how are we to 
explain the enormous thickness of its bottom sediments, according to Hans Pet- 
ri'.KSsoN a maximum of loooo feet, "representing a time-span of 300 to 400 
million years", which would bring us back to the Palaeozoic. If continental drift 
e\er occurred, UMliGROVE asserts, it took place some 3000 million years ago and 
C()nsc(iuently loses every shade of interest to the biogeographer. 

HakoM) Ji-.iikess, Mechanical aspects of continental drift and alteriiatii'e tJieories. 

ji;ill<i:\ s definitely rejects We(;ener's theory on geophysical grounds; 
W l.GlNl.K 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'ooi.iiKiix;!;, TJu- hearing of Late-tertiary history on vertical and Jiori- 
zo}ital ino/'cinoits of the co)iti?ients. 

The nature ot the incchanisni of the "uplift" may remain in doubt; its reality can 
hardly he gainsaid . . . in so tar as such comj)lementary foundering took place during 
Tertiary times, it is not directly relevant with the drift problem . . . Vertical movements 
otter an alternative solution to some of the 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. 8i). 

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 Jeff'reys 
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, The 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 if 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 conti^iental 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. Good, The distribution of the Flowering Plants in relation to theories of con- 
tinental drift. 
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 V2 to 4 times as many as the mono- 
cots), and the absence of an\' stron<^ly marked South Pacific flora south of Hawaii 
and east of Xew (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 j). 74: 

A coiiimon expression of the theory of continental drift postulates that the sunder- 
int; of various coiiiinental masses began in a relatively remote geological period and 
that it iuis 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 
ma^tcr-kcy 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 I'>rnandez, etc.) and Antarctica, Australia and New Zealand, vertical move- 
ments seem to otTer a less distant possibility. 

In a most cleverly written chapter Gooi) [log. 344-360) discussed land- 
bridge versus continental drift. If we cannot, he says, accept the former, nor 
put our trust in disj^ersal, 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 (//?) 
|)()ints 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 anv of the Polynesian or Melanesian groups, makes it impossible to deny 
both land bridges and the efficiency of transoceanic dispersal without providing 
for nngration with the help of shifting continents, (iooi) 

has run into an impissc f)vcr the Pacific islands like Hawaii. He has rejected the land- 
bridge hvpothcsis in fivour of ( outineutal drift.... Pnit he excludes continental drift so 
tar as the ishmds arc ( oik rrned, for he accepts them as truly oceanic, not continental frag- 
ments. \'et he will not ;i! i -pt oversi-as migration. Well, I can't see what explanation 
remains, if all the-e tlinr ;>!.■ e\( hided, but the plants are there (p. 148). 

W ri.l I {j<j/\ found that the biogeographers have good reason to support 
\\ i;(;i:m;Iv s tluM)r\-; much s|)eaks against it, but he trusts that the difficulties 
will be overcome by and 1)\-. He recognized that, with regard to the Pacific, 
they are ver>- considerable and call for a modification of the theory. 



DERIVATION OF THE FLORA AND FAUNA 33I 

Chapter VI. 

Transoceanic migration. 

In some of my earlier papers I touclied upon the great problem of long- 
distance dispersal and the supposed efficiency of the transporting agents; see for 
instance 2ji. 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 [8^) and Grisebacii 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 (j^j. 469): 

So merkwiirdig es auch sein mag, dass sogar einzelne Holzgewachse sich hier 
iiber das Stille Meer verbreitet haben, so ist ihre Wanderung doch aus der antarktischen 
Meeresstromung, den herrschenden Westwinden, oder durch Mithilfe der Seevogel, 
vielleicht auch durch alte Verkehrswege wohl hinlanglich zu erklaren, 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 Besiedeliing einer Insel erfolgt, haben die oben angefiihrten Beobachtungen 
bei der Wiederbesiedelung des Krakatau gelehrt . . . Aber die Entfernung des Krakatau 
von den nachst gelegenen Inseln ist keine allzugrosse, sie betragt nur etwa 18—40 Km, 
also Entfernungen die auch durch die Flugtiere nicht allzuschwer iiberbriickt 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 zunachst gelegenen Festlandern (und Inseln) erhalten haben, 
wenn auch vor undenklichen Zeiten und ganz allmahlich. Dafur spricht auch der Um- 
stand, dass die Flora dieser Inseln keineswegs von der ubrigen 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 weitcrer Aiistaiisch der Florenelemente zum mindesten wesentlich erschwert 
iind eine weitere /uwandcriing von Elementcn der Festlandsflora wenig wahrscheinlich. 

If tliis be true, lunv was immigration over tbousands of miles ever possible? 

Several writers who have {)aid special attention to Pacific problems occupy, 
more or less do^^matically, the same standpoint as Havek. Setciiell, with whom 
I had the j)ri\ile<;e to discuss this subject on various occasions, was already quoted 
{). 271 ; I shall add here what he says, in the same paper, about migration (2ig. 300). 
He found that 1 was "too narrow" in my allowances for migration possibilities; 
he belie\ed 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 Tertiar)', but in his summary pp. 307-309 admitted the possibility of their 
being considerabl}' 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 difi'erent 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 (ji'p. 537): 

'{"he seeds of j)lants 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. 

r'l.oKiN has, he writes, shown that conifers of the south hemisphere have migrated 
freely from Australasia to South Ainerica and vice versa, w-hereas mammals are 
unable to pass and are absent from oceanic islands — but is it not customary to 
place them on a j)ar.' S'lEHl^.lxs' 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 
o{)|)osite side the width of oj)en 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 h'()Siii;K(; : 

. . . transo(eani<; migration across at least 2500 miles without stej^ping-stones is not 
only a possibility hut a relatively common occurrence (99.867). 

Im)SI!I:i<(.s subject was the American element in the Hawaiian flora, but in order 
to exj)lain the j)resence f)f the dominant Australasian element we must count with 
still gieater distances. AxKl.RoD (7^) quoting F'o.SBERG takes a modified position. 
In case ot distances not exceeding some 200 or 300 miles there are no difficulties, 
"a coni|)]ete 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 probabiHties 
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 m.uch 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 [2JI, 318, 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 (121), 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. I expressed my doubts 
that it had ever been effective, in any case with regard to seed-plants. 

I have already remarked that Setctiell 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 {2ig. 300). His ideas are clearly expressed in 218 (\). 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 Setciiell, 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 
{26J. 572). In recent volcanic islands are many unstocked habitats to begin with, 



334 ^- SKOTTSBERG 

but each new species would mean competition, and the entry of an additional 
species would be very difhcult and consequently of rare occurrence. This may 
be so, but tiiere are nian\- 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, suj)posing that he likes climate and soil. Experience shows that 
numerous aggressive plants brought by man, not only herbs but also trees and 
shrubs, tind suitable living conditions even in undisturbed natural communities. 
In Juan Fernandez I have had occasion to follow the invasion by Aristotelia maqui 
(chilensis) and to witness the fabulous ease with which it crowds out the native 
vegetation, and to observe I'i^Jii Moliiiac springing up on the ridges where the 
j)lant cover was o|)en and, from there, to enter the dense native brushwood. 
And they are only two of the many successful weeds, a third one, equally dan- 
gerous but of quite recent introduction, is Rubus nluiifolius. All three have fleshy 
fruits and are eaten by man and birds and propagate themselves rapidly. They 
are conunon on the opposite mainland (where, of course, the brambleberry was 
introduced from luirope), but man, no bird, carried them across to the islands. 
Similar examples are, I presume, oftered by almost all oceanic islands. I just hap- 
j)ene(l to read a book on Cape Verde Islands, where a naturalist tells us about 
La)ita)ia caniara sj^reading like wildfire and menacing the little there is left of 
natural savanna and steppe [326). 

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 ofl" 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. 

Pl-.KKINS, in his introduction to Fauna Haw-aiiensis (j^j. XLVl), wrote: 

All the islands being volcanic and having been built up from a great depth of ocean 
at various j)eriods, 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 Hke the flightless insects and jirobably the land Mollusca. 

Dritting 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 
ser\e no |)urpose because it is unlikely that the passengers would become acclim- 
atized in ilawaii. The fauna must have come from the warmer parts of America, 
from Australia, Polynesia etc. "at rare intervals from the F^ocene until now". If 
we have to belie\e the geologists, no Ilawaii existed in the F^ocene — and how 
did those, who arrive "now", manage to become endemic genera and species? — 
non-endemic (lowering plants not brought by man are few. 

In some instances (ii LICK [Tig] admits the possibility of land connections, 
but Ilawaii, Juan hernandez, (ialaj)agos, St. Helena, etc. etc., have always been 
isolated. 'I he (juestion 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 Galapagos 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 (iig.414) first. 

It is possible to go far toward a first diagnosis of the degree of a land's insularity by 
noting how exclusively it is 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" — GULlCK 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 [lyg] includes 



336 C. 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. 

liirds are excellent rivers and thus cai)able of ra])id 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 untjuestionably 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. Mavr continues p. 198: 

'i'he possil)ility of transport by floats or in logs is not to be underestimated. Many 
tro])ical currents have a sj)ced of at least 2 knots, that is, 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 Kaster Island, juan Fernandez and Saint Helena is almost unassailable proof 
that such a method of disj:)ersal is a reality. Tropical hurricanes carry for hundreds and 
even thousands of miles. . . . I'he result of the recent surveys in the Hawaiian Islands, 
the Mar([uesas, 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 Mavr 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 hclj) 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, 
hf)wcver, ])Ostu]ates even today continental connections between all or nearly all the is- 
hmds where land shells exist. 

As we shall see below ([). 350) Mayr declared himself unable to explain how 
land shells arc (lisj)crsecl. 

It seems that, also with regard to the birds, Mavr contradicts himself, for in 
the same paper (j). Kj.S) he asserts that most birds, particularly on tropical islands, 
|)recisely the islands we are discussing, are extraordinarily sedentary, and as an 
examj)le he mentions that of 265 species known from that part of New Guinea 
which is ()j)posite 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 Cirri'V's fruit-eating pigeons which were 
thought to be res|K)nsible 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 Geiitiajia prosirata Haenke as the best example. Its distribu- 
tion is due, he says, to the wanderings of Diomedea exulans which, 

abweichend von der Lebensweise der meisten anderen Zugvogel, iiber beide Hemispharen, 
von Kap Horn bis zu den Kurilen und Kamtschatka, wandert und die Standorte jener 
Pflanze in der arktischen und antarktischen Flora in Verbindung setzt. Mil der Beute, 
die dieser Vogel verschlingt, kann er auch Samen von Pflanzen, welche, mil den Fliissen 
ins Meer gespult, in den Magen der Fische ubergehen, in einzelnen Fallen ausstreuen, 
so dass sie an fernen Kiisten aus seinem Dunger aufkeimen [32^. 469). 

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 Gentiana 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. prosirata 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 (^/c^'. 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 [26 j), 
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- 

^ According to Taylor (p. 570) the two truly Antarctic phanerogams, Deschampsia ant- 
arctica and Colobanthus crassifolius, 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, Tavlor states, propagules suited to bird transport. It is sur- 
prising that Wkrhi [jiy), 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 ( -,v). 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 
intlications 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 Ta\i,()R 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). 

W'ali.ack calls attention to sea birds breeding on islands in the tropics; 
Phaeioji 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 GiUHS (?.?/), 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 subaerial mountain 
range {2()8. 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. Pie 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 'j^y 
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 supi)ly 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 (^/j), describing the distribu- 
tion of Heteroptera, also looks for convenient stepping-stones. 

Divergent opinio7is 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 (20^), 
a firm believer in the great value of all kinds of dispersal mechanisms, some of 
which are, of course, very, wonderful. WULFF (2^1), in his 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). GoOD observes [log) 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 (21'/) was inclined to give considerable credit to 
migratory birds. The occurrence of identical species of flov.ering 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 
j^enera. It is not improbable that the secret of their origin and early history lies 
hidden in the far soutii, but tiiis is all we can say. Hemslev (727. 66) when dis- 
cussing the endemic Coni[)ositae of Saint Helena and Juan Fernandez said that 
"wind seems at first to be the most probable agent"; still he doubted its efficiency. 
In Setciiell's view storms seem to ofler more than a possibility in many in- 
stances, particularly tropical cyclones and vertical thrombs able to carry even 
lieavy diaspores to a great altitude. In another paper (218) 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 anomal}' together with adverse biotic factors, but I fail to see why they 
shouldn't ofter the same difficulties for diaspores coming from the west. 

Andrews (^.615) paid s})ecial 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 const of South America, where it is joined by the cold 
uprising water along the South American coast. This gives rise to the north-moving 
Peruvian C'urrent which sweeps by Juan Fernandez, Peru, Central America, and Mexico, 
whence there is a detlection 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... (2 5 genera are enumerated). The influence 
of ocean currents is suggested i)articularly in the peculiar distribution oi Acaena, Gunnera, 
Acrfcra, On-obolus, Sautalinn, Sophora, and so on. 

Juan hY-rnandez 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. I 
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 ( iVeS). 

\\ ith regard to the cyclonic storms several authors have, as we have seen, 
em[)hasized their j)r()minent role 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 trans|)()rte(l, and it is useless to deny that such events have 
taken j)lace and still take })lace, 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 c>'clones 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 j)apers \'lsiiER has summarized his studies on cyclonic storms in the 
Pacific. Three chief centres of origin are distinguished [322): (i) Western N.Pacific, 
originating some distance east of the Philippines in lat. 8° to 25°; (2) Western 



DERIVATION OF THE FLORA AND FAUNA 34I 

S. Pacific, particularly between Australia and Samoa in lat. io° 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 (i) is WNW, 
recurring NE, and of (2) WSW, recurring SE. To what extent plant distribution 
runs parallel to cyclone tracks has, as far as I 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" (j^J. 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 (j^^) 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 ViSllER 
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, Coniogramtne and Loxogra?nme, have flown the north Pacific from 
Japan to Mexico, each in one single instance. It is not exacdy impossible that direct 
colonization has occurred between Chile, New Zealand, Tasmania, the Cape, and Tristan 
da Cunha. 

It should be mentioned that CllRlST (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 Irmsciikr tried to prove that the distribution of plants strongly 
su|:)ports \\'K(;fnkr'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 [14.3. 291): 

Dass bei den Hliitenptlanzen die J^eforderung der Samen und Friichte durch Wind, 
Wasser und I'iere ganz wescntlich eingeschrankt werden muss und fiir geschlossene 
i-'orniationen auf grossen Kntfernungen hin nicht in Frage kommt, ist heute von den 
rtlan/.engeograi)hcn allgeniein anerkannt. 

Irmsciikr 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 s{)ore-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. iRMSCiil'.R remarks that already in 1903 SlEPllANl denied that liverworts 
are able to make long and successful jumps. Attention should be paid to Domin's 
valuable j)aper [yd] in which he brings together numerous facts illustrating the same 
definite distribution patterns in this as in other groups, and Miss FULFORl) (e.g. 
10 ;] 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 ra[)idly when exposed to the air — a promising field for experiments. Regard- 
ing mosses I refer to Herzog's work [i2g) where he speaks against the belief in 
the importance and great range of dispersal through the air. We find, IRMSCIIER 
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- 
seli)cn \'erhrcitungszuge ihrer Disjunktelemente ergeben, zeigt wohl einwandfrei, dass 
hierfur der "Wind" ebenso wie andere iiussere Krafte als Ursache abzulehnen sind. 
Wiiren sic in aiisschlaggcbender Weise an der Ausbreitung beteiligt, miisste die Besie- 
dclung entsj)rechend der Verschiedenheit der Friichte, Samen und Sporen ebenso ver- 
schiedcnartig ausgetallen scin, d. h. in den einzelnen Gruppen dieser biologischen Ver- 
schiedenheit entsprcchcnde charakteristische Merkmale zeigen. Dies ist aber nicht der 
i''all. Der alien vicr druppen ge m e i n sam en hochdisjunkten Ausbildung so vieler Areale 
nuiss vichnehr cine andere Ursache zu (irunde liegcn . . . 

This conniion cause was, in Irmsciier's opinion, continental displacement in the 
sense of Wi;(;ener. Those who disagree with him will have to look for vertical 
movements, emergence and submergence of land. 

Lichens, fungi etc. were not included in Irmsc:her's discussion. A survey of 
their distribution patterns is something to be asked for. Lichens are said to de- 
|)en(l on their vegetative reproduction bodies more than on spores. 

It is maintained that, with certain exceptions, terrestrial animals spread 
less easily than plants. 1 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 sup[K)sed 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 [dj) 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. 

W/iat kind of invertebrates are likely to withstand transoceariic 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 



344 C. SKOTTSBERG 

size, flight capacity, habits, mode of reproduction, sensitiveness to changes of 
miUeu, and so forth, and I am afraid that we have httle knowledge, founded on 
facts, in most cases. 

liarth-iroDus 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 leeches 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 (z/^. 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 criistaceaiis 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 spiders 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. Hkki.anI), pointing to the general distribution of genera and the high spe- 
cific endemism, is in favour of former land connections [2j. 1052). 

L'isolation doit etre assez ancien pour ce que cette fauna ait pu acquerir les carac- 
tcres d'endemisme (ju'elle presente. Un botaniste a fixe vers le Pliocene cet isolement, 
mais jc serais porte a croire (pi'il est i)lutc)t ])lus ancien, en me basant sur la lenteur 
de revolution des Araignees. 

AI.\\R, who rejects all land connections, remarks (27^.214): 

Considering the haj)ha7,ard manner by which these oceanic islands receive their popu- 
lations, it is rather astonishing how similar the faunas of the various islands are. Ber- 
i.ANi), 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 j)aradox of tlie similarity of the fiiunas of oceanic islands is solved in quite dif- 
ferent manner. Of all the j)Ossil)lc 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 [183) 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. Pseudoscorpio7is 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.e. 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 Myria- 
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 Collembola I 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 [i6y. 7 1 9) writes : 

The primitively wingless Collembola seem to constitute better material for distribution 
studies than any other insect order, because migration by flight is imp'ossible 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 Thysanuva 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 insects, 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. 



346 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 Sightless insects have, just as the flightless birds, given cause to 
much s[)eculati()n. 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 oft' 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 Ortlioptcra 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 
tciDiitcs 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. Tliysanoptera seem to be easily' spread with human 
traffic. Most of the 90 species found in Hawaii are adventitious, few indigenous. 
Two of the 4 species in Juan h'ernandez are endemic. How these delicate insects 
manage to get about and to reach oceanic islands I cannot tell. 

Tlie Hawaiian islands have a rich and peculiar fauna of Xcuroptera, some 
"among the most aberrant of all" (ZIMMERMAN p. "j6). 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 tlie same when we get to the Lepidoptera, about lOOO 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. vSo far only 26 indigenous species are known from Juan 
I^'ernandez, 70% of them endemic; when the list of Dr. Kusciiel's collection, which 
contains over 50 species, has been published, the figures will undergo alteration. 
In his survey of the Pacific lepidoptera Swezev [262) includes the Galapagos 
Islands, but does not mention Juan Fernandez. The Pacific islands were, he says 
p. 319, j)opulated from the Malayan and Oriental regions, and the fauna arrived 
in the main by accident, winds, ty[)hoons 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 Diptera 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 i 50 indigenous species (64 % endemic). Our 
knowledge of the dipterous fauna is in the main due to Dr. KuscilEl/s collec- 
tions; the 25—30 species not yet reported on will raise the total number. 

Over 1600 species of Colcopiera are reported from Hawaii, of which about 
'j^ % 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 (77^.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 taeevils, 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 (2yj) came to the conclusion that they must possess 
some unknown special means of dispersal. Brinck, who discussed the coleopterous 
fauna of Tristan da Cunha (316. 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 ofl"ers 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 

Xt'sotflcus is evidently a relict endemic genus of four closely allied species, and 
there aj)pears to be nothing like it elsewhere . . . Oodemus with its 58 species ... is 
the largest genus of the Hawaiian Cossoninae . . . together with its close ally Atiotherus 
(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 brini^ 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 yidviorJuui of .Vl KI\ILL11 s. 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 Brvan [j^g. 188) 
finds a Malayan origin most acceptable, GULICK writes (7/^.420): 

The history of Hawaiian land birds must have begun with the arrival of some form 
of troi)i(al American honey creeper, which became in due time the progenitor of all 18 
genera and 40 sj)ecies of the Drepanididae. 

Me seems to have forgotten that the islands are claimed to have cHved out of 
the ocean in late Pliocene and Pleistocene times. 

I lyinoioptcra 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, ZlM- 
MKRMAN pays special attention to the genus Xysius, which has its greatest known 
diversity in those islands. All are endemic and include the most divergent of all 
Xysius species. The ancestors are supposed to have come from the south and 
west Pacific. The genus is, according to UsiNGER (2'/j) common in the Australian 
and Oriental regions, extending through Melanesia to Fiji anci Samoa without a 
single re[)resentative east of this line, but important in the Hawaiian chain; the 
author seems to have overlooked its occurrence in Juan Fernandez. Xysius is 
su()posed 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, l^ut in other cases it is less 
eas)- 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 inexi)lical)le by present theories" (I.e. 315). Why not 
presume that all related genera have died out.-, which is the easiest explanation. 

Xysius is a widespread genus, well developed also in New Zealand, and the 
single Juan I^'ernandez 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 larid 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 [iig] 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 Partula 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 (I.e. 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: (i) our almost complete ignorance of the means of disi)ersal of 
snails, (2) our lack of knowledge of the speed of speciation in snails, and (3) faulty classi- 
fication, i)articularly generic classification. A. Gulick has already directed attention to the 
])resence of snails on most oceanic islands. 'J'hey were unquestionably carried there by 
some unknown means of transjiortation. . . . 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 eas\' to under.stand 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 wa\' 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, if we approach the problem 
differentlv, 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 si/e loses its weight. However, if small snails can be distributed overseas, 
then what is to j)revent eggs or tiny, immature specimens of large species from being 
similarly transported? (I.e. 61). 

In passing, ZiMMKRMAX quotes H. B. 13.\KKR, who thought that land-shells 
are carried along by migratory birds. After these speculations it is refreshing to 
read Hrvax (^o. 9): 

The j)resence 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, whi( h 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 ])lants or beneath fallen leaves and trash on the 
ground. How did their ancestors reach Hawaii if they could neither swim nor drift? 

( ^f^poju'iits ai^^aiiist the doi^iua of colonization across tJic oceans. 

Many biogeographcrs 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 l)ut leaving behind land fragments, is to refuse 
to accept overseas migration as the only possible means of colonization. Conse- 
(piently, a number of authors have already been quoted in the chapter on the 
history of the Pacific basin, (iooD, Ikmsciikk, WTij-f, Cami'BELL, etc., as well 
as my own contributions to the discussion, the latest in 195 1 [2^8). I shall add 
here that I never disclaimed every possibility of migration over wide expanses 
of water; see 226 and 2:12. X'arious writers have compared isolated peaks on 
continents with oceanic islands and have stated that under present conditions an 
exchange of biota is improbable. \\\x SlKKMS expressed his opinion when dealing 
with the Malaysian mountain flora, in very plain terms [2^8)\ too many facts 



DERIVATION OF THE FLORA AND FAUNA 35I 

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 (20j), referring to the history of the Chilean flora, 
called it "eine Kette von Unwahrscheinlichkeiten". Gordon (iij) 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 S4. 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 (2^^. 81), referring to Gigantodax kiiscJieli, arrived at a different 
conclusion. Gigaiitodax is an exclusively South American genus. 

Chapter VII. 
Biological characteristics of isolated islands. 

In his classical "Lecture on insular floras" [138) 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. SKOTTSBKRG 



1. /;/ all casts ciDisidired flcrislic relaliois exist beiwcen the island and one 
niot/ier coiitinejit. 

2. rite floras of all the islands in question are more temperate in character 
tha)i that of the mother cojitiiioit on the same latitude. 

This ina\' be true in most cases thanks to the influence of the surrounding 
ocean; it holds <;o()tl for Juan P'ernandez. 

3. All these islands show many biological peculiarities by which they are 
distifii^uished. 

Tlie chstin<^uishino characteristics are mainly expressed in endemism. HoOKER 
referred the etidemics to two cate<^()ries, such as do not show affinity to the plants 
on the mother continent, and such as, even if belongini^ 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 man\' cases also very remarkable endemics. Hawaii stand out above 
the others and so do Juan Fernandez and Desventuradas; the floras of Tahiti, 
Samoa, Marcjuesas etc., as well as of Micronesia, are less independent. The degree 
of s|)atial 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. F^n- 
(lemism in angi()sj)erms is 69% in the former and 50 in the latter; of the ferns 
about 30 "o in each, but only Juan P^rnandez has an endemic genus. 

4. the i!;e}ieral rule is that the species also fotaid in the mother continent are 
the most abitndant, the peculiar species are rarer, the peculiar j^enera of contineiital 
affinity rarer stilf but the plajits with 710 affijiity elseivJiere are often i>ery commoii. 

This is, I supj)ose, true of the islands examined by HOOKKR, with the 
excej)tion of St. Helena before the arrival of man, but not of Hawaii, nor of 
Juan l''ernandez. Some of the continental species — Libertia and a few grasses — 
are abundant, while others are rare, all according to the supply of suitable 
habitats; atuong the j)cculiar species of continental affinity are many quite 
common ones, e.g. the endemic species of Acaena, Driniys, Dysopsis, Es- 
callojiia, (iuniicra, Myrceui^enia (the leading forest tree on Masafuera), Pernettya, 
Rhafhithannius, ( ]<^n/\ flrijrcroji fruticosus, Uncijiia l)ouglasii\ and of the peculiar 
genera allied to South yVmerican ones, Xot/iomyrcia is the leading forest tree 
on Masatierra, where Ochaga/da is also common. To these may be added such 
common endemics as lioehmeria and the species of /v/^^'-<'7;'rt: and Cop rosm a, unde^v 
the assumption that related species of lloehnieria, and re{)resentatives of Coprosma 
and i-agara, once belonged to the neotropical flora. The plants of no affinity in 
the luother countr\-, that is the nearest continent, are as a rule very local, few 
are common and many extremely rare. 

Fossil)])- I have misunderstood llooKKK 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 rejjorted from a small number 
of localities only; and if we use "sj)ecies" in plural we mean the same thing. 
Hut what if HooKHK 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 ^/g 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 abse7it. 

Here "rare" must mean few species, and Therophytes are very few in Juan 
Fernandez 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 



SKOTTSBERG 



is considerably diftcrcnt; 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 frecjuent, and the individuals surviving and 
breeding, they keej) uj) 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 j)crj)etuate the continental races (p. lo). 

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 tvpe of flowering ]:)lants; 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. lo). 

11i:msi,i:v [12'j. 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. Fven 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. 'V\Tere the capacity of 
the dis{)ersal 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 (supj)osing 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 (.i""^?^): 

Turning to the natural agents of disi)ersion, winds are no doubt the most power- 
lul, 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 for very short i)eriods, but appear quite insufficient to transport seeds 
over 4000 miles (j). 13). 

Various j)henomenons . . . common to . . . Kerguelen, the Crozets and Marion, 
favour the supposition of these all having been peopled with land plants from South 
.\merica by intermediate tracts of land that have now disai)peared; 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. — The occurrence of numerous genera and species restricted to 
oceanic islands has caused much discussion. "Reichtum an Endemismen ist iiber- 
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 Ciiristensen {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 
{122) 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 WiLLlS [286) claims that wides are older 
than endemics, a rule with few exceptions; in another paper [283) he states that 
"insular endemic genera are as a rule young beginners, not relics". I have dis- 



350 C. SKOTTSBERG 

cussed his theories in an earher paper [jji) to which I refer. RlDLEV {20j) called 
the island endemics "newborn s[)ecies ", admittinf^ that all did not fall within this 
cate^ors- but were "epibiotics, relics at the end of their species life . . . unable to 
reach another suitable spot for their |j;rowth". It is noteworthy that so many of the 
"e[)ibiotics" are Composites, famous among diffusionists for their alleged effective 
dispersal mechanisms; in RiDi.KV's eyes they are, perhaps, pseudo-relicts. Cain (^2) 
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 tind, no island cases as clear as Giiikgo or Mctascquoia 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 stenotoj)ic, but others are what Cain (p. 230) calls "senescent"; such 
species occup\- 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 stenotoj)ic young beginners. If indeed old and senescent they ought to 
siiow 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 
'^ Robi)iso)iia assemblage" in Masatierra offers a good example (see 2ji). 

Xuiubcr of species per genus.— \Nq 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 
tlie numerical relation between species and genus approaches i 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 Ilawaiian flora is the most striking. Fos- 
i:Kk(;, who contributed a cha|)ter on the higher flora to Zimmerman's book (2^8), 
indicates <S3 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. Hiil.Kr.RAND (307) was conservative; from his Flora Drude 
^.^^\^- ^?>^^) g'^'t the ratio 6.2 : i, but scores of well-marked species have been de- 
scribed since i 8<S,S, 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 
ot modifications due to environment has been duly considered. Based on FOS- 
i;i:r(;'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 

1 .AinDiiK^ taxononiists particularly responsible for this alarming^ increase PL. P2. Sheri-F and 
H. .St. joiix should be mentioned first, but some others have also contributed and the writer 
cannot plead innocent. Shkrik, in addition to numerous new species, has described an endless 
number of varieties. 



PERIVATION OF THE FLORA AND FAUNA 357 

Zealand's continentality. According to CnEESF:MAN the number of families is 97, 
of genera 382 and of species 141 5; the relation in question is 3.7 : i. 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 disharmonic and fragmentary: 38 
families, 97 genera and 143 species; the ratio is 1.47 : i. 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, ividespread and '' successfur' 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 oi Jimiperus; 
perhaps junipers possess some kind of dispersal capacity and do not count, but 
we cannot get away from Finns cana^^iensis. 

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 SopJiora sect. Ediuardsia 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 



35^ 



SKOTTSBERG 



improbable. Araceac are pleiitihil in luiinid tropical climates, and some islands, 
the Hawaiian for instance, ouL^ht to offer a suitable environment; still there is not 
a sin<^le 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 Mala\'sia and Melanesia; cf. ?/cV, map 21. However, the seeds, even 
if carried far, rarely are able to retain their viability long enough to become 
established, should they haj)pen 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 
restrictetl areas; their advance over land is slow. A species introduced on pur- 
pose will, in rare instances, become naturalized, but I cannot remember ever 
luning 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 incomj)rehensible. Besides, the submerged links may, 
for all we know, have been {)oor in legumes, arum lilies and orchids. CAMPBELL, 
admitting that the total absence oi Araucana, Agat/us, Podocarpus, Ficus and Ara- 
icac 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 [^6. 181). 

'J he prcpcnidcraiicc of woody plants in oceanic islands. — Hemsley (727. 31) 
em|)hasized 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, (ialapagos, and Juan Fernandez, and are scattered over Polynesia (see 
,VcS'. map 20), and in the Atlantic (Macaronesia, St. Helena) and Indian oceans 
(.Socotra, the Seychelles, Mascarene Islands); arboreous Compositae are, as HemS- 
l.KV just])' remarks, by no means restricted to island habitats but numerous in 
tropi( al regions of the continents, particularly in America. Both Hawaii and Juan 
I'Y-rnandez offer good examples of woody plants belonging to otherwise herba- 
ceous families or genera, but Hkmsi.kv's statement p. 31 about Gwinera\\\]\x2S\. 
I'V-rnandez "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 tJierophytes. — 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 tlie original stock making u|) 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 exj)lain the high {)r()p()rtion of arborescent species in islands iso- 
lated since millions of \'ears. 

1-j'ideiice for a greater antiquity of lignifed angiosper)ns. — The angiosperms 
originated in the Mesozoic and are traced back to the Lower Cretaceous. The 
tree form was the res|)onse to a warm and humid climate. These problems were 
subjected to a comprehensive analysis by SiNNOTT and I. W. Bailey [22f), 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 paleobotany 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. WiLLlS 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 



36o 



SKOTTSBERG 



the first to note the predominance of trees on oceanic islands, tried to explain 
it as a result of natural selection. SlNNOTT and Baii.EV's criticism is absolutely 
C()nvincin<^. 

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 iiiiiabitants 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 j)ossession of large areas, but these were concentrated 
in the north; excej)t the Antarctic continent there was not much land in the 
far south. 

Several authors have expressed the same opinion as SlNNOTT and Bailev. 
Irms(IIi:k (//;) 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 fanfily Cruciferae descends from woody tropical Cap- 
paridaceae (IkMSCllKR might have called attention to the miniature Lepidiuni trees 
in Hawaii): 

Auch sonst ist die Abstammung gemiissigter Sippen von tropischen nachgewiesen 
worden. . . . Diesc Gegeniiberstellung der gemiissigten und tropischen Sipj)en lehrt aber 
auch ohnc wciteres, dass mit der Anpassung von Formen an die extratroi)ischen Zonen 
zuglcicii (lie F m priigung des Ty ])us des H o 1 z ge vviic h ses in den des Krautes 
vor sich ging, somit let/, tere als die j linger en Formen die Abkomm- 
1 i n g e \ n 1 1 o 1 /. g e w ii c h s e n s i n d (l. 2 oq). 

Dass (lie Thcorie dcr Abstammung der iibrigen Wuchsformen von der Gestalt des 
troj)is( hen liaunics immer mchr an l>oden gewinnt, geht zum Beispiel aus dem Buche 
von j'.cws hers or: "The mcgatherm hygrophilous forest of the tropics is probably the 
most an( ient t\ pc of habitat . . . and most recent of all (life-forms) is the annual type" 
(II. 321). 

The investigation of Ji:ki"RKV and ToKRKV [144) led to the conclusion that 
"the origin of the herbaceous type in dicotyledons is from woody or arboreal 
forms", and S 1 1 x K\\ i:i.L, in his monograph of Chacnaciis [2^g\ Compositae, 33 
sp.), states that, "within a family or genus, woody perennial species are more 
j)rimitive than herbaceous annual s|)ecies '. An evolution in the reversed direction 
is, however, postulated by HlKl iix.sox [140. I. 4) in the Polycarpicae. He regards 
the herbaceous Kanunculaccae as the most j)rimitive; a woody structure is, in this 
order, secondarw The current oj)inion is that the woody Magnoliaceae are among 
the oldest living angiosperms. 

The rosette trees. '\\\(i common t\'|)e 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 361 

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, j^j. I. ii), 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 (Grias), Meliaceae (Ca- 
rapa), Rutaceae [Spatkelia], 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 Piiya Rainiondii in Bromeliaceae. Hapaxanthic trees are few; 
Krause [162) called attention to the interesting rutaceous Sohnreya 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. Campa7iula Vidalii on the Azores, MusscJiia on Ma- 
deira, species of Aeoniuni, Dracaena, EcJiiiwi, Melanoselinum, Sonchus, Sinapide7i- 
dru7n, etc.). I 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. ScillMPER 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 [22^), and 
recent observations in the field have not made me alter my opinion. 

Ab:se)ice 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 o{ open water are an absolute obstacle and their absence 
is regardeil 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, 
graduall}' 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 b)' 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 multi{)ly unrestrainedly in virgin surroundings, their 
ravages would perhaps prove catastrophal to the native vegetation and, as a con- 
secjuence 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 herbixorous 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 
specuhitions. Xo 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 geograj)hical history of islands and 
archi|)elagc)es. Hut wc 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. 

-Xative reptiles and amphibians are also absent. CamI'BKIJ. (./^) mentions in 
j)assing that tiicre arc 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 j)erha])s later. That there are no frogs or toads would indicate, 
Cami'I'.kij, thinks, that the archi])elago became isolated before the modern kinds 
of these animals had been developed. lUit, as MlMFORD points out, many con- 
tinental islands lack all lower vertebrates (/cV-,\ 24<S). 

1 low can we exj)lain all these i)eculiarities in oceanic islands.^ HooKKR's answer 
was this: 

Thus, according to the hyi)()thc,sis of trans-oceanic nn'gration, and the theory of the 
derivative orij^nn of spec ics, wc can mulcrstand why the ancient tyj)es . . . should have 
survi\c(l on tlie islands to wliic h hut few of the superior race had j)enetrated ;• — we can 
understand how it comes about, that so many continental species and genera are represented 
on the island by similar hut not identical sj)ecies 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. ii). 

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 [2^8. 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 Wallace 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 
of 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 (/rtV. 11-12). 

As we have seen already, Hookkr's first point when summarizing the charac- 
teristics of island floras was that also their most peculiar genera had their origin 
in a HKUiier continent, that no oceanic island had been a centre for special creative 
forces. (iklSKliACll (II. 494) may have been the first to express another opinion. 
Referring to the systematically isolated endemics in St. Melena, he argued that they 
never had iiad any allies elsewhere: 

Na( h diesen 'I'hatsachen kann von einem Stammkontinent weder in dem Sinne die 
Kcde sein, class die Flora von daher durch natiirliche Einvvanderungen bereichert wurde, 
noch als ob die endemischen Arten aus Umbildungen von Pflanzen hervorgegangen wiiren, 
die in einer tViiheren Periode von auswiirts dahin gelangten. Nur von gewissen klima- 
tischen .\nalogieen ist ihrc Organisation der Ausdruck. S. Helena verhalt sich demnach 
ganz verschieden von den Kap-\'erden und liefert den voUgUltigen Beweis, dassdie Entste- 
hung der Pflanzen auf Inseln, cbenso wohl wie auf Kontinenten, unabhiingig von anderen 
W'getationscentren mogliih war. W'arum sollte auch der geographische Umfang eines 
(".ebiets auf die Kriiftc, welche die Organisationen erzeugt haben, von Einfluss sein? In 
dem kleinsten Raunie, wie im grossten, konnten sie in besonderer Weise sich entfalten: 
nur werden sie im erstercn Palle weniger zahlreich sein miissen. 

In his discussion of the Galapagos flora he admits that what he terms vicarious 
species, endemic in tiie islands, may have arisen through transformation of American 
species, but he doubts it. 

Diejcnigen, welche annehmen, dass die vikariierenden Arten aus Umbildungen von 
eingcwanderten hervorgegangen sind, konnen auch unter den endemischen (iewiichsen 
der (ialaj)agos lieispiele genug anfiihren, dass eine nahe Verwandtschaft dieselben mit 
Ainerika, als ihrem vorausgesetzten Stammkontinent, verbinde. Aber allgemein liisst sich 
dieser ( lesichtspunkt niclit durchfiihren. (ierade unter den geselligen Holzgewiichsen , . . 
finden wir die eigentiimlichsten Erzeugnisse, die Scalesien und andere Synanthereen, die 
nach ihrer systematischen Stelhmg dem Festlande ebenso fremdartig gegeniiber stehen 
wie (he Lobeliaceen des Sandwich-Archipels. . . . Alle P>eredsamkeit, womit die Abstam- 
uiung flcr Vegetation ozeanischer Inseln von den Kontinenten vertheidigt zu werden 
ptlegt, kann die Thatsache nicht verdunkeln, dass in solchen Fallen [the genus Scalesia 
et( . arc mentioned die Organisationen nicht anzugeben sind, aus deren Variation man 
sie hervorgegangen \orstellen mochte. Die nahe Verwandtschaft hingegen, welche zwischen 
vielen endemischen Frzeugnissen des Archipels und denen der amerikanischen Floren 
unleugbar besteht, kann aus dem Pildungsgesetz der riiumlichen Analogien ebenso wohl, 
als aus ei?iem genctis( hen /usanmienhang abgeleitet werden. Und warum sollte iiberhaupt 
das Festland vor den Inseln den \'orzug selbstiindig entstandener Organisationen gehabt 
haben, deren crste Erzeugung in den friihesten Perioden der Erdgeschichte jeder Moglich- 
keit einer \'ariation vorausging? waruin sollte sich nicht spiiter und an verschiedenen 
( )rten sich wiederholt haben, was urs])riingli( :h moglich war und wovon nur die Bedin- 
gungen ein noch ungelostes Kiithsel geblieben sind? (II. 512—513). 

Such ideas have little more than historical interest. Drudk, another leading 
authority on plant geograph)-, was more in accordance with modern thought: 

. . . dass die Flora der Inseln nicht nur als Transformationen der jetzt lebenden Kon- 
tinentaltloren erfasst werden darf, sondern dass auf vielen Inseln unzweifelhaft eine Weiter- 



DERIVATION OF THE FLORA AND FAUNA 365 

entwickelung alter, vielleicht den Charakter einer alteren tertiaren Periode repriisentieren- 
der Stammfloren stattgefunden hat, welche sich hier im Schiitze der Abgeschiedenheit 
fern von dem Einfluss kontinentaler Umwalzungserscheiniingen sich erhalten konnten 

[jos. 128). 

Drude agreed with Hooker that extinct species, known as fossils in Europe, 
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 (jo^. 252) expressed this very clearly: 

Dass diese eingewanderten Elemente infolge ihrer Isolierung eigene Entwickelungs- 
richtungen einschliigen, die sich in einer oft auftallend 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 Acacia, 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 ])lace, 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 (i.e.). 



366 



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 {)recijMtation be very variable ... if the soil be rich, but variable in porosity; if 
the j)lant 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. 

'i'he 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. 6i8). 

What .Vndki-.ws 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 
inimidit)' and hardly anything we can call soil, we need water before even the 
most j)rimitive 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 ma)' 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 Asplcniuui is the only living thing 
observed on the far-fiung 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, 
otter 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 Andki.ws sa\s. In order to start an evolution of new species and genera 
a ver\' great vertical relief, a variable preci[)itation and a rich soil are the conditions, 
l3ut in oriler t(; get a rich soil cover we must have a closed vegetation cover, also 
forest. Wa I.I.AC!; [2yS. 295), sj)eaking 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.' Uas 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. \)o 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 {lyg. 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 {2^8) 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 
lia\e 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 t\'pes. The living world in Hawaii is older than the rock — in a way, he 
sa)-s. ( )uite true, but it is not true that this possibility has, as he says, been entirely 
o\erl()oke(l in previous discussions; I think that 1 have, on repeated occasions, 
exj)ressed myself ver\' clearly on this point, even if 1 do not agree with ZIMMERMAN 
when it comes to exj)lain why and how it happened. 

The biota as we know it today is in part the ultimate product of a progressional 
devclopnient which has moved and evolved along great insular archipelagos over periods 
of time much longer than the ages rec^uired for the development of the main Hawaiian 
Islands and their contemporary biota. Various genera and stem forms of groups of species 
iiKu have evolved in islands — now atolls such as some of the leeward Hawaiian chain, 
the great Micronesian archi])elagos, 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 ])art develoj)ed by unique methods and in which are preserved 
forms whicli are the end ])roducts of species chains that carry back, through archi- 
])elagos 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 j)crceptible change. 

Zimmi:rm.\x does not hesitate to conjure up all the sunk archipelagos he needs, 
if only land connections arc 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 
(lues 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- 
l)clagos marked by (lusters or coral reefs. Surviving lines of middle Tertiary and of 
|)erhaj)s even older continental faunas nuiy have had their germ plasm filtered down 
through su( cessivcly 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 
ot time while tlieir continental j)rogenitors 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. V.ven 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.-Achaiinellinae. 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 (^218) 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 



SKOTTSBERG 



This is pure Lamarckian langua^^^c. \Vc may say, of course, that change of environ- 
ment may cause mutations, but our experience tends to prove that only a fraction 
of I °o are \aluable, the remainder, if not deleterious, at least indifferent. Mutations, 
gene losses. Inbridization, polyploidy and so forth, all may have their share, but 
they are only ripples on the surface. Xo theory has been able to penetrate to 
the nucleus of the j)r()blem. 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 
s|)eculati()n 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 (iS'6). 

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 recjuired an enormously long time to produce the present status, that from 
the Cretaceous until now nothing of a revolutionary character had happened, families, 
genera antl 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. 

l^'inally I shall quote some selected passages from Stebhixs' book on variation 
and evolution Uiy). 

The ditterentiation of orders and families of flowering plants through the action of 
natural selection under ])resent conditions is well-nigh impossible. . . . All the trends 
leading to the difterentiation of families of flowering plants probably took place simul- 
taneously and at a relatively early stage of angiosperm evolution. For instance, both 
distributional and ])aleontological evidence indicates that the Com])Ositae, the most highly 
sj)e( iali/.ed family of dicotyledons, already existed in the latter part of the Cretaceous 
])erio<l, and distributional evidence indicates a similar age for the most advanced families 
of niono( otyledons, the Orchidaceae and Cramineae. It is likely, therefore, that the 
major part of angios])erm evolution, involving the ])rincipal trends in the modification 
of the flowers, took i)lace during the Meso/.oic era (pp. 501-502). 

The g\-nmosj)erms 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 — angios|)ermic fossils are found in greater quantity from 
younger Cretaceous and then in many surprisingly modern types, l^^vidence is strong. 
Si KP.I'.INS has found, that evolution was rapid during one period and slower during 
another. At the end f)f the Cretaceous and the beginning of the l^Locene the number 
of modern types increased rapidly. In eocene deposits in North America and Eurasia 
"the majority of the s[)ecies 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 37I 

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 Eriogonum, Ceanoihus, 
Arctostaphylos, and various Conipositae'' (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 Stipa or Piptochaetium, an evidence 
from palaeontology which strengthens the theory of SiNNOT'r and Bailey already 
referred to. 

Isolation 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 {131, 132) 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 (^^^). The continental flora inherited 
by a Great Hawaii included numerous populations gradually taking possession of 
Ihe 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 



372 C. SKOTTSBERG 

done to examine the <^enoty})ical constitution of this astonishing assemblage of 
\oca\ forms; some may, for ail 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 
tlie majority t)f 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 W'ai.i.ack 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), 
I lad 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. 

Joiiow (/jo) based his opinion on Wallace; when discussing the dispersal 
agencies and the morphology of the diaspores his starting-point was /wo islands, 
Masatierra })lus Santa Clara and Masafuera, which has risen separately from the 
deef) 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 omnij)rcsent young basalt — ^JoilOW could not know that the interpretation 
of this stratum was false, as later shown by OUENSEL (J02). After his visit to 
the ncs\ enturadas Joiiow modified his opinion; the submarine ridge uniting these 
little islands with Juan T'ernandez had then been discovered: 

Die iiiUer [.'Icicher P.reite mit dem Hafen Caldera und in derselben Entfernung 
vom Kontinent wic Juan I'crnandc/. ^elegene Inselgruppe ist vulkanischen Ursprungs 
und stcilt, wie die von dcin Mitglicd der Expedition, Herrn Chaigncau, ausgeliihrten 
Eotungen crgahcn, die iiher Wasscr befindiichen hochsten (iipfel einer im Ubrigen 
iinterseeisch verlaufenden liergkcttc dar, welcher auch die Inseln der Juan-Fernandez- 
(iru])pe als siidli^ hstc (iiptel angehoren. Aus dem Vergleiche der Eloren und Faunen 
heider .\r( hipelc, \vcl( he trot/, der grossen klimatischen Verschiedenheiten frappante 
X'crwandtschaft autweisen, crgicbt .sich mit zwingender Notwendigkeit die Hypothese, 
dass die zwei Inselgriii)i)en in der Vorzeit mit einander in Landverbindung gestanden 
hahen und dass ihre Isolierung die Folge einer stattgehabten Senkung jener Bergkette 
ist isso- 259). 



DERIVATION OF THE FLORA AND FAUNA 



373 



In this assumption he may be quite right, but it finds, as already pointed 
out by Reiche (20J. 269), httle support in the flora, for the two groups have 
only a single species in common (2^1), 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, SopJiora ''tetraptera\ with winged pods, had drifted with 
the current. Five [Eryngium bupleuroides and sarcophyllum, Apiuin fernandezianum, 
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 (^j-j) 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 tetraptera, was recognized by 
JOHOW and recorded from Chile, Juan Fernandez, Easter Island and New Zealand. 
Genuine tetraptera 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 H. 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 erecta is, however, an endemic species; there are two more on Masa- 
fuera (see 22g, 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 
{199), GouRLAY [301), Burger [41), and Goetsch [124). Plate drew his con- 
clusions from a comparison between the littoral faunas of Juan Fernandez and 
the opposite coast: 



374 



C. SKOTTSBERG 



Die Seichtwasser-Faiina weist deutlicher als die geologischen Verhiiltnisse darauf 
hin, <lass Masatierra in der That als eine ozeanische Insel anzusehen ist, die nie in 
Zusamnienhang niit deni I-"estland gestandcii haben kann . . . (p. 228), 

but this did not prevent liiin from believing that Masatierra and Masafuera once 
formed a sini^le lari^e island, in spite of the distance (92 miles) and the deep 
water that sej)arates them (p. 222). No more will be said here about a connection 
with the mainland; I have already discussed tliis question from an algologist's 
view|)<)int, and with a difterent result (2jS). 

The reason why (ioLRl.AV 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 j)()inted 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 
sj)ecies cjuite without relations not only in Chile, but in all America, he said 
notliing. 

Hi K(;i:k, who had visited the islands, followed JoiJOW; they were typically 
oceanic. After telling us about some of the most remarkable endemics, he exclaims: 

W'er brachte die.sen entlegenen Stiitten solch kostliches Geschenk? Die Stromung, 
die \'(')gel iind vor allem die Winde. Sie beluden sich niit Sporen und Samen. . . . 
selhst voiii I'euerlande empfangenes gedieh. Doch aiis viel weiteren Fernen kamen die 
Kinwandcrer, auch vom tropischen x^merika, Polynesien, ja sogar von Aiistralien 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. 1 shall leave these speculations 
without comment; we have better reason to observe GOETSCIl's paper which, 
in sj)ite of being fairly recent, contains many amazing statements not supported 
by facts. Ignoring OLl'.NSlll/s report on the geology, he tells us that the islands 
rest "auf einem Sockel von grimlichem Andesit, der auch die Hauptmasse der 
Anden bildet". lie had seen my writings on the flora; the use he made of them 
mav l)e illustrated by a couple of examples. Among the plants introduced from 
luirope he mentions "Aromo aus Castilien" — this is a local name for the en- 
demic Arjai-a fcriiaiideziaiuA Me rejects all land connections, the sandal-wood 
came from the Past Indies; he had not observed that it belongs to a quite dif- 
ferent section than SautaliiDi albiDu. The arborescent Conipositae and Plaiitago had 
originated in the islands, '' Plaiitago feriiaiidezia wx^iX Skottsbcrgii, \-2 v[\\\oz\\vc\\\. 
20 cm langen und 3 ^2 cm breiten, an der Spitze des Stammes stehenden Blat- 
tern' . lie overlooked that the so-called /'. Skoitsbergi'i is a modest annual and 
a form of the common Chilean /rianata, a variable species. But GoETSCII knows 
what happened in the islands: "Dass der europiiische Wegerich auf Juan F'ernandez 
mcterhohe HUitenahren und fusslange Blatter tragt, weist auf die P^ntwickelung 
hin, die seine X'erwandten einstmal nahmen" (p. 29). This refers to P. lanceolala., 



DERIVATION OF THE FLORA AND FAUNA 375 

which luxuriates in the islands, but cannot prove that the arborescent P. fer7ian- 
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, I cannot tell, unless Joiiow'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 konnen. 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 nordlicher 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 GOOU says {log. 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 GOETSCli 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 
Meeresstromung entgegenwirkt, der sogenannte Nino, 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 (joo) — 
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 F'ernandez, but he does not 
appear to have profited very much by his visit. 



376 



C. SKOTTSBERG 



Where HkANClll [^^oj) got his idea of the island flora I cannot tell, in any 
case not from facts, in spite of having visited Masatierra: 

Coiiio curiosidad cientifica se piiede decir que Masatierra no seria sino una de 
las tantas vetas del gran continente sumergido en el Oceano Pacifico en epocas en 
(jue el continente sudamericano era un lecho de mar, y per consiguiente la isla serfa 
mas vieja (pie el suelo de la madre patria. Como comprobaci6n estil la ausencia de 
los \olcanes, los esarpados farallones que muestran las capas geol6gicas, el desliza- 
miento de las rocas de la epoca glacial, y sobre todo la flora unica, di versa del conti- 
nente y (juc i)uede solo acercarse a la flora polinesica y australiana (p. 14). 

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 (<?) 
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 I'^ernandez had been isolated since late Cretaceous times, so 
that onl}' 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- 
(iruppe einen der Ozeanis der siidpazifischen Landbriicke. Das beweisen nicht sicher 
die \'6gel . . . auch nicht die Schmetterlinge, von denen die Nymphaliden durch die 
chilenische J^ranieis carye vertreten sind, die Pyraliden durch Scoparia ragonoti, 
ncben denen noch weitere chilenische Formen genannt werden. Alle hiitten auch trans- 
marin die Inseln erreichen konnen. Wichtiger sind die Landschnecken, deren Gattungen 
fast alle auch in Chile vorkommen. Vertreten sind z. B. die Succineiden und Ferrusa- 
ciden, sowie die Kndodontiden. Diese sind hier mit Aniphidoxa vertreten. Deren typische 
Cntergattung ist auf Juan Fernandez endemisch. Stephatwda besitzt 6 endemische Arten 
aut den Inseln, andere auf Chiloe, in Chile, Patagonien, Feuerland, Argentinien, Para- 
guay, Brasilien und eine auch auf Kcrguelen. Dagegen gehort die auf Juan Fernandez 
endemische J^'ernandezia zu den sonst hawaiischen Amastriden, die auf Hawaii auch 
ihre ein/igen niiheren Verwandten in den Achatinellidcn besitzen. Hier kann nur eine 
Au>l)rcitung iiber Land angenommen werden. Von anderen Landtieren findet sich auf 
Juan Fernandez v.. W. noch der 'J'auscndfiissler Gecphilus latkollis. Endlich sind sie auch 
von cinem ( )ligo( haeten erreicht worden, der Ocnerodrilinen Kerria saltensis, ^\^ dMQ\\ 
in Chile vorkommt. Hier wiire eine jiingere, iiberseeische Einwanderung denkbar. Aber 
Kc) ) id ist au< h alt genug, dass sie auf dem Landwege nach Juan Fernandez gekommen 
scin kann. Auch die Flora der Inseln zeigt interessante Beziehungen (])p. 324-325). 

As could be expected, Arldt's south Pacific bridge was supported mainly by 
tile land molluscs, and they made him extend his bridge across the equator to 
ilawaii. There is in the Juan P'crnandez flora some perplexing affinities with 
Hawaii that scctn to defy all attempts to an explanation. The occurrence of two (not 
only one) endemic species of XcsogeopJiilus (formerly a subgenus of 6'r^////7//'j) is in- 
teresting, Kcrria is, j)erhaps, 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 Xi'sophilaoiioji, 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 F'ernandez and blaster Island" had escaped his notice. The leech would, 
I sup|)ose, have furnished him with one of the most eloquent evidences of 
former land connections. Xor did he know the botanical volume of this work, 



DERIVATION OF THE FLORA AND FAUNA 377 

for he gives a summary of my 19 14 paper (22^). He agrees on the great age 
of the "Palaeopacilic element"; second comes the small 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 VII — will now 
be answered. We have seen that endemism is very high among the phanerogams 
and that the various kinds distinguished by HoOKER are represented; we have 
even a primitive endemic family, Lactoridaceae, we have a proportionately large 
number of peculiar genera, some of them quite isolated, particularly among the 
Compositae {Centaur odendro7i, Dendroseris, Hesperosei'is, Phoenicoseris, Rea, Rhe- 
tinodendron^ Robinsonia, SympJiyochaeta, Yunqueci) but also in other families, 
Cuniiiiia (Labiatae), Juania (Palmae), Megalacfme and Podophorns (Gramineae); 
further there are a few endemic genera closely related to South American ones, 
Nothomyrcia (Myrtaceae), OcJiagavia (Bromeliaceae), and Selkirkia (Boraginaceae). 
As among the genera, so we find species of a strong character in Chenopodium, 
Coprosma, Eryngium, EupJirasia, Fagara, Peperomia, Plaiitago, Sanialum, Urtka, 
Wahleiibergia, well-marked but not very aberrant species in Berberis, Boehmeria, 
Car ex, Cladium, Colletia, Erjgeron, Escallonia, Gunner a, Halorrhagis, Hespero- 
greigia, Ranunculus, R/iaphitkaninus, Solanum and Ugni, and many not very 
different from their continental congeners in Abrotanella, Acaena, Apium, Azara, 
Cardami7te, Cliusquea, Drimys, Dysopsis, Galium, Luzula, Margyricarpus, Myrc- 
eugenia, Perneitya, Pkrygilanthus, Sopkora, 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 [Thyrsopteris] 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 [Sopkora] with two species, 
and no orchids. The proportion genus : species is i : 1.65. Mammals, batrachians, 
reptiles and fresh-water fishes are absent. The earth-worms are supposed to be 
adventitious with the possible exception of Kerria 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 VL 
Percentage of \v()ocl\- and licrbaceous species in Juan Fernandez. 



Indi.utMious species .... 

Not eiitleniic 

ICndeinie, ^enus not eiuleniic . . 71 
Als;) tlie <'enus endemic . . 



Total 


\N'()()( 


ly% 


Herbaceous % 


147 


68 


46.3 


79 


1 
53-7 


46 


5 


10.9 


41* 


89.1 


71 


37 


52.1 


34* 


47-9 


30 


27 


90.0 


3 


10. 



Four species suffruticose. 



comparison, and the Juan hVrnandez Islands were regarded as supporting their 
h>-|)othesis; this, to judge from tlie table, they certainly do. The five woody 
non-endemic species are liiiipctnDu nibnun and Saliconiia fruticosa (low, erect 
shrubs), and Myrtcola unnimularia, Riibiis gcoidcs and Calystegia tuguriorimi 
(trailing). 

The rosette tree form is observed in 16 genera with together 31 species, 
belonging to si.x families, Boraginaceae, Bromeliaceae, Chenopodiaceae, Compositae, 
Plantaginaceae and Umbelliferae; the palm Juania is of course excluded. As I 
have paid special attention to them in another paper (-^J/), where they were 
well illustrated, no more will be said here. 

Annual and biennial herbs, the therophytes of Raunklkr, are not completely 
lacking, but they are ver\' few: Cardamiiie chenopodiifolia, C/iae/otropis (2 species), 
Par'utaria, Plauiaoo tyuncata, I'etrao^oma and Urtica Masafuerae. In 2^1 I listed 
Cliartntropis among the hemicryj)tophytes; they give the impression of lasting 
more than one year antl are found green at all seasons, and the same may be 
true of Parictaria. I am not at all sure that PUxutago iruncala is native. The 
Caydanu}ic has been seen twice, last time in 1872, the Crtica not since 1854, 
when it was discoxercd. 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 exj)lained otherwise. iMulemism of a very high degree within a small 
area is no monopoly of isolated islands; it will be sufficient to mention the Cape 
fiorii or southwestern Australia. The large proj)ortion of woody plants can be 
understood if the islands became isolated before the myriads of herbs, particularly 
the annuals of Central Chile, had evolved. (iRlSKr,.\(ll's "Clavija" and related 
life forms are not confuied to oceanic islands; if my interpretation of this mor- 
phological t\'pe as an evidence of anti(juity is correct, we can understand why 
it takes such a pronuncnt part in old island fioras. The reason why therophytes 
are almost wanting is not climatic. This is amply |)roved by the innumerable 
anniuil weeds introduced with the traffic and thriving only too well. The climate 
is of a modified Mediterranean tyj)e, and from a purely climatic viewpoint we 
should ex{)ect a large percentage of native annuals and biennials; this question 
was discussed at some length in ^-r/. 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 role. 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 litde 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 
successfufly to get transported across the sea are very doubtful or, as far as we 
can see, none at all. 



38o 



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 X 
The Chilean coast line and the history of the Andes. 

I^^rom a look at the map we easily get the impression that the trend of the 
South American west coast is a j)roduct 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 ex{)osed 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. 

flic Cliaigticau Ridge. — The two archipelagoes lie within the 2000 m line, 
San I'elix and San Ambrosio on a plateau rising above the 400 m curve and 
extending a long way toward Juan Fernandez, as seen from CllAlGNEAU's table 
( -T^) 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 Meniivn Ridge. — 160 miles NW. of San Felix-San Ambrosio another 
ridge, called the Mcrriam Ridge, was discovered during the U.S. "Carnegie" 
campaign 1928-29 (//). It extends between 25° 3'. 2 S., 82° 20' W., and 24° 54' S., 
"^i" 13' \\'.; 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 i 1 86, 1188 and 1168 m were found. From the latter spot a series 
of soundings was taken SIv 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 
W'XW. the sloj)e is more gradual until a depth of 3000 m is reached. 

North of the Merriam ridge, in 21^40' S., 8i°4o' W., approximately, a sudden 
rise, bounded by the 2000 m curve and surrounded by deep water, has been 
discovered (see map). Mere the bottom rises to 972 m below the surface. I do 
not know if this remarkable j)lace has a name. 

The Caniegie Kidfi^e. — During the cruise of the "Carnegie" two soundings 
about 100 miles (jff the coast of Fcuador, lat. l°32' S., long. 82° 16', gave I 5 1 5 
and 1454 m, respective!)-, 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, jirobably 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 22y. 44, 45, show 



DERIVATION OF THE FLORA AND FAUNA 



381 



Table VII . 

The Chaigneau Ridge. 
r = rock, s = sand. 



Depth in 
m 


Bottom 


S. lat. 


W. long. 


Remarks 


38 


. — . 


26° 14' 


79° 56' 


14 km NE. San Felix 




80 


— 


26° 15' 


80° 4'.5 


4.5 km N. San Felix 




69 


— 


26° i7'.5 


80° 8'.5 


4 km W. San Felix 




179 


— 


26° i8'.5 


80° 8' 


5 km W. San Felix 




220 


r 


26° i8'.5 


79° 57' 


Between San Felix and San Ambrosio 




400 


s 


26° i8'.5 


79° 49'.3 


5 km NE. San Ambrosio 




250 


s 


26° 19' 


79° 52' 


3 km N. San Ambrosio 




150 


s 


26° 19' 


79° 51' 


3 km N. San Ambrosio 




215 


s 


26° 19' 


79° 49'.2 


4.5 km NE. San Ambrosio 




182 


— 


26° 25' 


79° 56'.5 


6 km S. San Ambrosio 




148 


— 


26° 26' 


79° 52' 


8 km S. San Ambrosio 




105 


s 


26° 30' 


79° 49' 


15.5 km S. San Ambrosio 




400 




26° 36'.5 


80° 8'.5 


33 km S. San Felix 




550 




27° 49'.5 


80° 14' 


c. 165 km S. San Felix 




675 




28° 3'.5 


80° 16' 


c. 200 km S. San Felix 




660 




28°33'.5 


80° 11' 


c. 260 km S. San Felix 




660 




29° 14' 


80° 15' 


c. 340 km S. San Felix 




1300 




30° 49' 


80° 24'.3 






1430 




31° 26' 


80° 10' 






1800 


— 


32° 45'.5 


80° 18' 


Between Masatierra and Masafuera, about i 


° N. 


1800 


— 


34° 34' 


80° 36' 


About 100 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 <-"• SKOTTSBERG 

the deep trench following the trend of the Andes, is a consequence of the gigan- 
tic mountain-builchng processes, tlie submarine ridges west of the trench can- 
not hel|> to get iniphcated. Tiiis idea is by no means new, it has been expressed 
by many: "the width of Soutli America may well be a good deal less now than 
before the Andes were uplifted", as G()(JI) says (lop. 349), but some of these 
writers did not tr\' to penetrate the complicated geographical-geological history 
of til is region of great tectonic disturbance. This is true of myself, when I tried 
to describe what I imagined having occurred (22/. 43), but it does not apply to 
Ikms(IIi:k (//i') who took pains to inform himself of the history of the Andes 
as told b\' 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 Tacihc land mass of hypothetical width; one opinion regarded 
the Coast Range as belonging to this land. Irmscher, who took \\T:c;ener's 
sitle. 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 sufticient to account for uplift and folding. Consequently, he was 
unwilling to accept Pexck'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, perha{)s, had been connected with California (p. 45). 

l'i.N( K stellte (lie Aiisbildiint( dor ozeanischen Tiefen am Rande des sudamerika- 
nischen Kontincntcs, also das \'ersinken angrenzender 'j'eile des Pazifiks, der aufwarts 
bewegten andinen Scholle gegeniiber und schliesst, dass die niichstliegende Erklarung 
liir die \'olumenanderungen unter der festen Kruste in dort stattfindenden Massenver- 
schicbungen zii suchcn ist. Wenn Massen aus der pazifischen Region in die andine 
iihertrcten, so nuiss in crstcrer 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: 

i)ie ine( hanis( hen Ursachcn des l-altenvorganges sind zweifellos anderer Natur, und 
(lie .Magniaintnision ist genau so eine Uirkung derselben wie die P'allung. Denn es steht 
test, dass die ( ichirgshildung niit dem Kmpordringen des Magmas synchron ist und die 
Intrusion somit gleic hzcitig niit der angcnommenen annuihlichen Lostrennung Siidamerikas 
von .Atrika. 

Where does the (■iiaigneau-Merriam bank come in.- \Ve(;ener left it unexplained, 
according to 1)1 Tori it was an "advance foltl". 

l^EKRV and SiN(;i:w.\i,i) (j^), in their review of the tectonic history of South 
America, describe the develoj)ment in the following terms. 

Some students regard the Cordillera de la Costa in Chile as remnants of an an- 
( ient massif, the bulk of \\hi( h has been downfaultcd beneath the waters of the Pa- 
cific. It (onsists of (rystalline ro( ks both igneous and metamorphic and these have 
(onimonly been assumed to be of great age — even Archean. This inference of antiquity 
rests, not upon their known rehitions, but upon the geosynclinal nature of the Andean 
seas which are clearly ej)icontinental and not shelf seas. 



DERIVATION OF THE FLORA AND FAUNA 383 

The sediments in this geosynchne are compressed between the Brazihan 
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 (jji). To begin with I shall allow myself to quote Florin's summary 
(^5. 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 Concepci6n-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 
Concepci6n Series is overlain by the deposits of the marine Navidad Series, which 
is upper Oligocene or Lower Miocene in age. Bruggen came to the conclusion that 
the Concepci(3n 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 Concepcion 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 BrCggen 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 



284 C. SKOTTSBERG 

disappeared under water to form the long series of the Patagonian channels, the 
Andean valle\-s 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 perioils of glaciation must, however, also be taken into account. 

The development of the coastal region, as told by BrCggen 1950, is ex- 
})lained 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. 

HkiciCKN 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 Maulh'n (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 PY^nandez Islands and, farther north, San Ambrosio and San Felix. 
Taking the 2000 m cur\e 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 tind in the Tertiary deposits of Parga and other places in the 
zone north of Rio Maulh'n. 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 I^rCggen's map, prob- 
ably copietl from Si pax. 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 Xavidad series south to 45°, and this was, as we have 
heard, referred to Upper Oligocene or Power Miocene. Also after the separation 
the Juan Fernandez land continued to exist until finally, presumably with the late 
Tertiary uplift of the yVikIcs, 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 h'elix^ the islands are young, probably Pliocene, 

( iiando cxistt'a todavia im resto de la antigua Tierra de Juan Fernandez, de la cual 
inm/xro la flora del lioccno? Ciiando mas tarde se hundi6 tambien este resto, sobre- 
salian solanicntc las i)artc's volcanicas, constituyendo las islas actuales de Juan Fernandez, 
(lue Servian j)ara rcfugif) de la flora. 

This is the process as I have described it (.^27. 43) and BrCgcjKN is of the 
same oj)inion from the geologist's viewpoint. And if we go back to Hooker's 
lecture, we shall [wmX that the same idea, ai)i)lied to a different region, was famil- 
iar with him. 

1 This refers to the siil,>inarinc eruption in 1835 "^<ir die coast of Masatierra; whether 
any signs are left I cannot tell, for tlie place has not been sounded. This eruption was simulta- 
neous with an (Mrthi|uake in ("oik epcion. I'ossilily there was a connection with the tsunami of 
\'allenar 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 &c, &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. 1 1). 

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 Xicotiaiia [ii2)\ it was, partly at least, based on WEEKS' paper 
{2jg). 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 



^Sb <-• SKOTTSBERG 

Chapter XI. 

The Tertiary floras of Chile and Patagonia. 

'rcrtiar\' 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, 37°-37°30' 
s. hit., in combination with the coal seams of Lota, Lebu etc., and east of Lake 
Xahuclluiapi on the Argentine side. 

I'lu- C 'o)iLt'pcio)i-Ayauco series. — The fossil flora was examined by ILngkliiardt, 
who described more than one hundred species of angiosperms, and later by BERRY 
\2j]. The determinations are, just as in all the other cases, based on leaf impres- 
sions, but although l^l'.KKV found that KxoKLllARDT's determinations "are usually 
to be relied upon" ({x 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, 
Hignoniaceae, Bombacaceac, Caesalpiniaceae, Combretaceae, Erythroxylaceae, Lau- 
raceae, Lec\'thi(.laceae, Myrtaceae, Palmae, Sapindaceae, Styracaceae, Vochysiaceae, 
and so forth. In general character the flora approaches that of the Amazon basin, 
e.xtended, 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 
eleiuents of the flora of Central Chile" (p. 106) — Cassia is, however, one of the 
genera mentioned, another is Myraugcuia, represented by several species in cen- 
tral and south Chile (to know Myrtaceae without flower and fruit is well-nigh 
impossible). Hl'kKV referred the flora to the Lower Miocene, "it is surely not so 
okl as lv)cene", and also younger than the XotJiofagus flora in the Magellanian 
zone (j). 115); this he considered to be of Low^er Oligocene age. 

There are two gynuiosperm genera in Berrv's list which seem to disturb 
the imj)ression of an otherwise homogeneous neotropical assemblage, Araucaria 
and St<//a'i(f. The material was reexamined by FLORIN (pj) who showed that 
Araiuana ara!/i(>r>fsis Berry is a species of Podocarpus, and Sequoia cJiilensis 
Fngclhardt p. p. another. These, together with a fern described by HallE (Lygo- 
({n())i, ;i2] are important additions to the Arauco flora. F^LORIN, in accordance 
with l^KrciCl.N, ref(,'rs it to the I'>)cene and characterizes it as follows. 

The ( oinpo.sition of the fossil c:onifer vegetation, and the distributional aspects, of 
it> < onstitucnts, iii(!i( ;itc that it derives from a \varni-teiuj)crate or sul)troj)ical rain-forest, 
more particularly a lowland i)od()c.arj)-evcrgreen dicotylous broad-leaved tree forest, 
growint,' on tin- ( oastal plain or j)erhaps partly on low hills not far from the coast. The 
( limatc was ])r()hal)ly ( harac tcri/.ed hy great humidity and rather uniform temperature, 
it was trostless, and wanner than the i)rescnt climate of the same district (p. 26). 

The possibility that plant material from the uplands had been carried down 
and become nnxcd with material frf)m the coastal plain is contradicted by the 
state ol preservation which is the same in all cases (p. 26). 

'I he rhiiileufu flora. — The fossiliferous beds of Rio Pichileufu are situated 
in 41 s. lat. about 30 miles east of Lake .Xahuelhuapi in a treeless steppe country. 



DERIVATION OF THE FLORA AND FAUNA 387 

Berry {2^) 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 Araucaria pichileufuensis, Fiizroya tertiaria, Libocedrus prechilensis and 2 
Podocarpus\ further, there is a species of Zaniia and Ginkgo patagonica, and of 
ferns 3 species, one of them a Dicksonia. Araucaria and Libocedrtis are, according to 
Florin, correctly named (pj), Ginkgo should be called Ginkgoites\ Fitzroya belongs 
to Podocarpus, and one of the Podocarpus sp. belongs to Acmopyle of PiLGER. 

There is no trace of Fagaceae and Berry referred the flora to Low^er 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 Pichileufu. Berry [zf] pointed out that the present South 
Chilean rain forest flora is not represented in the Arauco flora whereas the 
Pichileufu beds contain such Chilean genera as Azara, Berberis, Maytenus and 
Myrceugenia and, in addition, the following Antarcto-tertiary genera: Drimys, 
Embothrium, Eucrypliia, Laurelia, Libocedrus and Lontatia — 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 Araucaria-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. Araucaria and Nothofagus are absent, the only conifer 
found, Fitzroya 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 Laurelia and Peumus 
may be mentioned. The age is early Miocene according to BERRY (jj^). Eocene 
according to Frenguelli [337), the climate warm temperate. 

Berry {313) regarded all the fossil floras containing an abundance of Faga- 
ceae [Nothofagus, according to DUSEN also Fagus, which is questionable) as of 
approximately the same age and older than the Concepcion-Arauco series. 

The Nirihuao flora. — Three localities close together on Nirihuao river near 
Lake Nahuelhuapi. Some ferns, among them Alsophila australis, also known from 
Seymour Island, further Zamia, Araucaria Nathorstii, Fagus (?) and one species 



388 C. SKOTTSBERG 

o{ Xo/Ziofcii^iis. A^e proposed b\' HKkRV (jJ4) Lowest Miocene or Upper Oligocene, 
then \-oun<^er than the Ma^allancs flora. 

r/if Mdi^iil/cvns fiord. — DusKX [yQ) distin^uislied two plant-bearing horizons, 
an upper Ayaucafia horizon and a lower Xothofagus horizon. Of the remaining 
dicots'lons genera none were identified with living ones; DUSEX preferred to call 
them liscalliuiiipJirUiDii. I lydnvioeiphylliDu etc. There is no obviously tropical 
eletnent, it is a temperate flora. DlSKX regarded the two horizons as distinctly 
different in age, dating the upper to Lower ^Miocene, the lower to, perhaps, 
Oligocene. Hkrrv. who doubted the correctness of this distinction, regarded them 
as older than the Concepcion-Arauco flora, which seems improbable. 

riie Snuioin- flora. — To judge from DusKX's description (cVo) the tropical 
element is not conspicuous, whereas the actual South Chilean forest flora is well 
represented: Araucaria (nearly related to A. araiicaua), Drimys, Xotfiofagus., 
L'aldcluria, lAUirelia, I.oDiatia, all supposed to be of Antarctic parentage; lean 
see little reason for I^KKR\'s assertion that the Seymour flora contains "a large 
element of subtropical or warm temperate types like those found to-day in south- 
ern Hrazil" 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 temj^erate 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 Aonopyle [I^liyllites sj)., DlSEx) is of particular interest [338]. 

We have seen that Berry considered the Xotfiofagus beds to be older than 
the Arauco-Pichileufu deposits. y\ll the local fossil floras of Patagonia are, he 
says [333], older than the marine Patagonian transgression and undoubtedly pre- 
Miocene. P'rexcueli,! distinguished three epochs: (a) Late Cretaceous to early 
lv)cene, with a tropical flora, known from the Chali'a beds; (b) an intermediate 
period with subtropical and temperate types [Xotfiofagus) mixed; to this he would, 
1 suppose, refer the Seymour flora; (c) the youngest epoch, Miocene-Pliocene, 
evident!)' extending into Pleistocene: a temperate flora, now ranging along both 
sides of the southern Andes and characterized by the dominance of A^otfiofagus 
and ot a number of conifers. The more or less corresponding development of 
the Andes was according to I5ERRV (2S): (i) I'^ocene-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 HlRRV, the series of glacial and interglacial periods, a 
most important factor of disturbance. 

'I his is tlie background against which we have to discuss the history of the 
Juan I'ernandez 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 hterature, 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, i2y (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 [137) 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 
HOLTEDAIIL says, the existence of a land mass of considerable size where there 
is now sea. Joyce (<5-f) 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 iMacquarie 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 [14.. 183): 

Archijielagoes 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 ])layed 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 q{ Acuwpylc in Patagonia and a sec- 
ond fossil form in the TLocene of Seymour Island, he wrote [33S. 136): 

Fur cine ehemalige Verkniipfung der australischen Region mit der Antarktis spricht 
auch die Verhreitung der Gattung Acmopyle. Dass also die Antarktis in diesem Falle 
als cine 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 lY^riod, fcjr this island was, at least during maximum glaciation, 
entirely ice-covered and must, Tavlor 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. Dahe [^2. 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 39I 

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 
{^iig) 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 Lyallia\ 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 Ayaiicaria, Driniys, Laiwe/ia, 
XoiJiofagus etc. in Tertiary deposits on Seymour Island would lead us to infer 
that they are of Antarctic origin and have radiated from there, but Araucaria 
once iiad a world-wide distribution, Drhuys belongs to an order — Magnoliales — 
of Holarctic range, the same is true of Fagaceae, and even Laurclia is, Berrv 
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. '' Araji- 
caria stands as a per[)etual warning against forgetting that the past is the key 
to the {:)resent", l^KRRV wrote (I.e. 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 GlHHS appears to have been one of the very few experienced phyto- 
gcogra[)hers 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" [106. 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 
(kiinea. The highland of Tasmania, the subject of her survey, had received 
nothing from the south, all the so-called Antarctic plants, genera lika Ahroiajtclla, 
Astdia, C'arp/ia, Colobajiihus, Coprosnia, Driinys, Gaimardia, Gunner a, Lage?io- 
pliora, Xotliofagus, Orcobolus and so on, had come from New Guinea, and from 
there they had radiated to Polynesia, Hawaii, Juan P^ernandez, Tierra del PTiego 
and, I presume, Antarctica. Had she lived to hear of the discovery of A7v//^<7^//i' 
in New (luinea, where more species have been found than anywhere else, and 
in .\ew Caledonia, and of the rich development of the Winteraceae in New Guinea, 
she wf)ul(l have regarded such finds as a forcible proof of the correctness of her 
opinion. 

Many /ooge()gra{)hers have looked with much suspicion at the Antarctic 
continent as a centre of radiation. Sl.Ml'Sox, in his review of the theories involving 
Antarctica in the distribution of vertebrates (^^,?), 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. T^y): 

There is no known bioti( 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 
he exj)ected from the present northern connection known, or with considerably prob- 
ability inferred, to have existed at ai)propriate times in the past. 'J'here 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 Antafctic 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 (2j): 

Nous avons tire de notre etude cette notion importante que la liaison entre I'Aus- 
tralie et I'Amerique a eu lieu par une terre antarctique dont les temoins restent ac- 
tuellement, mais ni par la Nouvelle Caledonie, ni par la Nouvelle Zelande, ni par le 
groupe Samoa-Tonga-Fiji, et, par voie de consequence, qu'elle n'a pas eu lieu par le 
centre du Pacifique (p. 1053), 

Berlioz (2^), with examples off'ered 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 presente avec celles de I'Amerique 
du Sud surtout de la region andine et patagonienne, des affinites aussi etroites que 
curieuses. 

Finally Lindsay (idy), 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. 



394 



C. SKOTTSBERG 



Chapter XIII. 

The history of Juan Fernandez — a tentative sketch. 

In my first [)aper on tlie Botany of Juan Fernandez (22/^), 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: 

Frcilich haben wir keine Ahnung davon, wie schnell Arten oder (lattiingen ent" 
stehen, aber wir konnen uns kaum denkcn, dass in der kurzen Zeit, die seit der Ent" 
stehung der jetzigen Inseln verHossen ist, sich Typen wie Lactoris oder Rohinsojiia aiis 
"Keimen" entwickelten, die nach den Inseln gebracht wiirden, um sich in ungestorter 
Isolierung umzutormcn . . . Ich bin der Meinung, dass das alte Klement nicht auf Masa- 
ticrra oder Masafucra entstand, sondern alter ist als die jetzigen Inseln, und dass es 
wenig wahrschcinlich ist, dass die alten, endemischen (oder andere, eng vervvandte) 
(iattungcn und Arten von Juan Fernandez, noch nachdem die Inseln gebildet waren, 
die vielen vermeintlichen Ursprungsorte bewohnten, und dass Veriinderungen in der 
I'flanzenwelt von Polynesian, Neuseeland, Chile u.s.w. in (puirtiirer Zeit die isolierte Stel- 
lung bewirkt haben. Ich glaube also, dass in vor- und friihtertiarer Zeit grossere Ent- 
wickelungszentra existierten, und dass ihre Flora nunmehr als ein altpazifischer Rest 
fortlebt. Auch eine Resttlora wird sich aus vielen Familien und (iattungen aber ver- 
ba! tnismiissig wenigen Arten zusammensetzen. 

At that time our knowledge of the flora was incomplete; much fresh and new 
material was added in 19 16-17. 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 19 16-17 survey a rather 
strong Antarctic-bicentric group took shape, and in my 1925 sketch of the history 
of the flora (.?.?/) not only was this, but also the Old Pacific plants claimed to 
have "reached Juan F'ernandez 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 P'ernandez" of BrCckien, forming 
a westward extension or lobe of South America, reaching the actual site of Juan 
PY'rnandez 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 
uj)lift of the Andes, the j)resent coast line ought to be recent all along, but this 
is not in accordance with the opinion that the P^ocene Arauco flora was a coast 
flora. The conclusion would be that the great depths were initiated already during 
Cretaceous times, getting dec|)er and deeper with the successive periods of uplift. 
The Juan PY'rnandez land or peninsula formed part of the neotropical P2ocene 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 Pichileufii. As the Concepcion-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 Azai-a 
celastriniformis and tertiaria — 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 corynibosiflora is of still greater interest: 

I have seen leaves of all the South American species and the most similar is Berberis 
corymbosa Hook, et Arn. of Juan Fernandez (i.e. 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 Pichileufu 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 Myrceugenia, 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, Jiiaiiia, Ochagavia, Nothornyrcia and Selkirkia, and species 
of Chusquea, Hesperogreigia, Urtica, Phrygilanthus, Cheiiopodinin, Colletia, Dys- 
vpsis, Ugni, Eryngium, RJiaphithamnus, Solanum and Nicoiiana. 

Centaurodeiidron, Yunquea and the four endemic Cichoriaceous genera stand 
apart. They are montane and we have no clue at all to their history, but w^e 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 
ofl"ered by Ceiitaiirodendron and Yunquea, we are victims of wild speculation. 

Few endemic ferns belong to the neotropical element, Trichomanes bigae, 
Dryopteris maequalifolia, Asple7t2uni macrosorum and stellaium, Pellaea chilensis, 



396 



C. SKOTTSBERG 



Polvpodhini interniednnji and Ophioglossiim ferjiiutdezianmn, but PrZ/^rrt' and OpJiio- 
glossum do not grow in the forest, the former inhabiting the dry coast chffs, the 
hitter tlie open grass kind, 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 il/j'r/^^f^/rt 
)ni))n)nilaria, herbaceous. Myrteola is the only member of this group that extends 
south to the subantarctic zone. Of the others Dantlionia, Koeleria, Stipa, Piptochae- 
tiii))i, lUfocliaris, JuJiciis procerus (also douibeyanus and iiubricatusi), LUiertia, 
Pipiroj}iia fcniandcziaua, Parictaria and Miniulus 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 Stipeiuvi 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 1^rC(;gex 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- 
\idad transgression, separation from Chile occurred, 

. . . cl mar del polo sur entro en comunicacion con el mar que banaba las costas de Chile 
Central y cl primer antecesor de la corriente de Humboldt llevo las aguas inas frescas hacia 
el norte, dando principio a la gran zona desertica. 

HrCG(;kn 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 1 cannot believe that the present steppe-like communities 
date back to early or even middle Tertiary time. On the other hand it seems 
(luite unlikely that the s[)ecies of Stipa, PiptocJiaetiu))i, DantJionia, etc. were in- 
troduced with the traffic while, in this respect, Chaetoiropis (the endemic nature 
of i h. inibcrbis (juestionable), the two Cypcrus, jf uncus capillaceus, Paronychia, 
Ccnlclla. Ilcdyotis and Plantago iruncata are under suspicion. 

Of the ferns found elsewhere Polypodium lanceolalurn is pantropical and old 
enough to have belonged to the ancient flora, and this may be true also of 
7'richouiaues exsccfuiu, Hy))ieiiopJiyllum spp., Adianiuni chilcnse, Ptcris chiknsis and 
scmiad)iata. and lilaplioglossui)i\ 1 have suggested that the latter was carried 
directly to Masatierra by a northerly storm, and this could have been the case 
also with Polypodium Masafucrac, observed a single time lOO years ago and never 
again. P. tric/ioiuauoides remains doubtful in spite of the specimens still extant 
and labelled Juan hY'rnandez {2^(j. 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 juasafucraua, 2 sj). of Spergularia, 2 Berberis, Cardamine Krues- 



DERIVATION OF THE FLORA AND FAUNA 397 

selii, Galium masafuerajtuni and 6 Erigeron. The Boreal character of these genera 
is recognized. This element extended south along the precursors of the late Tertiary 
mountains; the species of Agrostisi^^ Spergularia, Berberis, Cardamine and Ga- 
lium have their closest relatives either in the tropical Andes [Berberis] or in Chile. 
Erigerofi 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: E.fruticosus and its cognate luteoviridis, the three 
herbaceous rosette herbs, and the peculiar E. rupicola 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, Trisctum, Carex 
Banksii, Pa^'onychia, Cardami^ie flaccid a, Callitriche, Rubus, Empeirum, Calysiegia 
and Gnaphalium, and, among the ferns, perhaps Cysiopteris. Rubus geoides forms 
together with R. radicans an isolated section but has a more southerly distribution 
but all may have reached Juan Fernandez from South Chile. 

Finally, Cuminia remains to be accounted for. Whether w^e 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 (i) is still represented in South 
America, the other (2) not. In (i) tw^o types can be distinguished, (a) not confined 
to subantarctic or alpine habitats and demanding a milder climate; to this lot I 
refer Uncinia Douglasii and costata, Drimys, Phrygila7itkus, Escallonia, Margyri- 
carpus, Sophora, Guiuicra, Apium, Pernettya, and possibly Planiago fernandezia. 
All have relatives in Chile. To these are added the species also occurring on the 
mainland: Danthonia, Koeleria, Juncus, 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 Araucaria, 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 COPELANI) many of the ferns also belong to i a: Hymenoglossum, 
five species of Hymeiwphyllum, Lopkosoria, Polystichum, Bleclnium, Hypolepis, 
Histiopteris, Gleichenia 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-knowm circumpolar assemblage of genera and species. Here we find 
Oreobolus, Uncijiia brevicaulis, phleoides and tenuis, and Lagenophora hirsuta, 
further three endemic species, Acaeiia masafuerana, Abrotaiiella crassipes, both 
with near relatives in West Patagonia-Fuegia, finally Agrostis 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, lIy))uniopliyllui)i falklandicuni, Polypodium (Grammitis) niagel- 
latiiciiiu, GlcicJioria quadripartiia, and Lycopodium inag€lla7naivi. 

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- 
cliactiu)n, Myrtio/a, Rn/uis, and so forth; the Boreal group is, as we have seen, 
also inxolved. 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 GrauDJiitis, 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 {2JI, 2^2, 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, 
AuthoxiDitJniui odoratiiDi and Rumex acctosella, 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 
u[) about 40 "o 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 Cakdoi' and Hkrz()(;: 56% of the mosses and 68% of the 
hej)atics 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 ha\e 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, Lobaria, 
Sticla and PscudocypIuUaria are frequent in tropical, subtropical and temperate- 
f)ceanic climates throughout, but the austral-circumpolar species are so many that 
we cannot exclude Antarctica as a possible source. 

We shall {)roceed to group 2. It comprises the genera or species which are 
foreign to the South American flora and have their relations in Australasia. JoilOW 
recogni/x'd 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 JoilOW thought, 28 species: Cladium, Carex bertero7iiana, Pepero- 
mia berteroana, margaritifera and Skottsbergii, Boehmeria, Santalum, Ranmuulns, 
Fagara (2), Halorrhagis (3), Euphrasia, Coprosma (2), Waklenbergia (5), Robinso- 
7iia (5), Syniphyochaeta and Rhctmodendron, the three last genera endemic. Objec- 
tions may be raised against including Carex and Euphrasia; the section to which 
Carex bertero7iiana was referred by KCkenthal is almost confined to New Zealand 
and barely represented in Australia, Tasmania and Norfolk Island, but one little 
known Chilean species is included, and Euphrasia formosissima is distantly related 
to E. 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, Santalum, Ranunculus, Halorrhagis and 
Coprosma. 

This element is conspicuous also among the ferns: the extremely old Thyr- 
sopteris, ArtJiroptcris, entirely unfamiliar with the neotropical flora, Dickso7tia, 
Blechnum ScJiottii 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, Cyatheoides tJiyrsopt- 
eroides in the Arauco flora, and Coprosma from Pichileufu, but the material is 
sterile. It is true that, to judge from Berry's illustrations, Cyatheoides suggests 
Thyrsopteris, but the author later [28. 57) compared it with h\s Dickso7iia 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. incerta, 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 Coprosma 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 



Jt is very easy to construct a liypothetical passage by which the "" Coprosnia 
group" readied Juan Fernandez without crossing the Antarctic or encroaching very 
much upon the surface of the Pacific Ocean. If we have reason to think that South 
America extended farther west I cannot see why this wasn't tiie case also with West 
Patagonia and Tierra del h\iego, 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 P^rnandez without finding their way east to what 
is n(nv 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 \: 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. 269). 

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 '' Coprosnia group" can be 
recognized. In IlKRZ()(;'s paper on the Hepaticae [130) a single species is indicated 
as restricted to Xew Zealand and Juan PY^nandez, Palla2>iciuia xiphoidcs, and two 
endemic s{)ecies are said to have their nearest allies not in South America but 
in Xew Zealand. \o case equal to Pallainci)iia is found among the mosses, but 
several endemic species are considered to be related, not to American ones, but 
to s{)ecies inhabiting the south-west Pacific region. 

With regard to tlie faioia I shall confine myself to some general remarks. 

The few land-birds are of neotropical origin. Of the seven species, three, 
liusttpluvius i:;aleyilus, 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 linstipluvms fcniaiidoisis. (ioiri sen's idea that it originated in 
the islands as a mutation of /:". o^alcritus 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 i)erhaj)s a late immigrant. Of the breeding 
sea birds the genus l^teroihoiiia forms an austral-circumpolar element and may, in 
j)reglacial time, have inhabited the coasts of Antarctica and adjacent islands. 

Little can be said as }'et about the invertebrates. The endemic leech, Ncso- 
philac))i())i, is an important case of non-American ancestry, and the only terrestrial 
amphi[)od 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 40I 

ism is high. In the fauna of the mainland Antarctic affinities have been stated 
to occur (Berland 2j. 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 Aulacodesimis and Nesogeophilus are austral genera 
and the species endemic; Schizotaenia 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, Pycnoinerodes and Eleusis. 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, Haplo- 
gonatus, Prenolepis and, perhaps, Metelia show Antarctic connections. 

The antiquity of the endemic land shells 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 P^ernandez [lOj. 995). 

S'il est bien ainsi, cette famille primordiale [the ancestors of Achatinellidae, Amas- 
tridae, Leptachatinidae and Tornatellinidae] doit avoir une tres ancienne origine et 
remonter au Paleozoique. . . . le peuplement malacologique de I'archipel des Hawaii 
est fort ancien et doit remonter a des temps primaires. II n'a put se faire, comme le 
preuvent les developpements precedents sur la repartition des genres et des especes, 
que sur une aire contenue, ce qui exclut la possibilite de considerer les iles Hawaii 
comme le resultat de I'activite des volcans sous-marins. 

26 — 557857 The Nat. Hist, of Juan Fernandez and EaJter Isl. Vol. I 



4o: 



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 BrCggen'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 hrst 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 
I'itzrova, SaxcgotJiaca, Pilgerodoidroji and Dacrydhim, 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 
XotJiofagiis flora. In my sketch of 1925 [2JI. 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 
XotJiofiii^us flora reached these latitudes", and BrCcgen 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 mezcia con una flora 
dc (lima mas fresco, (jue despues de la separaci6n de la Tierra de Juan Fernandez 
inmif^ro y (jue se caracteriza por los generos Nothofagus, Araucaria, etc. 

Araiicaria was found in the Pichileufu beds which are supposed to be Eocene, 
whereas the southern beeches appear in this latitude considerably later. In the 
Magellanian region Dl SKN, as we have seen, distinguished two horizons, a lower, 
Oligocene, with Xoiliofagus, and an upper. Lower Miocene, \m\\\\ Araucaria. Accord- 
ing t(j Hi<i'(;(;kn the separation of Juan Fernandez from Chile took place during 
the Oligocene Xavidad transgression. We would think that, if the bridge lasted 
longer, the XotJiofagus 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 Reiciie 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 403 

taken place as a consequence of the cooling of the chmate 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 P^ernandez- 
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 I75i> when also an earthquake occurred, and other earthquakes were registered 
in 1809, 1822 and 1835 (27 j). 

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 Stereocmilon 
[34-1), but a conditio sine qua 7wn 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 w^ork. 
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 Joiiow 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 wnth 
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 NoiJiofagus 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 NotJiofagus 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 liriireron, Euphrasia fonnosissi))ia, McgalacJme masafueraiia, PJwenicoseris regia^ 
]\anu7iculus capranuH and Robbisoiiia Masafuerae, all of them peculiar endemics, 
and of these Ranuuculus and JMegalacJine are found only along the high ridge 
above 1300 m. luen if some of them are of Antarctic ancestry, their history is 
another. When it comes to the Magellanian group, Orcoboliis, LageiiopJiora^ 
(ileichcnia, and so forth, it is difficult to exclude the possibility that they be 
glacial or postglacial immigrants. Carcx Baiiksii, limpeirnni ruhruni — only a single 
plant seen — (niliuui juasafueranimi, (iiiapJialiujii spiciforinc, Myrtcola yiuDiDiulai'ia 
and Rubus gcoides are, as was already told, not of Antarctic origin, but may have 
extended far south after the recession of the ice and accompanied Orcobohis, 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 
{:)revailing winds are westerly, storms from other directions occur, and there has 
been j^ilenty 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 l^\iegia were covered by inland ice [idf); 
if small, ice-free refuges occurred has not been definitely stated, but is not alto- 
getlier 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 Chiloe was not ice-covered, allowing the 
subantarctic fiora 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; scjueezed 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, 
Driniys, JiiaJiia, Cuminia, Escallonia, Rhethiodendron, 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. 

PaspaluDi L. More than 200; trop. to temp., most numerous in Amer. 

forstcria)un)i Muegge. X. Caled. 

scrobiculatnui L. var. orbicitlare (Forst.) Domin. X. Guin., Austral., N. Caled. ^ 
Polyn. 
Axouopus l^eauv. About 75, the majority in X. and S. Amer. 

paschalis Pilger. Related, according to I^ILGKR, to A. scoparius (Fluegge) 
Pilger, a S. American species. 
Siipa L.i 

horridula Pilger. Related species in S. Amer. and Austral. 
Sporoboliis R. Hr. Over lOO, trop.-subtrop., most numerous in Amer. 

elou(^atus R. Br. [as indicus (L.) R. Br. in 2jo\. S. As., Malays., Austral., 
X. Zeal. 
Calauiai:;rostis Adans. About 150, mostly temp., north and south. 

rctyofracta (Willd.) Link. (Agrostis W'illd., A. filitbrmis (Forst.) Spreng.) Aus- 
tral., 'i'asm., X. Zeal., Polyn., Hawaii. 
DichclacJuic ICndl. 2, Austral., X. Zeal. 

sc/Nna (R. \^v.) Hook. fil. Austral., X. Zeal. 
nauthojiia DC. 

paschalis Pilger. Perhaps most nearly related to /). cJiilciisis 13esv. 
I'j-(li^ras/is Host. ()\er 200, trop.-subtrop. 

i-loiigata Jacfj. \\. Ind. to Malays, and Polyn. 

Cyperaceae. 

C)pcri(s L. 

cragrostis Lam. \^ery 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. i and 
is not repeated liere. 



DERIVATION OF THE FLORA AND FAUNA 407 

polystachyus Rottb. Trop.-subtrop., wide-spread. 

cylindrostachys Boeck. As the former. 

brevifolhis (Rottb.) Hassk. Pantrop. 
Scirpus L. 

ripariiis Presl. N. Amer., Calif, to S. Amer., south to Fueg., Falkland. I can- 
not find that var. paschalis Kiiekenth. deserves to be distinguished. 

Juncaceae. 

J uncus L. 

plebeius R. Br. Colomb.-Urug., Austral, Tasm., N. Zeal. 

Piperaceae. 

Peperomia Ruiz et Pav. 

rejlexa Dietr. Pantrop., very widespread, north to Hawaii and south to 
N. Zeal.; also on Rapa and possibly Pitcairn. 

Chenopodiaceae. 

Chenopodiiim ambiguiun R. Br. Austral., Tasm., N. Zeal. 

Polygonaceae. 

Polygo7imn L. About 150, world-wide. 

acuminatum H. B. K. W. Ind., Centr. and S. Amer., Galap. Is., trop. and 
S. Afr., Orient. 

Nyctaginiaceae. 

Boerhaavia L. About 20. Pantrop. 

diffusa L. Widespread in the Pacific and a common weed in the Old and 
New World; probably of aboriginal introduction. 

Aizoaceae. 

Tetragonia L. 

expansa Murr. 

Cruciferae. 

Nasturtium R. Br. About 50. Widespread, mostly temp. 
sarmentosum (Sol.) O. E. Sch, Austral.-Polyn., Hawaii. 

Leguminosae. 

Caesalpiiiia L. At least 90-100; pantrop. 

bonduc (L.) Roxb. Trop. As.-Polyn. 
Soph or a L. 

toromiro (Phil.) Skottsb. Nearly related to the species from J. Fern. 

Euphorbiaceae. 

Euphorbia L. 1 500-1600; world-wide. 

hirta L. Indomal.-Polyn., often adventitious. 

serpe7is L. As the former, also common in trop. Amer. 



408 C. SKOTTSBERG 

Umbelliferae. 

ApiuDi L. 

prostration Labill. Austral-circuiii[)., incl. A. atistrale Thouars. 

Primulaceae. 

Satiiolus L. 9 (12?), I cosmop.. 2 (5?) X. Amer., 3 southern S. Amer., i S. Afr., 
I W. Austral., and tlie following. 

repejis (Forst.) Pers. Austral circump. 

Gentianaceae. 

Erythraea Neck. (Centaurium Hill.) 30-40, subtrop.-temp. 
australis R. Br. Austral., \. Caled., Fiji. 

Convolvulaceae. 

Calystegia R. l^r. 7-8, temp.-subtrop. 

stpiuin (L.) R. Br. forma. All continents, also reported from Australia, but 
possibly introduced in the s. hemisph. 
Ipoiuaca L. 

pes caprae (L.) Roth. Pantrop. 

Solanaceae. 

LyciuDi L. About 100, most numerous in S. Amer. 

carolhiiamiui Walt. var. scDuiuiccnsc (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" Solamini Insulae 
PascJialis Bitter, see 2^c)\ it was used as medicine. P'our species are endemic 
(13 %), but three of them belong to large grass genera needing monographic study. 
FoRSiKK mentions {347), beside some cultigens, only Panictmi filifonue Jacq. 
(-- Digitaria sanguinalis), a common weed, SJicfficldia ( = Samolus) rcpens, Aveim 
filiformis ( -- Calamagrostis retrofracta) and Solanioii nigruvi; in his journal (j^<?) 
he refers to Apiuni^ which he knew from New- Zealand, to ''JMivwsa' ( = Sophora), 
and also to the former occurrence of sandalnwod. In regard to the distribution 
of SiDitalion in Southeastern l\)lynesia, where S. hisulare Bert., including varieties, 
is known from Tahiti, Kaiatea, Marquesas, and Austral Is., and 5. li€ndersone7isc 
F. B. R. Brown (very chxse 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 l^Laster Island, perhaps introduced by the aborigines; not 
very long ago S. yasi 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 liaster 
islanders, took possession of the island, he brought various useful trees and other 
plants, a story first told by Fokster (,?^<^'. 583). Mhtraix [180. i^-xy) enu- 



DERIVATION OF THE FLORA AND FAUNA 409 

merates mahute [Brotissonetia papyrifera), makoi [Thcspcsia populnea), hauhau 
[Triumfetta semitfiloba), toromiro [Sophora), naunau (Saiiialum), and marikuru 
[Sapijtdus 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 Sophora wood, 
called toromiro, while the name of the tree with the fragrant wood was naunau. 
Cooke {34.3. 722) translated this word "bastard sandalwood". This is the name 
given to Myoporum sandvicense Gray in Hawaii, where the native name is naio 
(naeo, naieo); Santalmn is called iliaJii, in Marquesas puahi, on Rapa eahi, in 
Tahiti and Tonga ahi. Whether or not the words naunau and naio have the 
same base I cannot tell; on Raivavae, Rapa and New Zealand the name of 
Myoporum is ngaio, whereas true sandalwood, in this case Mida, is called 
maire, the Hawaiian name for the fragrant Alyxia oliviformis Gaud., transferred 
to Mida in New Zealand, where no Alyxia 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 {180. 17-18). 

My informant gave "sandal" as the Spanish equivalent of the word naunau — 
the correct Spanish word is sdndalo — 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 naunau 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 [33. ill). To judge from the distribution, jiaunau could 
just as well refer to Myoporum, a genus represented by endemic species in Rapa 
— where, as we shall learn presently, 7mu 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 Santalum 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 
Sebastl\N Englert, who has lived more than 1 5 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. 



^lO C. SKOTTSBERG 

yau o Naunaii (Santaliim): arbiisto de la familia de las santaldceas, tambien 11a- 
mado nan opata, poniue creci'a en los barrancos (opata) de la costa, entre rocas y 
I)iedras. Ahora ha desaparecido. I.os ultimos ejemplares (jiie algunos de los natives 
actuales recuerdan haber visto todavi'a, se ban secado hace unos 50 anos. El nau opata 
daba, como friitos, niieces del tamano de castafias, los "mako'i nau opata". Carl 
Friedrich Behrens nombra nueces entre los friitos que los islenos les regalaron en gran 
numero a el y a sus compaiieros. Hotii Matiia y su gente parecen haber trafdo gran 
cantidad de cstas nueces, i)orciue de ellas se alimentaron en los primeros meses des- 
pues de haber llegado a la isla. Al excavar la tierra en cuevas (jue estaban antigua- 
niente habitadas se encuentran cascaras de estas nueces. Estas cascaras generalmente 
no cstan ([uebradas sino ([ue han sido abiertas en forma de un pecjueno circulo, para 
ser usadas por los ninos en el juego del trompo. La madera del arbusto se utilizaba, 
j)or su cx([uisito aroma, para confeccionar un perfume, como los veremos en otro capitulo. 

This description does not at all fit either Santaluin or Myoportwi. The fruit 
of SaiitaluDi 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. pyrularhan Gray) the drupe was 16-18 mm 
long and the stone 12-15 mm. According to HlLLEBRANl) (joy. 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 Jiatinau which 
grew along the coast and produced the "cascaras" found in the caves cannot 
have been a species of Sajitaluui. Fortunately this could be proved. Mr. STEELE 
had told me that Father FLnglert had sent him two shells for his collection 
of l^aster Lsland 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 Saiitaliini. A passage in Me'I'RAUX's book, p. 353, 
j)ut 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 viakoi 
— 'riicspesia popiibiea cajxsules! A comparison with herbarium material showed 
that Mr. Steele's specimen is a typical capsule of yy/r.sy^ri-/"^, one of the "nuts" 
found in the caves. They made \ery poor food but good spinning-tops. We 
did not see Thcspesia on the island, but Metrai X observed it growing on the 
cliffs at Poike (the eastern headland). F^vidently the word nau or nauuau has 
been altogether misplaced by the blaster islanders, although both Metraux 
and bLxcLERi- were told that it was the name of Santaluni. We find the same 
word in Tahiti and the Tuamotu Islands for Lcpidiu))i bide7itaiuni Montin, and 
this is called iiaupata, strikingly like FLxglert's }iau opata, in Marquesas; in 
Tahiti naupata means Scaci'ola frutcscens, which is called naupaka in Hawaii and 
}ii^ai{}ii!;au in Rarotonga, and on Rapa }iau is used for So)icJius oleracens. All 
these {plants were used as medicine. 



DERIVATION OF THE FLORA AND FAUNA 4 II 

Geographical elements. 

In 1934 I made