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ACTA PHYTOGEOGRAPHICA SUECICA EDIDIT

SVENSKA VA.XTGEOGRAFISKA SA.LLSKAPET

42

NAHUEL HUAPI

PLANKTON OF SOME LAKES IN AN ARGENTINE

NATIONAL PARK WITH NOTES ON TERRESTRIAL

VEGETATION

BY

KUNO THOMASSON

UPPSALA 1959

ALMQVIST & WIKSELLS BOKTRYCKERI AB

SVENSKA VAXTGEOGRAFISKA ·sA.LLSKA.PET (SOCIETAS PHYTOGEOGRAPHICA SUECANA)

Adress: Uppaala UniversitetB Vaxtbiologiaka Inatitution, Villaviigen 14, Uppsala 8, Sverige (

Styrelse: Ordf. Prof. G. EINAR Du RIETZ, v. ordf. Prof. HuGo OsvALD, sekr. Fil. kand. BENGT M. P. LARSSON, skattm. Fil. mag. Sv ANTE PEXKARI, red. Doe. MATS W JERN, klubbm. Fil. lie·. ELIEL STEEN,

avr . . � Prof. Emo HULTEN, Prof. BERTIL LINDQUIST, Prof. JOHN AxEL NANNFELDT,

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ACTA PHYTOGEOGRAPHICA SUECICA 42

NAHUEL HUAPI

PLANKTON OF SOME LAKES IN AN ARGENTINE

NATIONAL PARK, W I T H N O T E S ON T E R RES T R I A L

V E G E T AT I O N

BY

KUNO THOMASSON

UPPSALA 1959 Almqvist & Wiksells Boktryckeri AB

The present paper constitutes no. 4 of the Studies on South

American Fresh-Water Plankton by the author, and no. 2

of the Reports of the Swedish Limnological Expedition to

South America in 1953-54.

C O NTE NTS

I. Preface 5 VIII . Tarns, ponds, and temporary waters . 51

II. Introduction 6', 1. Strand-pool 51

III. Climate 7 2 . Pond near military camp 52

IV. Geology 9 3. Inundation pool 53

V. Flora and vegetation 10 4. Tarn in Paso V uriloche 55

1 . Eastern zone . 13 5. Alpine tarns 56

2. Central zone . 1 5 IX. Running waters 56

3. Western zone 24 1. Brook at Lake Gutierrez 56

4. Alpine belt 28 2. Brook at Lake Mascardi 56

VI. Hydrography 34 3. Riv.er between L. Gutierrez and L.

VII. Lakes 36 N ahuel Huapi 57

1. Nahuel Huapi 36 X. Benthos 58

a. Morphometry 36 XI. Retrospective survey 59

b. Thermal relations 37 XII. Taxonomical comments 64

c. Plankton 42 1. Microphyta 64

2 . Gutierrez 44 2. Microzoa 76

3 . Mascardi 45

4. Guillelmo 46 XIII. Fishes and waterfowl 79

5. Frias 48 Acknowledgements 80

6. Hess 49 References . 81

A cta Phytogeogr. Suec. 42


;· ..... . .. .· ·213o

. . . ..

Jnternational boundary

Divortium

I. P R EFA C E

The National Park of Nahuel Huapi, situated between 71 o and 72° west longitude and 40°20' and 41 °35' south latitude, is the largest of Argentine national parks, its area being 785,000 ha. It lies along the eastern slope of the Andean chain and encircles the large Lake Nahuel Huapi and a number of smaller lakes. Some of the lakes within the N ational Park were the object of a short visit by the Swedish Limnological South-America Expedition 1953-54.

Our thanks are due to the Argentine Ministry of Defence for their generous support making it possible for us to obtain the maximum benefit. from our short 10-days visit. The representative of Ministry of Defence Captain CARLOS HoFFM.ANN

from Buenos Aires did all in his power to facilitate our studies on the natural features of the National Park of Nahuel Huapi. We are also indebted to him for his good fellowship during the often rather arduous excursions in the National Park.

Our sojourn in the National Park lasted from

January 19 until February 1, 1954. During our stay we studied the lakes in the southern half of the National Park. Naturally it was quite out of the question to carry out any thorough investigations in the too. short time at our disposal. Hence our studies were restricted to obtaining a general view of physical conditions and to collecting random samples from lakes in order to enable us to get some comparative data for our investigations on the lakes west of the Andean range, in the Valdivian area. Samples of plankton and bottom fauna were collected, in addition sporadic observations were made on the terrestrial flora and fauna along our excursion routes.

The following notes are of a sketchy character,

in spite of which we hope to contribute something to the knowledge of plankters in South American lakes. The few words on the natural features of the National Park of Nahuel Huapi are also fragmentary but give an indication of the nature of the terrain.

Fig. 1. Map of the Nahuel Huapi National Park and its situation in South America. The lakes surveyed by the

present author are N ahuel Huapi, Gutierrez, Mascardi, Guillelmo, Frias, and Hess.


I I . I NT R O DUCTI O N

The discovery of the Nahuel Huapi area can be

traced back to the legend about "Ciudad de los Cesares" which plays an important role in all stages of earlier exploration history of South America.

The conquerors were followed by Jesuits, some of

whom are perpetuated in the names of lakes, e.g. Mascardi and Guillelmo.

Dr. FRANCISCO P. MoRENO was the first to record the natural history of the Nahuel Huapi area� Dur� ing his many exploratory expeditions and finally as leader of the Argentinean-Chilean frontier-commission he was seized by the extraordinary charm of the scenery around Lake Nahuel Huapi. He fully appreciated the value of this area· as a significant natural region and proposed in 1903 that the Nahuel Huapi area should be proclaimed a national park. According to a decree of February 1, 1909 the Nahuel Huapi National Park was founded. Its territory was later extended from about 9 square miles to 3030 square miles.

The principal centre for Nahuel Huapi National Park is the town San Carlos de Bariloche, the well known startingpoint · of the ride of the Swedish Patagonian and Tierra del Fuego Expedition

.(1907-

09) in 1908, down to the Straits of Magellan. The settlement was then made up of a few primitive ramshackle houses only few years old. A visitor today finds a very handsome up-to-date settlement, modern hotels and private villas. Bariloche has been developed as the main tourist and winter sport resort in Argentina. There are more than 10,000 inhabitants, the German element being important, and this is reflected in the style of architecture. Tourist organization is well-arranged in the National Park, a park service being responsible for it. Supervision over this huge area of 785,000 ha is carried out by numerous wardens.

We have perhaps nowhere else a better opportunity of seeing combined and concentrated such rich variety of the most wonderful and varied examples of beautiful scenery and magnificent panoramas of the immense Andean mountains than in the Nahuel Huapi National Park. One is often

A cta Phytogeogr. Suec. 4.2

inclined to make a comparison with the Swiss scenery; on the other hand the number of rockbound lakes hidden in the interior of the Cordillera, gleaming between the precipitous rock-faces has very often duplicated the nature of Norwegian fjords. The lakes are surrounded by the greenery of luxuriant forests, dominated by Nothofagus. On the slope of the Chilean watershed the forest is at its best; here we meet the exuberant temperate rainforest, known as the V aldivian forest. The extraordinary overgrowth of trees and shrubs the imposing intert·wining of lianas artistically twisting round the trunks and branches of the massive N othofagus

dombeyi and the Lomatia obligua recall impressions of the cloudforests of the Urubamba-valley in Peru.

The western boundary of the park is formed by the immense barrier of the snow-capped Cordillera. Above the serried lines of haughty peaks towers the white massive cupola of Mt. Tronador (3554 m a. s.-1.), of volcanic origin, the highest mountain in Northern Patagonia. From this giant of the Cordillera there issues enormous glaciers whose formidable bulks hanging from its steep walls emit hoarse growls and thunderous roars when the sun melts the snow and ice. These glaciers terminate in the magnificent belt of green woods below. Mt.

Tronador is the only mountain exceeding 2500 m. The number of peaks exceeding 1800 m is about 55. Looking from Tronador these summits form a "Gipfelflur", residuum of an old peneplane from the Miocene. This on the whole uniform range of mountains is interrupted on the horizon by the Chilean volcanoes Puntiagudo and Osorno.

The National ·Park of Nahuel Huapi contains six nature sanctuaries (areas intangibles) of extensive area which are wholly exempted from human in

fluence and activity. These areas on one hand represent the most characteristic stretches of

natural scenery; on the other hand they are established for preservation of the rarer species of autoctonous fauna and flora which demands special care and attention to be preserved from extermination.

8 Climate

25 c· •••••

•

Patagones Pilcaniyen Bariloche

40°48'5 41° 10'5 41° 10' s

20 •• •·•

•

••

•

•

• \ •

.

•

\

15

10

5

\ I I I \ \ \ \ \ \ \

\ \ \ \ \

0 :

' ' '

··· ..... ··

" .J ................

62°40'W 70°41'W 71° 21'W

/ / I I

I I I

I I

. ·

I /

11 Ill IV V VI VII VIII IX X XI XII

Fig. 3. Mean temperature at three stations situated at

abo ut the same latitude between the Andean range and

the Atlantic.

40

r-30

20

0 � ...

0

- Temperatvre

--- Precipitation

)/ -� ��

..,.I/ ..... """"

. .. _ - .. ·

XI XII


� /

i'' / _, '� /

Ill IV

I I

I I

I \ I \ I I \ I \

\ \

' \ '

' ' '

... � / 1"--.. .,

VI VII VIII IX

The amount of rain is very low during the growing season, e.g. in Bariloche 264 mm, consequently the drought is of greatest importance for plant life, because the growing season coincides also with the high temperatures, cf. fig. 4 .

As with precipitation the available observations on other elements of climate are restricted to Bariloche. A more detailed network of meteorological stations in the National Park would be of great importance in respect to the varied meteorological conditions of this region.

The mean temperature for Bariloche is 8.1 °0, the average value for temperature maxima is 13.4°, and for minima 3.4°. The highest temperature ever recorded in Bariloche is 35.3°, and the lowest one -14°. The amplitude for summer temperatures is

12..4°, and for winter temperatures 6.7°. Of great importance for the vegetation are the frost nights which may occur over the greater part of the year, viz. in the interval between February and December. The duration of snow cover is about 70 days.

Prevailing winds are the strong descending winds from W and SW, whereas the winds from N and NE are of no importance. The effect of prevailing winds is reflected in the shape of the timber line, e.g., on the mountain slopes round Bariloche where the catch of rain has decreased considerably (pl. 6) in relation to the western parts of the national park, we notice that the timber line has a serrated shape. The forest extends upward along the ravines and gullies which provide protection from the violent forces of the winds. Furthermore the conditions of moisture are more favourable in these localities, partly because the parching effect of the winds is eliminated, and partly depending upon the fact that a large amount of snow accumulates in these depressions and remains there for a long period.

Fig. 4. Monthly mean active temperature and monthly

mean effective precipitation in San Carlos de Bariloche.

From PEREZ-MOREAU, 1 948.

I V. G E O L O GY

From the geological point of view the Andean range is a relatively young formation. The tectonic processes which have folded up this majestic wall along the western coast of the ·continent are still active, as is evident from the long line of volcanoes bordering the western base of the Andean range. Some of them are still active, the other ones are extinct at least at the present time. The unsettlement of the earth's crust is also reflected by the

frequent earthquakes. Considering the tectonic activity there is nothing

to be wondered at in the particularly intricate character of the geological structure of the Andean range. The tectonic movements, volcanism and erosion etc., have resulted in rather mosaic-like geological patterns.

The volcanism in Northern Patagonia was localfzed during the last interglacial period to a more easterly zone in the high Cordillera, though still west of the axis of maximum heights which cuts

the central part of Lake Nahuel Huapi. The volcanic activity at that time was obviously very intense, as evidence has been found that the preexisting relief in the Nahuel Huapi area was wholly drowned under the eruptive masses. After the ceasing of the interglacial volcanism these products

were removed, largely by fluvial and glacial erosion. A great area of these rocks is preserved with Mt. Tronador, the highest mountain of this district, as the most important volcanic centre.

Eastward of these so-called Tronador series a

large area is covered with a granodiorite series which forms the greater part of the precipitous shores of Lake Frias. Also the Rio Manso valley running down from the V entisquero Grande to Lake Mascardi is excavated in a granodiorite series.

Lake Mascardi is surrounded mainly by the highmetamorphic basement and by the above mentioned granodiorite series which also form the basin of Lake Guillelmo, a little lake just south-east of Lake Mascardi. Lake Hess is surrounded by the same series. Lake Fonck, which has not been visited by us, lies wholly surrounded by the Tronador series.

As this lake drains into Lake Hess, it is not out of the question that this is a factor to be reckoned with when considering the divergent character of Lake Hess. Lake Gutierrez is surrounded in its south-western half by gneisses, highly metamorphic, regarded by LJUNGNER (1931) as belonging to the basement of the Andean geosyncline. The northern half of Lake Guatierrez is surrounded by the Nahuel Huapi series, which also form a broad continuous

zone with northwesterly strike through the central part of the Lake Nahuel Huapi. The Nahuel Huapi series consist mainly of tuffs of varying appearance interspersed with lava flows of dacitic, andesitic or basaltic composition. The eastern parts of Lake Nahuel Huapi, approximately east of Bariloche longitude are surrounded by glacial formations. Around the Brazo Tristeza, the southwestern arm of Lake Nahuel Huapi, the Tristeza granite series occurs. It also occurs on the northern shore of Brazo del Viento, the western arm of Lake Nahuel Huapi. Besides the above mentioned Tristeza granite the shores of Brazo del Viento are made up of N ahuel Huapi series, Milleaqueco formation and granodiorite series.

The relief conditions of the Nahuel Huapi region at the end of the Pliocene period were in their main features the same as those of the present time. The modifications which subsequently ensued are to be ascribed to a large extent to the partly eroding, partly depositing activity of the Pleistocene glaciations and to the volcanic activity recommencing in interglacial time. During the latest glaciation the whole region was covered by an immense ice-sheet, except some of the most elevated mountain peaks which projected through the ice-sheet forming nunataks. The constructional effects of glacial action have resulted in deposition of morainic material and sheets of sand and gravel mainly in the valleys, consisting of outwash fans, outwash deltas and kames. One ought to mention the two large systems of terminal moraines at the eastern end of Lake Nahuel Huapi; moreover one can find numerous very beautiful moraines of different origin in the


10 Flora and vegetation

valleys reaching far into the heart of the Cordillera, cf. pl. 8. Some of the minor lakes are dammed up behind the moraines, e.g. Lake Frias. The basins of small lakes are to a great extent the result of the glacial action during the ·last glaciation. Consequently they are all relatively young, as opposed to the ancient large Lake Nahuel Huapi. The deep basin of the latter lake is characterized by numerous inlets bounded by steep mountain slopes, a basin which according to LJUNGNER (1931) existed before the last glaciation.

The soil is, except in the alpine region and recent alluvian fans, formed by an eolian cover. It consists partly of volcanic ash and partly of loess, and is poor in calcium.

For thorough information on the geological features of the Nahuel Huapi area see LJUNGNER 1931, CoRDINI 1939, LARSSON 1940 and AUER 1958, where an extensive geological bibliography on the Nahuel Huapi region can be found.

V. FLO R A A N.D V E G ETATI O N

The National Park of Nahuel Huapi extends over the area between the Patagonian steppe and the ice-covered peaks of Andean chain. The diversity of its biotopes is reflected in the rich flora and great variety of vegetation.

The . following, unfortunately too superficial survey of the vegetation of the Nahuel Huapi area has been ]ncluded partly as a complement to the physical features and partly to contrast with the planktic vegetation of the lakes. The phanerogamic flora is exclusively composed of plants with strange names, though there are many which have a somewhat familiar appearance for the European botanist. In striking contrast the phytoplankton is mainly composed of plants well known or at least closely related to those occurring in lakes in the northern hemisphere.

The following sketch illustrates the vegetation only in the southern half of the National Park, and by no means does it set out to be a complete description of the vegetation, but recapitulates orily impressions and notes, and also includes an account of a part of some collections made during a 10-day journey in the National Park of Nahuel Huapi.

There are quite a number of publications on the flora and vegetation of the Nahuel Huapi National Park and it� adjacent areas, e.g. HossEus (1916)_, SKOTTSBERG (1916), LJUNGNER (1939), CABRERA

(1939), KALELA (1941), PEREZ MoREAU (1944),


KALELA ( 1946) and A UER ( 1958). A monograph on the vegetation in this area, however,-is still required, as is also a flora, for about 1000 vascular plants occurring in the National Park, of which the Oompositae constitute some 25 per cent.

According to the most recent phytogeographical survey of the Argentine flora by CABRERA (1953), the following floristic regions are represented in, or are adjacent to, the National Park of Nahuel Huapi:

Region N eotropical Provincia Altoandina

Distrito Altoandina austral Provincia Patagonica

Distrito Patagonico subandino Region Austral

Provincia Subantarctica Distrito del Bosque Caducifolio Distrito Valdiviano

The earlier attempts to divid� up the Argentine flora have been lucidly presented and fully discussed in the above mentioned paper by CABRERA.

First it is necessary to emphasize a few of the ecological features which are reflected in the character of the vegetation.

The maritime influence increases towards the west and the continental towards the east, while the northern slopes of the mountains are more parched than those in the south for we are in the

Flora and vegetation 11

mm m

4000 4000

3000 3000

2000 2000

1000 1000

0 50 lOO km Fig. 5. Distribution of the yearly average precipitation (mm) in the Nahuel Huapi area between the frontier and

Pilcaniyen.

southern hemisphere. The borderlines, depending on the

.exposure, are very distinctive.

Mountains grow wetter from their bases upward, because, as the atmosphere grows steadily cooler with altitude, more and more moisture condenses into rain or snow. On the high mountains most of the moisture in the air may condense and fall at levels below the summits; thus the tops of such mountains, towering above the highest rain clouds are often dry. This is reflected in the fact that some evergreen plants common in the western part

of the National Park, e.g. Drimys winteri, Maytenus

magellanicus, Berberis pearcei, · etc., occur in the eastern part of the National Park only in the decidous N othofagus pumilio belt, on the slopes of the mountains. In the western part these plants are to be found also in the lower altitudes in the N othofagus

dombeyi b�lt as well as higher up on the slopes of mountains, in the N othofagus pumilio belt. The increase in precipitation with altitude is also re-

sponsible for the phenomenon that at high altitudes the decidous forest reaches further towards the east than at low altitudes; here the pampas vegetation has spread considerably to the detritment of the forest.

The snow -line rises from the west to the east, partly depending on the decreasing precipitation, and partly due to an increasingly continental influence which is reflected in high summer temperatures. The timber-line which, on the average, lies 1600 m a. s.-1., and about 800 m above the lake level, rises from the west to the east. The rise of timber-line is not so pronounced as the rise of the snow-line. Regarding the upper belt of deciduous forest which is independent of winter temperatures the effective climatic factors are increasing summer warmth and drought. The lower belt of evergreen forest is sensitive to the increasingly continental influence of falling winter temperatures. Therefore the belt of �vergreen forest shows from west to e�st falling



upper boundary. The vacated space is occupied by decidous forest and consequently the belt of decidous forest widens towards the east.

Above the timber-line there is a narrow belt of steppe vegetation whose members are also found growing in the upper parts of forest-belt, as will be illustrated below. Nothing equivalent to alpine meadows occurs in the mountains of "suissa Argentina". This high-Andean steppe belt, which from the ecological point of view is characterized

Oligohumid

as frost-desert, narrows towards the east, and varies in breadth depending on exposure.

The forests of Nahuel Huapi National Park were the object of investigations by the Finnish Patagonia-Expedition in 1937-38. Unfortunately only a part of the results has been published. The following survey on the forest types according to A. KALELA is here reproduced, somewhat modified, from E. K. KALELA, 1941:

Mesohumid Poly humid

l . Berberis pearcei type

Subalpine

Montane

A. Alstroemeria variety I B. Berberis linearifolia variety

2. Osrnorrhiza-Rhacoma type

A. Valeriana laxiflora variety I B. Berberis linearifolia variety ·

l . Vicia-Alstroemeria type 2. Osmorrhiza-Mutisia type

3. Rhacoma-Lathyrus type

4. Oarex patagonica type

l . Blechnum-Viola reichei type 2. Osmorrhiza-Adenocaulon type

3. Rhacoma-Berberis darwinii type

4. Gaultheria type

l. Dioscorea-Alstroemeria type l . Azara-Blechnum- Viola reichei type

Sub- A. Variety rich in grasses 2. Azara-Osmorrhiza-Adenocaulon montane type

B. Variety poor in grasses 3. Azara-Rhacoma-Berberis dar-winii type

l . Asteranthera-Nertera type 2. Asteranthera-Luzuriaga

type 3. Desfontainea-Gaultheria

type

While studying the flora of a distant area of unfamiliar character the taxonomic work on the collections is as a rule the most time-consuming and difficult aspect. However, I have had the privilege of -receiving generous assistance in the identification of a part of my collections from C. G. ALM

(Uppsala), M. BARROS (Buenos Aires), ELENA 0. BoRSINI (Tucuman), A. BuRKART (San Isidro ), A. L. CABRERA (La Plata), L. A. GARAY (Toronto), G. LoosER (Santiago), ALICIA LOURTEIG (Paris), R. SANTESSON (Uppsala), H. 0. SLEUMER (Leiden), B. SPARRE (Stockholm). I am extremely grateful for their �indness and generosity.

In the following survey, the vegetation of the National Park of Nahuel Huapi has been divided into four areas:

A cta Phytogeogr. Suec . .42

l. The eastern zone-pampas. 2. The central zone-it is in reality only a transi

tional zone between the first one and the following one.

3. The western zone-the Valdivian forest. 4. The alpine belt-this is of rather uniform

character and extends above the central and western zones.


L. BRUNDIN Fig. 6. Steppe east of Bariloche. Vegetation of Festuca and Mulinum spinosum.

1. EA S T E R N Z O N E

This zone includes a part of the Distrito Patagonico suba:ridino CABRERA (1953) which extends between Distrito Valdiviano and Distrito Patagonica accidental and embraces only the very eastern part of the National Park of Nahuel Huapi. LJUNGNER (1939) designates it as "The fourth meridional zone". In comparison to the remaining districts of Provincia Patagonica, the Distrito Patagonico subandino is cooler, the average tem7 perature of the year is 8°0, and has a higher average precipitation of 200-350 mm. The present notes and collections were made in the surroundings of Bariloche which lies on the boundary between the present zone and the following one.

The western part of the Patagonian plain and the

eastern offsets from the Andean range are covered with pampas vegetation. The pampas vegetation passes in the east into semidesert vegetation, the latter being sparse and of low growth, without tall shrubs. It is made up of xerophilous, small-leaved scrub plants, e.g. Nardophyllum, Nassauvia, Ver

bena, Chuquiragua. Contrary to pampas vegetation the grasses are here without any importance. Instead of tufts of grasses, the· semidesert vegetation is characterized by cushion plants, like Adesmia,

Azorella, Benthamiella, Bolax, M ulinum, N assauvia

and only along the streams is shrub-vegetation encountered. This semidesert presents a very monotonous landscape-"region maltida o Paso del Diabolo".

Acta Phytogeogr. Suec. 42


L. BRUNDIN

Fig. 7. Steppe east of Bariloche. The tussocks of Festuca and Stipa, in the background Baccharis shrub.

The pampas vegetation in contrast to the semidesert vegetation is more compact, the open space between the plants being less than in the semidesert. The grass vegetation is luxuriant, and the whole landscape is often dominated by the plumes of tall grasses waving in the wind. The most important species are Agrostis pyrogea, Bromus macranthus,

Danthonia sp. , Deschampsia antarctica, Festuca

argentina, F. monticola, F. pallescens, Hordeum

comosum, Poa lanuginosa, P. ligularis, Stipa hirti

flora, S. humilis, S. neaei, S. speciosa, all of which grow in large tu;:;socks. When proceeding from the east to the west, i.e. towards the increasing precipitation, one crosses first the Senecio-, then the Stipa-, and

at last the Festuca-steppes. Small shrubs are abundant. Tall shrubs and also trees grow in the favourable localities, e.g. river valleys and precipices, cf. pl. 2. Also in these localities grow shrubs of the genera


Adesmia, Anarthrophyllum, and Berberis; here and there a few trees occur, e.g. Austrocedrus chilensis

and N othofagus antarctica. In the foremost eastern offsets of the Andean range the Festuca-steppe turns into a kind of park-like landscape. Here the patches

of steppe vegetation are intermingled with patches . of forest vegetation forming a mosaic. The vegeta

tion is here of mesophilous character. The groves are formed by Austrocedrus chilensis, Maytenus

boaria, N othofagus antarctica, N. pumilio, and Schinus patagonicus. Amongst the shrubs and dwarf shrubs mention should be made of: -

Berberis buxifolia B. cuneata B. empetrifolia B. heterophylla Azara microphylla Adesmia campestris

A narthrophyllum patagonicum

A . rigidum Escallonia virgata Ribes cucullatum Ovidia pillopillo


Embothrium coccineum Lomatia hirsuta Schinus polygamus Colliguaya integerrima Colletia spinosa Discaria serratifolia D. trinervis Buddleja globosa Fabiana imbricata Diostea juncea Oorynabutilon bicolor M ulinum spinosum

Baccharis lanceolata B. rosmarinifolia v.

subandina ·

B. umbelliformis ' H aplopappus· pectinatus N ardophyllum

parvifolium N. obtusifolium N assauvia aculeata Senecio coxi S. microcephalus

As in the case of the trees and shrubs, the her

baceous flora; too, is rich. Intermingled among

the above-mentioned grasses are a variety of herbs,

some of which also occur in the eastern part of

pampas. We find here inter alia: -

Agrostis leptotricha A . pyrogea Bromus stramineus Oortaderia selloana Elymus patagonicus H olcus lanatus Hordeum murinum I mperata condensata Luzula chilensis

M icrosteris gracilis N asella chilensis V ulpia eriolepis V. megalura. Oarex subantarctica Ephedra andina Amaryllis elwessi H abranthus bagnoldii Sisyrinchium junceum

S. striatum Anemone decapetala A. multifida V i0la maculata Oajaphora silvestris Oardamine integrifolia Draba australis Acaena acroglochin A. pinnatifida A . poeppigiana A . pseudopoeppigiana A . splendens Astragalus cruckshanksii Adesmia palenae Arjona tuberosa Euphorbia portulacoides

ssp. nahuelhuapiana Geranium sessiliflorum

Oerastium arvense Oolomia biflora Gilia gracilis Oalceolaria lanceolata 0. polyrrhiza Veronica peregrina ssp.

xalopensis H eliothropium

paronychioides Plagiobothrys verrucosus Azorella nucamentacea A. tri furcata Oruckshanksia glacialis V aleriana carnosa H aplopappus prunelloides N assauvia glomerulosa

·Senecio coxi Triptilon achillea

The spinosity of many of the plants is most

striking. The desmochore plants are abundant.

Gradually one passes over into a truly transitional

zone betw_een pampas and temperate rainforest

for no sharp and obvious boundaries between these

two exist. The clumps of trees become more and

more luxuriant and broader; the transition is one

of succession.

2. CEN T R AL Z O NE

This zone is appropriately divided into two sub

zones:

Subzone A is obviously influenced by pampas vege

tation. It contains a considerable number

of elements representative of the pampas

vegetation.

Subzone B bears the impress of the Valdivian dis

trict. It contains a rich variety of re

presentatives from the Valdivian flora.

The forest is prevailingly compact, and

could be designated as mesophytic

forest.

These subzones together form a gradual transi

tion from the pampas in the east to the temperate

rainforest in the west, in respect to the flora, as

well as in the character of vegetation. The whole

transition zone could be designated as a major eco-

tone. This is a contact zone between two climaxes,

where small changes in climate produce a contam

ination of one zone by the ot.her.

S ubzone A

The pampas vegetation is replaced about the

latitude of Bariloche by a mosaic-like vegetation.

There are ·no sharp boundaries, the steppe vegeta

tion ramifying deeply towards the Cordilleras .

Likewise the mesophytic forest penetrates along the

streams and slopes of the mountains into the pam

pas. The result is a rather broad area with both of

these elements intertwined. This area has been de

signated by LJUNGNER (1939) as the "Third merid

ional zone-fringing forest".

The average precipitation varies within the limits



of the present area between 700 and 1300 mm per an

num The boundary between the steppe and forest

coincides on the whole with an annual rainfall of

1000-1 100 mm.

The distribution of the different elements with

in this transition zone is determined mainly by mi

croclimatical, oreographic, and historical factors.

Among the last mentioned the following two are

the most important and pronounced. The first one

is the increasing aridity, cf. KALELA, 1941 , p. 141.

The second and probably more important is the

human interferences.

It is a pity that there are no observations avail

able on precipitation for a long enough period. But

a comparison between the sparse records from the

very beginning of the century and those of recent

years show that the rainfall has somewhat di

minished. The increasing aridity is more obviously

manifest in the retrogression of forest, cf. KALELA,

1941 . The rejuvenation is insignificant, and in

general gives one the impression that it is degener

ating, this being a zone of struggle for the forest.

Here on the boundary towards the steppe, the forest

is-involved in a constant battle against the drought,

cf. p. 27 and fig. 12 in AuER, 1958. When the forest

is destroyed by fire or lumbering, its place is taken

by steppe vegetation, for it seems to be unable to

recolonize an area from which it has been removed.

A further important factor to reckon with is

overgrazing, which results in the prevention of

rejuvenation through the young plants being dam

aged and produces a profound change of biotope.

The overgrazing in this zone, where the forest even

under the natural conditions has difficulty in

upholding itself, has an annihilating effect. As

mentioned before, the indications of increasing

aridity are very vague, and one is more inclined to

attribute the degeneration of the forest to the

human interference. For grazing see BOELCKE, 1957.

KALELA (1941) has illustrated the retrogression

of forest, characterized by the movement of the

forest boundary towards the west by the following

schema:

The pampas vegetation occupies in this zone the

arid and open ground, e .g. valley bottoms, and

slopes above timber line. Also the places where the

forest has been exploited for timber are, due to

the altered microclimatic conditions, suitable ha

bitats for pampas vegetation. Along the shores and

on the old strand-lines an abundant occurrence of

plants characteristic of pampas vegetation was

noted. The pampas vegetation extends along glens

far into the Andean range, e.g. Pampa Linda at

the foot of Mt. Tronador. Here we find steppe

plants growing on the terminal moraines of the

glaciers of Mt. Tronador. These glaciers penetrate

deeply into the Nothofagus forests, cf. pl. 9.

Characteristic of these pampas fragments is

the growth of trees and shrubs which results in the

very pleasant park-landscape extending along the

valley bottoms and hills. Here there occur inter

alia, some shrubs which are missing west of this

zone, e.g. Fabiana imbricata and Diostea juncea.

The steppe vegetation occurring a hove the timber

line has up to an altitude of 1000-1 100 m the

character of grass-steppe, while the similarity to the

pampas vegetation increases downhill. Here grow:-

Blechnum penna-marina Polystichum adiantiforme P. mohrioides var. elegans Berberis empetrifolia Oardamine macrostachya Acaena ovalifolia ssp.

austral is Ovidia andina Lomatia obliqua Nothofagus antarctica Schinus patagonicus M aytenus boaria ])iscaria discolor ]) . trinervis

Buddleya globosa Fabiana imbricata ])iostea juncea Azorella sp. M ulinum spinosum Baccharis magellanica B. patagonica B. rosmarinifolia var.

subandina H aplopappus glutinosus H. pectinatus N assauvia aculeata Senecio coxi S. sericeonitens

Above the altitude of 1000-1 100 m the steppe

vegetation has the character of the dry variant of

altoandean steppe, v .i. Here we meet H ierochloe

juncifolia, Berberis empetrifolia, Ribes cucullatum,

Discaria sp., Acaena ovalifolia ssp. australis, and

some other Acaena spp. , Baccharis magellanica,

Ohilotrichum rosmarinifolium. · ( N. dombeyi ( A ustrocedrus ( A ustrocedrus ( N. dombeyi

F. . �tzroya Steppe-+ -+ -+ Austrocedrus -+N. dombey�-+

. . Saxegothea steppe N. antarct�ca N. antarctwa Laurelia


PL. I

L. BRUNDIN

Cerro Tronador seen from Casa Pangue. In the foreground Cortaderia dioica and Nothofagus dombeyi.

PL. 11

L. BRUNDJN

Plates I/ and J/l-Xerophilous vegetation in the valley of Rio Limay near Bariloche. Vegetation

of Stipa, Senecio, Di$caria serratifolia, Haplopappus pectinatus, Mulinum spi�osum.

PL. 111

L. BRUNDIN

PL. [V

L. BRUNDIN

The valley of Rio Limay. Vegetation of Mulinum spinosum, Stipa, Discaria serratifolia, Haplopappus pectinatus.

PL. V

L. BRUNDIN

Lake Mascardi. In background Cerro Catedral (2388 m); on the right, branches of Austrocedrus chilensis.

PL. V I

K . THOMASSON

Timber-line formed by Nothofagus pumilio on the slopes behind the · military camp at Bariloche.

PL. V I I

K. THOMASSON

R i o Manso and Lake Mascardi surrounded by the forest of Nothofagus domhey.

PL. V I I I

K. THOMASSON

Val ley of Rio M anso. In the foreground the terminal moraine of Ventisquero Tronador.

Ventisquero Tronador. The snout of the glacier with mounds of ice covered with debris on the surface.

On the left, terminal moraine which extends across the valley.

PL. I X

K . THO MASSO N

PL. X I

PL. X I I

Abo l'e: Lake Fri as . Jn the background Cerro Frias (1 800 m).

Opposite page: The shores of Lake Frias are bordered by luxuriant Valdivian forest ;

Nothofagus dombeyi is the dominating tree .

L B R U N J) [ N

PL. X I I I

l . . llR U N D l N

PL. X I V

L. B R U N D I N

Plates XI V and XV-Nothofagus pumilio forest on the slopes of Mt. Tronador. The trees

are hung all over with Vsnea beards. Bamboo is frequent in the ground vegetation .

PL. X V

L. ORUND i l\

PL. XVI

L. B I� UN D I N

Alpine tarn near Rigi. In the background Cerro Tronador.


Here and there grow interspersed in the andean steppe shrubs of N othofagus antarctica. They reach an altitude of about 1400 m a. s.-1.

The forest within this su bzone has on . the whole the same character as in the following zone with the single exception of the more frequent occurrence of conifers. At the low levels as well as at' the more elevated habitats it is a mesophytic forest. As a rule the fragments of forest adjacent to steppe show a considerable number of steppe plants in the ground vegetation.

In the. dry habitats close to the steppe grow the forests of A ustrocedrus chilensis but of these often only minor clumps remain for this timber is highly coveted. These groves are mainly found· along the streams and on the slopes of mountains, preferably on the northern slopes where they often occur in pure stands. A ustrocedrus chilensis has its main distribution located in the area between the pampas and N othofagus forests. Towards the west it becomes rare and is replaced by N othofagus dombey{

A ustrocedrus chilensis and N othofagus dombeyi are in some ways vicarious species, their presence being determined by water supply. Austrocedrus chilensis

is less demanding and able to survive under poorer conditions, on good soils with an abundant supply of water it is · injerior, however, when competition is present. In this transition zone, A ustrocedrus

chilensis often forms the undergrowth in N othofagus

dombeyi forests while the bamboo, which is so characteristic in the forests of the following zone, is lacking here. A ustrocedrus chilensis is in its vertical distribution confined to N othofagus dombeyi belt,_ i .e. to · the lower slopes of the mountains. In the very rainy regions in the west Austrocedrus chilensis

is rare and grows in the dry locations in N othofagus

dombeyi belt being without any competition in such places .

Besides A ustrocedrus chilensis also the rather eurytopic N othofagus antarctica participates in forming the border of the forests towards the steppe. It extends, like A ustrocedrus chilensis, along the streams and humid slopes into pampas in the east, though the main part grows at the alpine timber-line occupying the habitats unacceptable to Nothofagus

pumilio. Also on the malinas and swampy grounds on the bottom of valleys where the soil conditions

2 - 588829 Kuno Thomasson

are unfavourable for Nothofagus dombeyi and A ustrocedru.s chilensis these are both replaced by N othofagus antarctica. If the demanding N othofagus

dombeyi forest is destroyed by fire or some other devastation, its place is taken by N othofagus

antarctica. The deciduous N othofagus antarctica

covers extensive areas in the National Park of Nahuel Huapi. The total distribution along the eastern slopes of the Andean range extends over the whole woodland from its northernmost to the souther�most point. The deciduous N othofagus

pumilio has a similar distribution. N othofagus

antarctica is inferior in competition and it makes little demands on the habitats. It grows on all kinds of soils from the valley bottoms up to the timber-line and from the Patagonian steppe to the most humid parts of the National Park of Nahuel Huapi. Nothofagus antarctica on good soils is as a rule superseded by more competitive trees. It is the smallest of the N othofagus species occurring in the National Park. The length is about 3-6 m, seldom more than 10 m, while quite often it is only a tall shrub.

The forests on the northern slopes of the mountains within this subzone are composed of N otho

fagus dombeyi and A ustrocedrus chilensis. On the southern slopes we have forests of Nothofagus

pumilio. Intermingled in these forests are various small trees, e.g. Adesmia boronioides, Lomatia magel

lanica, L. obliqua, N othofagus antarctica, Schinus

patagonicus, Maytenus boaria, M. magellanica,

Colletia spinosa, Discaria foliolosa, D. serratifolia.

Mention should also be made of the conspicuous parasitic plants Phrygilanthus tetrandrus, Myzo

dendron linearifolium var. contractum, M. recurvum.

These forests are to be regarded as ramifications from the N othofagus dombeyi and N othofagus pumilio

forests respectively, growing in the west where they are fully developed. Therefore they �ll be treated in the following paragraph.

Along the forest skirts and in the open localities grows very rich brush-wood, many of the shrubs mentioned here have also been frequently noted in the forest vegetation. The following species should be specially mentioned: Aristotelia maqui, a widely distributed shrub, and Embothrium coccineum, a shrub with extraordinarily beautiful red blossoms,

Acta Phy togeogr. Suec. 42


Chusquea culeou, Berberis darwinii, B. heterophylla, B. parodii, Escalonia rubra, Ovidia pillopillo, M yoscilos oblonga, M aytenus magellanica, Discaria discolor, Buddleya globosa, Fabiana imbricata, Diostea juncea, Baccharis patagonica var. palenae, Senecio neaei. Entwining these shrubs are the lovely blooming Iianes M utisia decurrens and its var. patagonica, M. oligodon, M. retusa var. glaberrima, and Scyphanthus elegans.

The herbaceous flora includes many representatives from the flora of pampas. The following plants were noted: · -

Ophioglossum vulgatum v. valdivianum ·

N assella exserta Polypogon australis Sisyrinchum sp. Gavilea glandulifera Ranunculus peduncularis R. peduncularis v.

erodiijolius Acaena pinnatijida


Fragaria chilensis M f>,dicago lupulina Adesmia boronioides Astragalus palenae Polygala salasiana Oalceolaria llaimae Erigeron mysotis v.

fuegiae H ieraciurn glauci folium Senecio bracteolatus

Fig. 8. Mulinum spinosum on the

beach of Lake N ahuel Huapi. K. THOli!ASSON

S. goldsacki S. gymnocaulos S. linariaejolius

S. sericeonitens Solidago microglossa

On the damp soils the forest is in the main composed of N othofagus antarctica or N othofagus dombeyi, but Lomatia obliqua is also frequent. The shrub vegetation in such habitats is composed of Azara microphylla, Ribes sp ., Myoscilos oblongus, Berberis darwinii, B. heterophylla.

A few gleanings from the flora of herbaceous plants on the damp soils are as follows: -

Blechnum penna marina Alopecurus antarcticus Bromus unioloides Deschampsia elongata Glyceria multiflora Polypogon australis Stipa filiculmis J uncus bufonius J. depauperatus J. stipulatus 0 arex fuscula v. distenta

0. macloviana 0. patagonica 0. schedonautos Eleocharis albibracteata E. melanostachys Scirpus inundatus S. macrolepis Uncinia sp. Ranunculus minutiflorus Viola maculata v. pube-

scens f. grandidentata


V. reichei Oardamine glacialis Acaena ovalifolia ssp.

australis Epilobium valdiviense Primula farinosa v. ma

gellanica Plantago barbata P. carrenleu fuensis

M imulus glabratus v .

parvi florus Plagiobothrys

calandrinioides Hypsela reniformis Aster vahlii Erigeron spiculosus Senecio tri furcatus

Finally, there is the flora along predominantly sandy and coarsely graded morainic shores of the lakes to be considered. The shores are bordered by N othofagus pumilio and N othofagus dombeyi, and here and there are isolated trees of A ustrocedrus chilensis. The border of shrubs along the lake shores is composed of Berberis barilochensis, B. buxifolia, Escallonia virgata , M yrceugenella apiculata, M yrceugenia exsucca, Fuchsia magellanica, Coriaria ruscifolia, Baccharis lanceolata, B. rosmarinifolia var. subandina.

The herbaceous vegetation on the shores is K. THoMAssoN

composed of the following plants: - Fig. 9. Myzodendron linearifolium var. contractum on No

thofagus antarctica.

Ephedra andina H olcus lanatus N as sella exserta Polypogon australis J uncus burkartii J. stipulatus v. corralensis Oarex andersonii 0. darwinii 0. patagonica Ohloraea pichiquen Anemone multifida Ranunculus minutiflorus R. peduncularis R. repens Viola maculata v .

maculata V. maculata v. pubescens

f. grandidentata V. maculata v .

microphyllos f . septendrionalis

Sisymbrium officinale Gaultheria tenuifolia Pernettya poeppigii P. poeppigii v. linifolia Acaena cylindrostachya A. ovalifolia ssp.

australis Adesmia sp.

Arfona chubutensis Quinchamalium chilense Euphorbia protulacoides

ssp. nahuelhuapiana W endtia gracilis Pycnophyllum subulata Anagallis alternifolia v.

re pens Oollomia biflora Phacelia magellanica ssp.

eumagellanica Oalceolaria tenella Euphrasia meiantha M imulus cupreus M. luteus Veronica anagallis-

aquatica V. serpyllifolia Eryngium paniculatum H ydrocotyle chamaemoru,s · M ulinum morenonis M. spinosum V aleriana clarionifolia V. laxiflora V. virescens Hypochoeris sp. Senecio trifurcatus S. zosteraefolius Solidago chilensis

Subzone B

This subzone is included by CABRERA ( 1 953) in his Bosque Caducifolia. LJUNGNER (op. cit . ) designates it as "The second meridional zone-Mesophytic forest" .

The annual precipitation i s higher than i n the foregoing subzone, 1 300-1400 mm per annum . The vegetation is more luxuriant an� the forest more compact than in the east, though there are a few glades with steppe vegetation. Only on the bottoms of valleys, above the timber-line and along the shores do representatives of the xerophilous flora occur, the plant communities on the whole being of mesophytic character. Nothofagus dombeyi forests predominate, while small clumps of Austrocedrus chilensis occur on the parched localities, often with Lomatia obliqua forming a subordinate layer.

The mesophytic forest forms, together with the temperate rainforest, the narrow border of forests along the eastern slopes of the Andean range. This border is only about 60-70 km broad, its boundary towards the steppe coincides on the whole with


2 Flora and e;;etation

mm

uth in th

oler l ima , n id ra ly 1 w r amount of rain

fal l ar uffi ien for he grm h f f r t . I n th

w t hi bord r f N othofagus for t i bound d

y And an nowy p ak . Through h pa it i i n

contact with th Valdivian fore t t o which it i

1o ly akin fr m th flori tic point of view.

In t hi zon of m o hyti for N othofag

pur im

. l endtia gracilis on t h f Lak ahu l Huapi.

l ina and wamp ground . u h localitie are

w d d b N thofag anta1'Ctica. gr at part of

N othofag dombeyi for t growin on t he mountain

lope and along t he . hor of lak b long t

0 morrhiza-A denocaulon typ . l A L ELA .

Abov th belt of N othofagus dombeyi on t h

mountain i d t her o ur a lt o f N othofag

pumilio and N othofagus antaT tica. Th bord r lin

b tv n the belt of J-.. othofag domb yi and N otho

fag outh to t he north .

I n ituat d at

breadth f

m . Oft n

road nin toward h

dombeyi and th N otho-

un- lt N othofagus anlaT -

A ta Phy logeogr. ue . 42

t ica ft n gr v and pr ad i lf a Nothofagus

pu milio r tr at u pward and N othofagus domb yi

r tr a d v n' ards. LELA ( 1 94 1 ) ha int rpr t d

thi ph nom n n a r u lt f th all ged in rea ing

aridity. Thi impr ion i ti l l mor complicat d

Flora and vegetation 2 1

Fig. 1 1 . Quinchamalium chilense on th shor of Lake ahu l Hu pi.

by the effect of for t fire which favour N otho

fagus antat·ctica. N othofagu antarctica is one of

the pioneers on burnt locations l ike the birch

in candinavia. Be ide Nothofagu antantica thi

belt of teppe contain ome hrub , e .g . Lomatia

and Embothrium, and c l umps of Chusquea are a lso

frequent. On the very rainy lope in the we tern

part of the ahuel H uapi ational Park thi narrow

belt of teppe vegetation i lacking.

Returning to th Nothofagu dombeyi belt we

note here a variety of small tree and hru b ,

e.g. Nothofagu pumilio and Nothofagus antat·ctica,

both of them having their main di tri bution above

the belt con ider d. Furthermore Nothofagu.s

procera and the le demanding N othojag1t.s obliqua

al o occur, and in addition Lomatia hirs·uta, L.

obliqua, A ri totelia maqui, Embothrium coccineum,

A zara microphylla, chinu patagonicus, Discaria

erratifolia, D. trinervis, olletia spino a, Maytenus

boaria, M. di ticha, Myrceugenella apiculata, Myt·

ceugenia ex ucca, Myo cilus oblongus, Berberis dar-

winii, Escallonia rubra, E. virgata, Fabiana imbri

cata. Very charact ri tic for th N othofagu.., dombeyi

belt i the growth of the l iane Cynanchum lancifolium,

the common occurrence of bamboo ha already been

pointed out. The climbing grandiflorous compo ite

Mu ti ia retu a var. glaben·ima and M uti ia de

currens ar frequent. Among t the para ite pecial

mention hould be made of Phrygilanthu tetrandrus

while on the branche of Nothofagu the pecie of

M yzodendron are common. On the ame ho t

para ite al o Cyttaria, e .g . Cyttaria darwinii, C.

harioti, and C. hookeri. The e a comycete ac

company the g nu N othofagus even to the oppo ite

ide of the outhern hemi pher . The epi phytic ferns

are in the pre ent zone le prominent than in the

fol lowing one. The tree-trunk are covered mainly

with mo e and lichen . The ground vegetation

con ist of many plant common with the previou

zone. The damp habitat accom modate be ides

mo e , e .g. Dicranoloma mbu tum, a variety of

fern , e.g. Botrychium australe p. negeri, Ophio-

A cta Ph ytogeogr. Suec. 42


K. THOMASSON

Fig. 1 2. Chlorea piquichen on the moraine slopes at Lake N ahuel Huapi, Ba:riloche.

glossum crotalophoroides, Blechnum penna-marina, Oystopteris fragilis, Polystichum mohrioides. Furthermore we note: -

Alstroemeria aurantiaca Codonorchis lessonii Viola maculata

Loasa argentina Pernettya poeppigii Acaena spp. Fragaria chilensis V icia nigricans Gunnera magellanica

Phacelia magellanica Oalceolaria spp. Osmorrhiza berteroi 0. chilensis Adenocaulon chilense Leuceria thermarum Senecio otides S. spp.

In the open places, where inter alia a considerable number of steppe plants occur, we have noted: -Berberis empetrifolia Gaultheria tenuifolia

A cta Phytogeogr. Suec. 4-2

Pernettya mucronata v.

angustifolia

P. poeppigii Embothrium coccineum Nothofagus antarctica Schinus patagonicus M aytenus boaria lJiscaria discolor

Fabiana sp. Lippia sp. Azorella sp. M ulinum spinosum H aplopappus glutinosus N assauvia aculeata

The vegetation of the lake-shores consists of a mixture of different floristic elements. Of the plants growing on the shores of the lakes visited by us, the following should be named: -Ophioglossum vulgatum

v. valdivianum Poa bonariensis J uncus stipulatus Oarex aphyllus 0 . . macloviana Gaultheria phillyreaefolia Pernettya poeppigii Acaena glandulifera Adesmia boronioides Lathyrus cabrerianus Fuchsia magellanica

Gunnera chilensis W endtia gracilis Oollomia biflora Phacelia magellanica ssp.

eumagellanica 0 alceolaria tenella Ourisia poeppigii Eryngium paniculatum Baccharis patagonica Hypochoeris sp. Perezia linearis

The forest of N othofagus pumilio which extends above the belt of Nothofagus dombeyi has a more favourable light-climate than the later one. The bamboo which is of great importance in the N othofagu.s dombeyi belt reaches only the lower parts of the N othofagus pumilio belt. The forest of N othofagus pumilio comprises the alpine timber-line under a stretch of 2000 km along the eastern slopes of the Andean range. In the National Park of the Nahuel Huapi the timber-line is situated between 1400 and 1600 m a. s . -1. It drops towards the south and lies in Tierra del Fuego at an altitude of 0-500 m. In this region the forest of N othofagus pumilio reaches sea-level. The breadth of the N othofagus pumilio belt varies in the National Park between 400-500 m, its lower limit adheres to the Nothofagus dombeyi belt (v. s . ) . The deciduous Nothofagus pumilio is obviously confined to damp and cool habitats, because it shows no tendency towards colonizing the lower habitats. Nothofagus pumilio grows as a rule in pure woods which at the timber-line are interspersed with N othofagus antarctica. It was interesting to note the similarity between the Scandinavian alpine birch forest and the upper border of the Nothofagus pumilio-Nothofagus antarctica belt. Also here the influence of snow clearly manifests itself, the trees are curved and often damaged by


the fall of avalanches. A similarity to the Scandinavian mountains is contributed by N othofagus antarctica which often shows vegetative rejuvenation like Betula tortuosa. It has already been pointed out in connection with the meteorological survey, that the timber-line forms a zigzag line depending on exposure. The forest vegetation avoids bleak habitats and is confined to depressions and ravines and in such places the timber-line is forced upwards, as is the case on the leeward slopes of the mountains.

The forest of N othofagus pumilio with the above mentioned character also extends above the Valdivian rainforest in the west.

In the Nothofagus pumilio forest the shrubs of Berberis pearcei, B. linearifolia, Maytenus magellanica, and a low-grown form of Drimys winteri are common. Amongst the ground species the following

have been noted: Gavilea lutea, Ranunculus peduncularis var. erodiifolius, Pernettya poeppigii, Adesmia fernandezii, Epilobium sp., Valeriana lapathifolia, V. laxiflora, Erigeron philippi, Leuceria thermarum.

At the timber-line the trees are as a rule covered with magnificent U snea rnagellanica. In its bearded appearance, this part of the present belt resembles to a considerable degree the forests at the timberline on the East African mountains. The tree-trunks are covered with lichens, like Nephroma antarcticum var. lobuligerum, Pseudocyphellaria durvillei, P. gilva and Sticta damaecornis.

The N othofagus pumilio forests in the National Park of Nahuel Huapi belong partly to the Berberis pearcei type A. KALELA, and partly to the Osmorrhiza- Rha.coma type A. KALELA. The first one is represented by a mesohumid Berberis linearifolia variety and by a oligohumid Valeriana laxifolia variety.

Finally let us single out a few differences between the central zone of mesophytic forests, and the following zone of temperate rainforest. In both Nothofagus dombeyi constitutes the most important woody species. Since the Valdivian rainforest is characterized by an exceptional richness of its flora, its main distinguishing features are negative ones. The mesophytic forest lacks Saxegothea conspicua, Fitzroya cupressoides, Laurelia phillipiana, L. sempervirens, Gaultheria tenuifolia, Caldcluvia paniculata, W einmannia trichosperma, Amomyrtus

K. THOMASSON

Fig. 13. Gavilea lutea in the Nothofagus forest on the slopes

of Mt. Trona.dor.

luma, Myrceugenia chrysocarpa, M. ovata, Tepulia stipularis, Lomatia ferruginea, Pseudopanax laetevirens, Flotowia diacanthoides. All these species are frequent in temperate rainforest. Berberis pearcei, Drimys winteri, and M aytenus magellanica grow only in elevated habitats where the rainfall is plentiful. The conifers are of greater importance in the mesophytic forest than in the rainforest, especially in the case of A ustrocedrus chilensis which prefers dry places.



K. THOMASSON Fig. 14. Pampa Linda. In the background Cerro Tronador ( 3554 m).

3 . WE S T E R N Z O N E

The westernmost part of the National Park of Nahuel Huapi is covered with the luxuriant Valdivian rainforest. It forms a part of Distrito Valdiviano CABRERA, 1 953. LJUNGNER ( 1 939) has designated this area as ' 'The first meridional zone� Rain forest" . Our studies in this zone were carried out along the international route between Puerto Blest and Casa Pangue.

The climate of this zone is characterized by great amounts of precipitation, 2000-6000 mm per annum, cf. fig. 5.

On the valley bottoms the temperate rainforest occurs. Through the passes it is connected with the main area of this formation, i .e. with


the temperate rainforest which extends west of the Andean chain. This temperate rainforest belongs from the ecological point of view to the subhygrophytia, DEL VrLLAR, 1 929. There is a great variety of forest trees; many of them broad-leaved of magnolia- or laurel-type. RuBEL ( 1 930) has designated such vegetation as laurilignosa. The predominating tree is here N othofagus dombeyi as was the case in the mesophytic forest. Often intermingled in the N othofagus dombeyi forest grows Fitzroya cupressoides or, like at Puerto Blest, Saxegothea conspicua. In the dampest glens the forests of Saxegothea conspicua and Laurelia sempervirens occur, sometimes interspersed with N othofagus


dombeyi. In the last case Nothofagus dombeyi forms the highest layer of tree crowns at 40-45 m level and beneath at about 20-25 m lies the layer of Saxegothea and Laurelia crowns. In damp habitats there is often also a forest of Fitzroya cupressoides interspersed with Podocarpus nubigena.

Fitzroya cupressoides is without doubt the most magnificent tree in Patagonia. In its appearance it somewhat resembles Sequoia, and it attains a height of 50-60 m and an age of 2000-3000 years. From ancient times Fitzroya cupressoides has had its residence in this rainy region. MEYER (1950) has reported su bfossil finds from the western side of the Andean range in about 8 m depth, in a sandlayer of fluvioglacial origin. He estimates the age of these finds to at least 1 0,000 years. Fitzroya cupressoides is rare in the National Park where it grows in open stands on swampy soils in the valley bottoms between Brazo de Tristeza and Lake Frias. It prefers the dampest habitats, growing in such places in unmixed stands. On drier places it often grows together with N othofagus dombeyi, Saxegothea, Laurelia, and Pilgerodendron uviferum, e.g. in the neighbourhood of Puerto Blest.

The following list of the trees in the temperate rainforest is in no way a complete one.

Podocarpus nubigena Saxegothea conspicua A ustrocedrus chilensis Fitzroya cupressoides Pilgerodendron uviferum Drimys winteri Laurelia philippiana L. sempervirens Persea lingue Azara microphylla Eucryphia cordifolia Oaldcluvia paniculata W einmannia trichosperma Amomyrtus luma M yrceugenella apiculata

M yrceugenia chrysocarpa M. exsucca M. ovata Guevina avellana Lomatia ferruginea L. hirsuta L. obliqua Nothofagus antarctica N. dombeyi N. pumilio Schinus patagonicus Aextoxicum punctatum M aytenus boaria M. magellanica Pseudopanax laetevirens Flotovia diacanthoides

The tree-trunks are covered with a rich epiphytic vegetation of mosses and lichens and in addition the epiphytic ferns from the genus H ymenophyllum are abundant, e.g.· Hymenophyllum pectinatum, H. plicatum, and H. tortuosum. Furthermore among the epiphytes are Polypodium billardieri var. magel-

lanicum, P. trilobum, Serpyllopsis caespitosa, and last but not least, the charming Luzuriaga radicans belonging to lridaceae. Parasitic plants are also common, e.g. the widespread Phrygilanthus tetrandrus with its beautiful red flowers, and the genus M yzodendron, represented by various species and in great numbers. Moreover tree-trunks are decorated by lianes, like Mitraria coccinea, Sarmienta repens, Asteranthera ovata, Bogailia trifoliata, Griselinia r uscifolia, Dioscorea brachybotrya, Oampsidium valdivianum, and Hydrangea integerrima which is often as thick as one's arm.

One of the most noticeable features of these woods is the large number of arboreal and shrub by species which occur in the association. The luxuriant growth of bamboo is almost impenetrable. The bamboo, Ohusquea argentina and higher up Ohusquea culeou, often clings along the tree-trunks. The impenetrability of these woods is added to by shrubs and, should any space be left, by bamboo. Moreover, the growth of shrubs is luxuriant along the forest skirts and on the glades. In such localites we noted: -

Philesia magellanica Berberis darwinii B. linearifolia B. pearcei Azara lanceolata Gaultheria phillyreaefolia G. tenuifolia Pernettya mucronata v .

angustifolia Escallonia spp.

Ribes magellanicum Ovidia pillopillo Myrceugenia spp.

M yrteola nummularia v.

barneoudi

U gni molinae Fuchsia magellanica Embothrium coccineum Lomatia ferruginea M yoscilos oblongus A ristotelia maqui Orinodendron

hookerianum Ooriaria ruscifolia M aytenus disticha Buddleya globosa Desfontainea spinosa Pseudopanax laetevirens Baccharis racemosa B. umbelliformis

The ground is as a rule covered· with a dense mat of mosses, the most common of which are Acrocladium auriculatum, Breutelia subplicata, Dendroligotrichum dendroides, Dicranoloma robustum, Hypopterygium didictyon, Lepyrodon implexus, and Rhizogonium mnioides. Among the bryophytes there are many endemic species.

The herbaceous flora is rich, especially in the open localities. Ferns are represented by Ophioglossum vulgatum var. valdivianum, Asplenium sp.,

A cta Phy togeogr. Suec. 42


Blechnum chilense, B. magellanicum, B. pennamarina, Dryopteris sp. , Gleichenia quadripartitia, Gleichenia sp. , Polystichum sp. , and the imposing

Lophosoria quadripinnata, often as tall as a man.

Furthermore we note among the herbs Codonorchis lessonii, a conspicuous vernal flower with white

flowers giving to these forests an air of springtime

as in the woods of the northern hemisphere when

the Anemone nemorosa is flowering. Especially

along the streams grows Gunnera chilensis with huge

leaves, and moreover many Calceolaria species,

Gavilea cardioglossa var. patagonica, G. lutea, Chlorea sp. , Nertera depressa, Lobelia tuppa, Gentiana sp., the yellow-flowered Mimulus spp . , and

the fascinating saprophyte Arachnites uniflora, a

characteristic plant in summertime. Also present

are:-

U ncinia phleoides v.

trichocarpa Alstroemeria aurantiaca Azara lanceolata Viola reichei Gaultheria caespitosa G. phillyreaefolia Acaena ovalifolia Rubus radicans Vicia sp. Dysopsis glechomoides Oxalis corniculata 0. valdiviensis

Benthamiella aurea Ourisia ruelloides Osmorrhiza chilensis N ertera depressa V aleriana clarionifolia V. lapathifolia Adenocaulon chilense C otula acaenoides Lagenophora hirsuta M acrochaenium gracile P erezia palustris Senecio otites S. smithii

It must be emphasized that while botanizing in

the short time at our disposal, the personal factor

is of great importance and tends to the production

of a rather arbitrary list of plants from which quite

important species may have been omitted.

Owing to the tremendously rich flora it is rather

puzzling to arrange these forests into the schema

of Patagonian forest-types by A. KALELA (op . cit. ) .

The woods of Laurelia sempervirens are very close to

Asteranthera-N ertera type, and the woods of Saxegothea conspicua represent in addition also the

Desfontainea-Gaultheria type.

Along the lake shores, e .g. Lake Frias and the

innermost shores of Brazo Puerto Blest, a mixed

community of plants occurs. The following plants

have been noted:-


Equisetum bogotense A ustrocedrus chilensis Deschampsia antm·ctica Polypogon australis J uncus stipulatus Carex banksii 0. magellanica Cyperus sp. Scirpus cernuus Berberis linearifolia B. parodii Azara lanceolata A . microphylla Viola reichei Cardamine glacialis Gaultheria florida G. phillyreaefolia Pernettya mucronata v.

angustifolia P. poeppigii P. poeppigii v. nana Crassula moschata Fragaria chilensis Adesmia boronioides Escallonia alpina E. rubra E. virgata

A momyrtus luma, M yrceugenella apiculata M yrceugenia exsucca Tepualia stipularis U gni molinae Fuchsia magellanica Gunnera chilensis Lomatia obliqua Nothofagus antarctica N. pumilio M aytenus boaria Colletia spinosa Discaria discolor Arenaria serpens v.

andicola Anagallis alternifolia v.

re pens Desfontainea spinosa Prunella vulgaris Eryngium paniculatum H ydrocotyle chamaemorus H. chamaemorus v.

valdiviana H. marchantioides V aleriana chubutensis Baccharis sp. Senecio philippii S. tri furcatus

The marshy bottoms of the valleys are character

ized by Saxegothea conspicua, Fitzroya cupressoides, Pilgerodendron uviferum, Drimys winteri, Laurelia sempervirens, Myrceugenia ovata, Nothofagus antarctica

Other parts of these woods, although not really

marshy are nearly always very moist, and such

places have a rich and exuberant flora of arboral

and shrubby species, e.g. Chusquea spp. , Myrceugenella apiculata, and N othofagus antarctica.

In addition to the above mentioned indigenous

trees and shrubs there occur the following moisture

loving plants which are characteristic of marshy

places: -

J uncus cyperoides J. diemii J. involucarius Carex andersonii C. darwinii Eleocharis melanostachys

Scirpus inundatus Escallonia virgata Aster vahlii P erezia palustris Senecio jistulosus S. tri furcatus

Naturally peat-moses, Sphagnum spp. are frequent.

When moving upwards towards the higher re

gions, the N othofagus dombeyi forest becomes grad-


Fig. 1 5 . Acaena ovalifolia.

ually impoverished in tree species. According to

LJUNGNER ( 1939) the disappearance occurs in the

following sequence: W einmannia trichosperma, Flotowia diacanthoides, Lomatia ferruginea, Pseudopanax laetevirens, Azara lanceolata. No personal

observations were made to prove this statement.

At about 1200-1300 m the Nothofagus dombeyi forest is replaced by N othofagus pumilio forest. At

about 1400 m the bamboo vegetation ceases. Drimys winteri which is a tall tree at low altitudes forms

in the N othofagus pumilio belt only low shrubs, a

lifeform which has sometimes been designated as var.

andina. Interspersed here and there, within the

Nothofagus pumilio belt are a few trees of Saxegothea conspicua, Fitzroya cupressoides, Pilgerodendron uviferum, A ustrocedrus chilensis, and naturally

N othofagus antarctica. The shrub-layer is made up

of shrubs which obviously thrive in these light

forests. During the ascent to Rigi and Paso

Vuriloche the following shrubs and dwarf shrubs

were noted: -

Drimys winteri Berberis buxifolia B. empetrijolia B. linearifolia B. microphylla B. pearcei Azara microphylla Pernettya sp.

K. THOMASSON

Escallonia alpina Ribes cucullatum R. magellanicum M yrteola barneoudii Embothrium coccineum Discaria sp. Desfontainea spinosa Baccharis magellanica

The fern Blechnum chilense is of common occurrence,

but lianes are missing, only Ast1·anthera ovata ought

to be mentioned. Due to the favourable light

conditions and thin growth of shrubs, the ground

vegetation is rich. Here grow: -

Lycopodium paniculatum Blechnum auriculatum Agrostis montevidensis v.

submutica Festuca sp. Oarex nigra Tristagma nivale Alstroemeria aurantiaca

Sisyrinchium graminifolium

S. junceum Gavilea sp . Pogonia sp. Viola sp . Gaultheria caespitosa Pernettya poeppigii



Empetrum rubrum Acaena adscendens A . ovalifolia A. tenera Fragaria chilensis Rubus radicans Vicia sp.

Gunnera magellanica Erigeron philippii Gnaphalium andicola G. serranoi G. spicatum

M acrachaenium gracile Lagenophora hirsuta L. nudicaulis Perezia brachylepis

-P. pedicularifolia P. pili/era Senecio acanthifolius S. chilensis S. subdiscoideus S. subpubescens S. trifurcatus 8. triodon

The two Sisyrinchium_ species are originally steppe plants and prefer dry locations.

On precipices, there grow: -

Hymenophyllum tortuosum

Serpyllopsis caespitosa Sisyrinchium sp.

Berberis darwinii Draba sp.

Acaena t.enera Escallonia rubra

0 olletia spinosa Galceolaria spp . .

Ourisia sp.

Pinguicula antarctica Abrotanella linearifolia Perezia spp.

Senecio trifurcatus

Along the rapid streems there is Juncus lesueurii,

Caltha andicola, Gunnera magellanica, N othofagus

antarctica, Calr;eolaria tenella, Mimulus spp. , Ourisia

alpina, 0. poeppigii, Pinguicula antarctica, Perezia

megalantha.

The timber-line is situated at about 1600 m a. s . -1. Its somewhat varying altitude is due partly to the exposition lying higher on the northern than on the southern slopes, a feature partly dependent on edaphic conditions. The timer-line is formed by dwarf-sized Nothofagus pumilio, which is often pressed down by the snows. Below Rigi N othofagus

pumilio has been observed growing in creeping shape -like Betula nana in the alpine belt of the Scandina-

vian mountains. Interspersed among the N othofagus

pumilio a few specimens of N othofagus antarctica

occur here and there and just below the timber-line the trees · often bear magnificent beards of U snea

on their trunks and branches, cf. pl. 14 & 15 . In dry places at the timber-line there occur·

Festuca sp.

Garex similis Tristagrria nivale Sisyrinchium

graminifolium S. iunceum Berberis empetrifolia Gaultheria caespitosa Rubus radicans

Ribes cucullatum Discaria sp.

Baccharis magellanica Ghilotrichium

rosmarinifolium Perezia pilifera Senecio subdiscoideus S. triodon

The open marshy locations at the timber-line , e.g. the mire in the Paso Vuriloche, have a vegetation dominated by Carex spp . , e .g. Carex canescens,

C. decidua, C. magellanica, C. banksii. In addition there is Sphagnum fimbriatum. Here and there grow the shrubs of the white-flowered composite, Chilio

trichum rosmarinifolium, and the trees of N othofagus

pumilio of low growth. The following have also been noted: -

Deschampsia sp.

J uncus stipulatus M arsippospermum

grandiflorum M. philippi Oreobulus obtusangulus Schoenus antarcticus Ranunculus peduncularis V icia magellanica Gunnera magellanica N anodea muscosa Euphrasia subexerta

V aleriana carnosa V. chubutensis V. fonckii V. laxiflora Aster vahlii Erigeron philippii Leuceria sp.

Perezia lactucoides Senecio hieracium S. parodii Troximon poeppigii

4. T H E A L P I N E B E LT

The andean semidesert extends between the altitudes of 1600 and 2 100 m. This regia alpina

has been designated by CABRERA ( 1953) as Distrito altoandino austral.

As it has been already mentioned the alpine belt contains nothing to correspond with the alpine meadows in the Alps. The vegetation is sparse and


occurs in patches between rocks and blocks of stone. This is the natural home of the condor ( V ultur

gryphys L. ) , which is often rather inquisitive about the activities of a wandering botanist in these lofty regions with its most wonderful panorama extending over the Andean mountain scenery.

In the following I have listed plants noted


from the alpine belt. Observations were carried out mainly in the surroundings of Rigi at the border between the Argentine and Chile. An attempt has been made to distinguish between dry and damp habitats, and the lists are by no means exhaustive ones. A full list would have needed a much longer stay and studies of many localities in the Andean range.

In dry habitats have been noted: -

Oalamagrostis erythrostachya

Dantonia violacea F estuca scabriuscula Hordeum pubiflorum Poa sp. Trisetum sp. Oarex canescens Berberis empetrifolia Viola auricolor V. tridentata V. reichei Gaultheria caespitosa Pernettya poeppigii v.

nana P. pumila v. crassifolia P. pumila v. leucocarpa Empetrum rubrum Adesmia longipes A . retusa Oxalis adenophylla M elandrium aff.

magellanicum 0 olobanthus subulatus Euphrasia subexerta Oalceolaria biflora Ourisia fragrans Azorella mesetae

M ulinum microphyllum Oruckshanksia glacialis V a,leriana fonckii V. philippiana A brotanella emarginata A. linearifolia A. trichoachaenia Aster vahlii Baccharis magellafl,ica Ohaetanthera villosa Ohiliotrichum

rosmarinifolium Gnaphalium affine G. nivale H ypochoeris humilis Leuceria papillosa Lucilia araucana N assauvia argentea N. dentata N. lagascae N. lagascae v. lanata N. pygmaea v. intermedia Perezia pedicularifolia Senecio fulieti S. poeppigii S. subdiscoideus S. tri furcatus S. triodon

The superficial similarity between the alpine vegetation of the Andean range and the Scandean range is conspicuous. Empetrum rubrum grows in large patches and it resembles to a considerable extent the crowberry Empetrum hermaphroditum, except that the fruits are red. Instead of the lingonberry there is Pernettya pumila, an ericaceous plant having two varieties, one with red and the other with white berries. Like the dwarf-birch is the habit of the trailing Baccharis magellanica, while M elandrium aff. magellanicum is most suggestive of the M elandrium apetalum of the Scandina vian mountains. As is the case in Scandinavian mountains,

solifluction is also of common occurrence in the alpine belt of the Andean range.

Besides vascular plants mentioned above and below there are a great number of lichens, like Oladonia laevigata, 0. vicaria, Neuropogon auran

tiacoater, Stereocaulon speciosum; and mosses, like Bartramia aristata and Plagiochila remotidens,

growing on the ground and on stones. In addition the following list of plants growing in

damp habitats contains many forms which resemble their relatives in Scandinavian mountains, e.g. Pinguicula, Euphrasia, Oardamine, Epilobium,

Ranunculus, etc. Among the plants growing on wet soil I 4ave noted the following ones: -

Deschampsia atropurpurea

Festuca acanthophylla Poa obvallata M arsippospermum

philippii Schoenus antarcticus Barneoudia major v.

patagonica Oaltha appendiculata 0. dioneifolia 0. sagittata Ranunculus peduncularis R. semiverticillatus Oardamine nivalis Epilobium magellanicum

E. nivale Oalandrinia rupestris Primula farinosa v.

magellanica Euphrasia andicola E. chrysantha Ourisia alpina 0 . coccinea 0. poeppigii 0. pygmaea Pinguicula antarctica V aleriana fonckii Aster vahlii H ypochoeris andina Lagenophora nudicaulis Perezia variabilis

Thus concludes the brief sketch of this botanical foray which extended from the arid semi-desert in the east to the humid rain forest in the west, and from the shores of the bluest lakes up to the heights where lie eternal snow and ice. It is indeed unfortunate that words alone cannot justly describe the full beauty, grandeur and wealth of colour of the various regions contained within the National Park of Nahuel Huapi. Certainly it is quite impossible to present the wonders of this area merely by the presentation of lists of plants, or by prosy descriptions of vegetation types, even when aided by black and white photographs.

In conclusion the following list of the vascular plants mentioned above is given in alphabetical order, and this, it is hoped, will pro:vide some systematic orientation to the foregoing account.



LIST OF VASCULAR PLANTS

Abrotanella emarginata CASS. - Compos. - linearijolia GRAY - trichoachaenia CABR.

Acaena adscendens V AHL. - Rosac. acroglochin BITT.

cylindrostachya R UIZ et P A v.

glandulifera BITT.

ovalijolia spp. australis BITT. pinnatifida RUIZ et PAV.

poeppigiana GAY pseudopoeppigiana KALELA

splendens HooK et ARN.

tenera ALBOFF

Adenocaulon chilense LEss. - Compos. Adesmia boronioides HooK f. - Legumin. - campestris (RENDELE) RoWLEE

- jernandezii PHIL.

- palenae PHIL.

- retusa GRISEB .

Aextoxicum punctatum Rmz et PAv. - Aextoxic. Agrostis leptotricha DESV. - Gramin. - montevidensis var. submutica DO ELL.

- pyrogea SPEG.

Alopecurus antarcticus V AHL. - Gramin. Alstroemeria aruantiaca DoN. - AmaryJlidac. Amaryllis elwessi (WRIGHT) TRAUB. - Amaryllidac. Amomyrtus luma (MoL . ) 1LEGR. et KAUS. - Myrtac. Anagallis alternifolia var. repens ( D'URv.) KNUTH. -

Primulac. Anarthrophyllum patagonicum SPEG. - Legumin. - rigidum {GILL . ) BENTH.

Anemone decapetala L. - Ranunculac. - multifida PoiR.

Arachnites uniflora PHIL. - Burmanniac. Arenaria serpens var. andicola GILL. - Caryophyllac . Aristotelia maqui L'HERIT. - Eleocarpac. Arjona chubutensis Dus:EN. - Santalac. - tuberosa CA v.

Aster vahlii (GAUD.) HooK et ARN. - Compos. Asteranthera ovata (CAv.) HANST. - Gessnerac . Astragalus cruckshanksii (HooK et ARN.) GRIS.

Legumin . - palenae (PHIL. ) REICHE

A ustrocedrus chilensis (DoN.) FLORIN et BouTELJE -

Cupressac. Azara lanceolata HooK f. - Flacourtiac. _:_ microphylla HooK f. Azorella mesetae SKOTTSB. - Umbellif. - nucamentacea (PmL.) HAUM.

- trifurcata (GAERT. ) HooK

Baccharis lanceolata H. B. et K. - Compos. magellanica (LAM.) PERS.

- patagonica HooK et ARN.


Baccharis patagonica var. palenae (PHIL.) REICHE

- racemosa (RUIZ et PAv. ) DC.

- rosmarinijolia var. subandina HEER

- umbellijormis DC.

Barneoudia major var. patagonica (SKOTTSB.) LouRT.

- Ranunculac. Benthamiella aurea SKOTTSB. - Solanac. Berberis barilochensis JoB. - Berberidac.

buxifolia LAM. cuneata DC. darwinii HooK

empetrijolia LAM. heterophylla Juss.

linearifolia PHIL.

- parodii JoB.

- pearcei PHIL.

Blechnum auriculatum CAv. - Polypodiac. - chilense METT.

- magellanicum METT.

- penna-marina (PoiR.) KuHN

Boquilia trifoliolata (DC.) DEENE - Lardizabalac. Botrychium australe ssp. negeri (CHRIST.) CLAUSEN -

Ophioglossac. Bromus macranthus MEY. - Gramin. - stramineus DESV.

- trinii DEsv.

- unioloides H. B. K.

Buddleya globosa HOPE - Loganiac. Oajophora silvestris (PoEPP.) URB. et GILG. - Loasac. Oalamagrostis erythrostachya DEsv. - Gramin. Oalandrinia rupestris BARN. - Portulacac. Oalceolaria biflora LAM. - Scrophulariac.

lanceolata CA v.

llaimae KR.ANzL.

polyrrhiza CA v.

tenella POEPP. et ENDL.

Oaldcluvia paniculata (CAv.) DoN. - Cunoniac. Oaltha appendiculata PERS. - Ranunculac. - dioneifolia HooK f. - sagittata CA v.

Oampsidium valdivianum (PHIL. ) SKOTTSB. - Bignoniac.

Oardamine glacialis DC. - Crucifer. integrifolia PHIL.

- macrostachya PHIL.

- nivalis GILL.

Oarex aematorrhyncha DESV. - Cyperac. - andersonii BooTT

- aphyllus KUNTH

banksii BOOTT

canescens L. darwinii BOOTT

decidua BooTT


Garax juscula var. distenta (KUNZE) KuKENTH.

- macloviana D'URv.

- magellanica LAM. - nigra (L.) RErCH.

- patagonica SPEG.

- schedonautos (STEUD . ) em KA.LELA

- similis D'URv.

- subantarctica SPEG.

Gerastium arvense L. - Caryophyllac. Chaetanthera villosa DoN . - Compos. Ghiliotrichum rosmarinifolium LESS. - Compos. Ghloraea piquichen LINDL. - Orchidac. Chusquea argentina P ARODI - Gramin. - culeou DESV.

- quila KNTH.

Codonorchis lessonii (BRONGN. ) LrNDL� - Orchidac. Colletia spinosa GMEL. - Rhamnac. Golliguaya integerrima GILL et HooK - Polygalac. Collomia bijlora (Rurz et PAv. ) BRAND . - Polemoniac. Golobanthus subulatus ( D'URv.) HooK f. - Caryo-

phyllac . Coriaria ruscijolia L. - Coriariac. Corynabutilon bicolor (PHIL. ) KEARNEY - Malvac. Cortaderia selloana (SCHULT.) AscH. et GRAEB .

Gramin.

Gotula acaenoides (HooK et ARN.) ALBOFF. - Compos. Grassula moschata FoRST . - Crassulac. Grinodendron hookerianum GAY - Elaeocarpac. Cruckshanksia glacialis POEPP. et ENDL. - Rubiac. Gynanchum lancifolium HooK et ARN. - Asclepiadac. Cystopteris fragilis (L.) BERNH. - Polypodiac. Danthonia violacea DEsv. - Gramin. Deschampsia antarctica DEsv. - Gramin. - atropurpurea (W AHL.) ScHHUL.

- elongata (HooK) MUNRO

Desjontainea spinosa Rurz et PAv. - Loganiac. Dioscorea brachybotrya POEPP. - Dioscoreac. Diostea iuncea ScHAu. - Verbenac. Discaria discolor (HooK f. )BENTH. et HooK -

Rhamnac. - foliosa MrERS.

- serratifolia (VENT. ) BENTH. et HooK

- trinervis (POEPP. ) REICHE

Draba australis R. BR. - Crucif. Drimys winteri FoRST. - Magnoliac. Dysopsis glechomoides (RICH. ) MuLL. ARG.

Euphorbiac. Eleocharis albibracteata NEES - Cyperac. - melanostachys ( D'URv. ) CLARKE

Elymus patagonicus SPEG . - Gramin. Embothrium coccineum FoRST. - Proteac. Empetrum rubrum V AHL. - Empetrac. Ephedra andina PoEPP. - Ephedrac. Epilobium magellanicum PnrL. et HAUSSKN. -

Onagrac. - nivale MEYEN

Epilobium valdiviense HAUSSKN.

Equisetum bogotense H. B. K. - Equisetac. Erigeron myosotis var. fuegiae VIERH. - Compos. - philippii ScH. BrP.

- spiculosus HooK et ARN.

Eryngium paniculatum CAv. et DoMB. - Umbellif. Escallonia alpina PoEPP. - Saxifragac. - rubra (Rurz et PAv.) PERS.

- virgata (Rurz et PAv.) PERS.

Eucryphia cordijolia CAv . - Eucryphiac. Euphorbia portulacoides ssp. nahuelhuapiana CROIZAT

- Euphorbiac. Euphrasia andicola (GILL . ) BENTH. - Scrophulariac. - crysantha PmL.

- meiantha CLOS.

- subexerta BENTH.

Fabiana imbricata Rurz et PAv. - Solanac. Festuca acanthophylla DEsv. - Gramin. - argentina (SPEG.) PARODI

- monticola PHIL.

- pallescens (ST. YVEs) PARonr

- scrabriuscula PHIL.

Fitzroya cupressoides (MoL.) JoHNSTON - Cupressac. Flotovia diacanthoides LEss. - Compos. Fragaria chilensis DucH. - Rosac. Fuchsia magellanica LAM. - Onagrac. Galium aparine. L. - Rubiac. Gaultheria caespitosa P. et E. - Ericac. - phillyreaefolia (PERS.) SLEUMER

- tenuijolia (PHIL.) SLEUMER

Gavilea cardioglossa var. patagonica (GARAY) CoRREA

- Orchidac.

- glandulijera (POEPP.) CORREA

- lutea (PERS. ) CoRREA

Geranium sessiliflorum CAv. - Geraniac. Gilia gracilis (DouGL.) HooK - Polemoniac . Gleichenia quadripartitia (Porn.) MoO RE - Gleicheniac. Glyceria multijlora STEUD . - Gramin. Gnaphalium affine D'URV. - Compos.

. - andicola PHrL.

- nivale PHIL. - serra.noi PHIL.

- spicatum LAM. Griselinia ruscijolia (CLos . ) TAUB . - Cornac. Guevina avellana MoL . - Proteac. Gunnera chilensis LAM. - Haloragac. - magellanica LAM.

Habranthus bagnoldii HERB.

Halopapp'l,ts glutinosus CASS. ___.:.. Compos. - prunelloides DC. - pectinatus PHrL.

Heliothropium paronychioides DC. - Boraginac. Hieracium glaucifolium POEPP. - Compos. Hierochloe iuncifolia (HACKEL) PARODI - Gramin. Holcus lanatus L. - Gramin. Hordeum comosum PRESL. - Gramin.



Hordeum murinum L. - pubijlorum HooK f. Hydrangea integerrima (HooK et ARN. ) ENGL.

Saxifragac. Hydrocotyle chamaemorus CRAM. et ScHLEqHT -

Umbellif. - chamaemorus var. valdiviana (PHIL.) REICHE

- marchantioides CA v.

Hymenophyllum krauseanum PHIL. - Hymenophyllac. - pectinatum CAv. - tortuosum HooK et GREY. Hypochoeris andina (DC.) BENTH. et HooK - Compos. - humilis (PHIL.) REICHE

Hypsela reniformis (H. B. K.) PRESL. - Campanulac. I mperata condensata STEUD . - Gramin. Juncus bufonius L. - Juncac. - burkartii BARROS - cyperoides LAHARPE

- depauperatus PHIL.

- diemii BARROS - involucarius STEUD apud BuCHENAU

- lesueurii BoL.

- stipulatus N EES et ME YEN

- stipulatus var. corralensis (PHIL) BucHENAU

Lagenophora hirsuta PoEPP. - Compos. - nudicaulis (CoMM. ) Dus:EN

Lathyrus cabrerianus BURKART - Leg�min. - magellanicus LAM. Laurelia philippiana LoosER - Monimiac. - sempervirens (Rurz et PAv . ) TuL.

Leuceria papillosa CABR. - Compos. - thermarum (PHIL . ) REICHE

Loasa argentina URB . et GrLG. - Loasac. Lobelia tupa L. - Campanulac. Lomatia jerruginea (CAv.) R. BR: - Proteac. - hirsuta (LAM.) DIELS

- obliqua R. BR.

Lophosoria quadripinnata {GMEL . ) CHR. - Cyatheac. Lucilia araucana PHIL. - Compos . . Luzula chilensis NEES et MEYEN - Juncac. Luzuriaga radicans Rurz et PAY. - Iridac. Lycopodium paniculatum DESV. - Lycopodiac. Macrachaenium gracile HooK f. - Compos. Marsippospermum grandiflorum (L. f . ) Hoort f .

Juncac. - philippi (BUCHENAU) HAUM . Maytenus boaria MoL. - Celastrac. - disticha URB. - magellanica HooK f.

Medicago lupulina L. - Legumin. Melandrium aff. magellanicum FENZL.

Caryophyllac. Microsteris gracilis ( DouGL.) GREENE

Mimulus cupreus REGEL - Scrophulariac. - glabratus var. parvijlorus (LINDL. ) GRANT - luteus L.

A cta Phytogeogr. Suec. 4 2

M itraria coccinea CA v. - Gessneriac. Mulinum microphyllum PERS. - Umbellif. - morenonis (KUNTZE) SPEG.

- spinosum (CAv.) PERS. Mutisia decurrens CAV. - Compos. - decurrens var. patagonica (PHIL.) BLAKE

- oligodon POEPP. et ENDL.

- retusa var. glaberrima PHIL.

Myoscilos oblonga Rurz. et PAv. - Santalac . Myrceugenella apiculata (DC.) KAus. - Myrtac. Myrceugenia chrysocarpa (BERG) KAus. - Myrtac. - exsucca (DC.) BERG

- ovata (HooK et ARN.) BERG

Myrteola nummularia var. barneoudi BERG - Myrtac. Myzodendron linearifolium var. contractum SKOTTSB.

- Myzodendrac. - recurvum VAN TIEGH.

Nanodea muscosa GAERTN. - Santalac. Nardophyllum obtusifolium HooK et ARN. - Compos. - parvijolium PHIL.

Nassauvia aculeata (LESS.) PoEPP. et ENDL. -Compos. argentea PHIL.

dentata GRISEB . glomerulosa (LAG . ) DoN.

N assauvia lagascae (DoN�) BE NTH. et Hoo:rr

- lagascae var. lanata (PHIL.) SKOTTSB. - pygmaea var. intermedia (PHIL.) CABR. Nassella chilensis (TRrNrus) DEsv. - Gramin. - exserta PnrL. - Gramin. N ertera depressa BANKS. - Rubiac. Nothojagus antarctica (FoERST. ) 0ERST. - Fagac. - dombeyi {MIRB . ) BLUME

- obliqua (MIRB . ) BL UME

- procera 0ERST. - pumilio {POEPP. et ENDL.) KRASSER Ophioglossum crotalophoroides W ALT. - Ophioglossac. - vulgatum var. valdivianum (PnrL.) LICHTENST.

Oreobolus obtusangulus GAUD . - Cyperac. Osmorrhiza berteroi DC. - Umbellif. - chilensis HooK

Ourisia alpina PoEPP. et ENDL. - Scrophulariac. - coccinea PERS.

- fragrans PHlL.

- poeppigii BENTH.

- pygmaea PHIL.

- ruelloides (L. f . ) Dus.

Ovidia andina (POEPP. et ENDL. ) MEISSN. -Thymelac.

- pillopillo (GAY) MEISSN. Oxalis adenophylla GrLL . - Oxalidac. - corniculata L. - valdiviensis BARN.

Perezia brachylepis PnrL. - Compos. lactucoides (V AHL. ) LEss.

- linearis LESS.


Perezia megalantha SPEG.

- palustris (PHIL. ) REICHE

- pedicularifolia LEss.

- pilifera HooK et ARN.

- variabilis (PHIL.) REICHE

Pernettya mucronata var. angv,stifolia (LINDL. ) REICHE

- Ericac. - poeppigii (DC.) KLOTZSCH.

- poeppigii var. linifolia SLEUMER

- poeppigii var. nana SLEUMER

- pumila var. crassifolia (PHIL.) SLEUMER

- pumila var. leucocarpa (DC.) KLAUSEL

Persea lingue NEES. - Laurac. Phacelia magellanica ssp. eumagellanica Hydro-

phyllac. Philesia magellanica GMEL. - Liliac. Phrygilanthus tetrandrus (Rmz. et PAv. ) ErcHL. - .

Loranthac. Pilgerodendron uviferum (DoM . ) FLOR. - Cupressac. Pinguicula antarctica V AHL. - Lentibulariac. Plagiobothrys calandrinioides (PHIL. ) J OHNST.

Boraginac. - verrucosus (PHIL. ) JOHNST.

Plantago barbata FoRST. - Plantaginac.

- carrenleufuensis SPEG.

Poa bonariensis (LAM. ) KuNTH. - Gramin. lanuginosa Pom.

- ligularis N EES.

- obvallata STEUD. Podocarpus nubigena LINDL. - Podocarpac. Polygala salasiana GAY - Polygalac. Polypodium billardieri var. magellanicum ( DREV . )

CHR. - Polypodiac. - trilobum CAV. Polypogon australis BRONGN. - Gramin. Polystichum adiantiforme (FoRST) SM. - Polypodiac. - mohrioides (BoRY) PRES.

- mohrioides var. elegans (REMY) CHR.

Primula farinosa var. magellanica (LEHM.) HooK f. -Primulac.

Prunella vulgaris L. - Labiat. Pseudopanax laetevirens (GAY) SuM. - Araliac . Pycnophyllum lechlerianum (PmL. ) RoHRB . - Caryo-

pyllac.

Quinchamalium chilense MoLINA - Santalac. Ranunculus minutijlorus BERT. - Ranunculac. - peduncularis SM.

- penducularis var. erodiifolius (GAY) REICHE

- repens L. - semiverticillatus PHIL.

Ribes cucullatum HooK et ARN. - Saxifragac.

- magellanicum Pom.

Rubus radicans CAv. - Rosac. Sarmienta repens Ruiz et PAv. - Gessneriac. Saxegothea conspicua LINDL. - Podocarpac. Schinus patagonicus (PmL.) JoHNST. - Anacardiac.


Schinus polygamus (CAv.) CABR. Schoenus antarcticus HooK f. - Cyperac. Scirpus cernuus V AHL. - Cyperac. - inundatus POIR.

- macrolepis PHIL.

Scyphanthus elegans DoN. - Loasac. Senecio acanthifolius HoMBR. et J AQ. - Comp()s.

bracteolatus HooK et ARN.

- chilensis LESS. - coxi PHIL.

,---- fistulosus PoEPP.

- goldsacki PHIL.

- gymnocaulos PHIL.

- hieracium REMY

- julietii PHIL.

- linariaefolius PoEPP.

- microcephalus PHIL.

- neaei DC. - otites K UNZE

- parodii CABR.

- philippii ScH. BIP.

- poeppigii HooK et ARN.

- sericeonitens SPEG.

- smithii DC. - subdiscoideus ScH. BIP.

subpubescens CABR. trifurcatus (FORST.) LESS.

triodon PHIL.

zosteraefolius HooK et ARN.

Serpyllopsis caespitosa (GAUD . ) CHR. - Hymeno-phyllac.

Sisymbrium officinale (L.) ScoP. - Cruciferac. Sisyrinchium graminifolium LINDL. - Iridac. - junceum MAY. - striatum SMITH

Solidago chilensis MEYEN - Compos. - microglossa DC. Spartium junceum L. - Legum. Stipa filiculmis DEL. - Gramin. - hirtiflora HACK. - humilis CAv. - neaei NEES

- speciosa TRIN.

Tepualia stipularis (HooK et ARN.) GRISEB. - Myrtac. Triptilion achilleae DC. - Compos. Tristagma nivale PoEPP. et ENDL. - Liliac. Troximon poeppigii (DC.) WmEM. - Compos. Ugni molinae TURcz. - Myrtac. Uncinia phleoides var. trichocarpa (C. A. MEY.) CLARK

- Cyperac. Valeriana carnosa SM. - Valerianac. - chubutensis SPEG.

- clarionifolia PHIL.

- fonckii PHIL.

lapathifolia V AHL.

- laxiflora DC.


34 H ydrography

V aleriana philippiana BRIQ. - virescens CLos. Wendtia gracilis MEYEN - Oxalidac. W einmannia trichosperma CA v. - Cunoniac. Veronica anagallis-aquatica L. - Scrophulariac. - peregrina ssp. xalopensi.<� (H. B. K.) PENN. - serpyllifolia L. Vicia magellanica HooK f . - Legumin. - nigricans HooK et ARN. Viola auricolor SKOTTSB. - Violac.

Viola maculata var. maculata CA v.

- maculata var. microphyllos f. septendrionalis (KALELA) WEIBEL

- maculata var. pubescens f. grandidentata (KALELA) SPARRE

- reichei SKOTTSB. - tridentata MENZ. Vulpia eriolepis ( DEsv.) BLOM. - Gramin. - megalura (NUTT. ) RYDB.

VI. H YD R O G RA P HY

The inter-oceanic watershed, which runs through

the national park, coincides north of Mt. Tronador

with the frontier between the Argentine and Chile,

it runs about 35 km west of the general axis of

elevation. South of Mt. Tronador the watershed

turns abruptly to the east and forms a pass between

the Lakes Gutierrez and Mascardi. Then it makes

a wide curve through the Paso . de Villegas and

coincides again with the frontier about 75 km

south of Mt. Tronador. Consequently the major

part of Nahuel Huapi National Park belongs to the

catchment-basin of Rio . Limay which forms the

Rio Negro to run into the Atlantic.

The Nahuel Huapi region is rich in lakes; most of

them, covering an area of l l49 km2, belong to the

Rio Limay drainage area. The most expansive of

them is Lago Nahuel Huapi (740-767 m a. s . -1 . ) ,

which i s the largest lake i n the national park, its

area being 523-531 km 2• The second largest lake is

Lago Traful (790 m a. s.-1 . ) which also belongs to the

same drainage area, as do the following: Lago

Espejo (748-:-816 m a. s.-l.) 40-44 km2, Lago Cor

rentoso (748-8l l m a. s . -L) 13.3-27 km2 and Lago

Gutierrez (760-800 m a. s . -l . ) 14.34-28 km2• The

three last named lakes have a catchment-basin of

290, 337 and l l2 km2 respectively. Besides the

already mentioned lakes there are a lot of smaller

lakes and lakelets belonging to the drainage area

of Rio Limay, e.g. Lago Frias (770 m a. s . -l . )

5.3 km2, Lago Moreno 10 .61 km2, Lago Falkner

(926 m a . s . -1 . ) .

A Cta Phytogeogr. S u e c . 42

As will appear also from the following figures the

statements about the areas and altitudes vary

considerably depending on the different sources of

information. The same matter is illustrated by

fig. 4 in LARSSON, 1940, as to the altitude of Mt.

Tronador.

The southwestern and minor part of the National

Park belongs to the basin of Rio Manso which via

the Rio Puelo discharges its waters into the Pacific .

Rio Manso arises from the glaciers on the eastern

slopes of Mt. Tronador and after a meandering

course through the Cordillera falls into the Estuario

de Reloncavi to the west of the mountains. Among

the lakes belonging to its catchment-basin is the

large Lago Mascardi (796 m a. s . -l . ) its area being

36 km2, together with a number of minor lakes,

e.g. Lago Guillelmo (820 m a.s.-1. ) , Lago Fonck

( 740 m a. s . -1 . ) , Lago Hess (730-750 m a. s . -1 . ) ,

Lago Steffen (590 m a. s . -1. ) , and Lago Martin.

Chemical conditions have a central position in

the study of aquatic biota. It is to be regretted that

the chemical conditions of the waters in Nahuel

Huapi National Park have not hitherto been

studied, since they are of major importance to a

knowledge of the ecology of plankters. Unfortun

ately there was a stoppage in the ferry service across

Lake Frias during our j ourney; therefore we were

forced to leave our equipment for chemical analyses

behind in Chile. No investigations corresponding

to those in Chile and Peru (LoFFLER, 1959) have

been carried out; only a few figures based on the

H ydrography 35

water-samples we had with us when returning from Nahuel Huapi National Park were analysed and

·are quoted here from L6FFLER, 1959.

Conduc-Lake tivity Ca Mg Na K Si02

Nahuel Huapi 23 .5 1 .2 1 . 1 2.0 0.7 1 2 p.p.m.

Frias 18 .6 Hess 34 4

The low value for calcium was expected: in SALMI (1941 ) we find a figure of 2.58 p .p.m. for CaO present in a soil sample from the beach of Lago Espejo. In the bottom deposites of Lago Nahuel Huapi CoRDINI ( 1939) found a CaO content from 2 . 1 to 2.24 p.. p. m. For comparison the following d1,1ta from BLACK (1929) concerning the sediments of North American lakes may be of interest: hardwater lakes 20-25 p. p. m. and medium to soft water lakes 0.6-3 .7 p. p. m .

Some lakes are fed b y glacier-waters rich in ooze, e.g. Lake Frias, and the influence of the large quantities of ooze on the chemical composition of the water is of great interest. It is well known from pedology that clays affect the properties of water largely through the exchangeable ions on or near the surface of colloidal particles. If the water is free from electrolytes, or contains only very low concentrations, as glacier waters do, a proportion of exchangeable ions dissociate from the surface of the clay particles a:nd form a diffuse double layer around the colloidal particle. While the anionic groups form the outer layers of the colloidal particle, the corresponding cations arrange themselves with respect to the particle and the surrounding water in a distribution depending on their own ionic activity, and their degree of hydration. The distribution is also governed by the presence of electrolytes. The water carries a net positive charge, and the surface of the clay particle carries a net negative charge, due to isomorphus replacement in the lattice. This negative charge is caused by the dissociation of hydrogen ions into the surrounding water from hydroxyl groups situated at the edges of the crystal lattice. The above mentioned double layer prevents the free interchange of cations between the surface

of clay particle and the bulk of the solution. Anion exchange differs from cation exchange in that it is not a quantitative exchange. The total quantity of anions held depends on the closeness with which they are held and the power of the cationic double layer to displace the anionic groups. Phosphate anions are held close to the . positive charges, so that the double layer due to the negative charges cannot expel them. Hence, these anions cannot be washed out by distilled water, but need to be displaced by other strongly adsorbed anions, e.g. bicarbonate or silicate. It is probable that the only inorganic form of phosphate taken up by plants is the H2P04-. Calcium clays adsorb water-soluble phosphates · [Ca(H2P04)2] , the phosphate being converted to the relatively insoluble dicalcium phosphate CaHP04 at the surface of the · clay particles. But this phosphate is sufficiently soluble to maintain a high enough concentration of HP04 -

ions in solution to allow an adequate proportion of H2PO 4- ions to be present. This is so only under acid conditions; under neutral or alkaline conditions dicalcium phosphate becomes converted to still more insoluble phosphates, which maintain a still lower HP04-- or P04--- ion concentration. The proportion of soluble phosphate ions in the H2P04-form becomes very small if the pH rises above 8.

Nitrates are only weakly held by clay particles and can be removed completely by reducing the electrolyte concentration around the particles, so that the double layer d11e to the negative charges on the colloidal particles interpenetrates and replaces that due to the positive charges.

It is evident that the influence of ooze dispersed in glacier-lakes on the chemical conditions of the water is a rather complicated problem . It seems to me that . apart from the mechanical action of the floating ooze this chemical effect is of great importance for the biota in such lakes. Interactions between the clay particles dispersed in water and substances dissolved in water may be a factor to be reckoned with in connection with the ·theory launched by BRUNDIN ( 1956) concerning the influence of glacial errosion on the productivity of waters.


36 Lakes

VII. LA K E S

1 . L A K E N A H U E L H U A P I

Lake Nahuel Huapi forms the grand central feature of the National Park. In shape it resembles a gigantic amoeba with enormous tentacles extending in all directions to form the reaches and fjords. The lake is almost completely encircled by verdureclad hills and mountains. The coastline which is extremely indented is of great length. It consists of a succession of bays and coves, shingle and sandy beaches, perpendicular rocky cliffs, steep promontories, wooded isthmuses and peninsulas. It is studded with islands and islets, the majority covered with dense vegetation. The luxuriant forests of A ustrocedrus and N othofagus surrounding its solitary fjords contrasts with the scanty aquatic vegetation of macrophytes. The scarcity of the latter is partly dependent on the very limited shallow-water areas due to the precipitous slopes of the lake basin and partly to the considerable wave action along the shores of Lake Nahuel Huapi. Only in a few sheltered and shallow locations has the occurrence of thinly grown Scirpus californicus (S . A. MEY) BRITTON been noted. Moreover the occurrence of Isoetes savatieri FRANCHET should be mentioned.

a. Morphometry

From the investigations carried out by CoRDINI

( 1939) on Lake Nahuel Huapi we are able to form a picture of the morphometry of the largest lake in the national park. Lake Nahuel Huapi is the only one which has hitherto been studied in this respect, though only partially. More thorough investigations with the aid of echo-sounders have been planned.

Altitude m Area sq. km Maximum depth m Maximum length km Maximum width km Length of shore line km Shore development (-8-)

2 v;; Acta Pl1yto_geogr. Suec. 42

764 529 438 i

74.4 10.2

357.35

4.3

S is the length of shore line and c:x is the area of lake. , The term shore development indicates the form of shore line according to HALBFASS ( 1923) and STR0M ( 1937) referring to the proportion between the actual shoreline and circumference of a circle covering the same area as the lake surface.

Lake Nahuel Huapi has about the same depth as the Norwegian lake Mj0sa (449 m), which is the fourth deepest of the European lakes. The first three places are also occupied by Norwegian lakes, the deepest being Hornindalsvatn, 5J4 m. With its fjord-sceneries Lake Nahuel Huapi is suggestive of Norwegian lakes. But considering heat economy it is better to compare Lake Nahuel Huapi with Lago di Como (v.i . ) which is also a deep lake, 410 m. The position of Lake Nahuel Huapi in the Andean range corresponds to the position of Lago di Como in the prealpine mountains. Like Lake Nahuel Huapi which extends its enormous tentacles among the mountains, Lago di Como penetrates with its fjord-like arms into the Bergmasque Alps. Even the temperature of water is evidence of resemblances; viz. the mean temperature of winter ( T mw) is for Nahuel Huapi 7 .2° and for Lago di Como 6.8°, the mean temperature of summer (T ms) for both of lakes is 8.5°.

The depth relations of Lake Nahuel Huapi are illustrated in fig. 17 , they are based on data given by CoRDINI ( op. cit.) considering the hitherto sounded 281 km2 of the total area of 529 km2• In spite of incompleteness these figures will give a

1 MARCHETTI, 1 949, reports that the depth of the lake

is not well known as yet but it is in some places greater

than 1000 m. With regard to the geological history of the

Nahuel Huapi region this s tatement seems to be scarcely

plausible, though not absolutely out of the question. Such

great depths occur in lakes which are of quite different

origin than Lake Nahuel Hu,api, viz. in lakes which occupy

tectonic troughs, e.g. Lake Baikal and Lake Tanganyika.

Until the report of MARCHETTI has been confirmed by

exact soundings it seems to be j ust a story about bottom

less lakes.

Lakes 37

Fig. 1 6. Lake Na.huel Huapi. In the foreground Eryngium pani

culatum.

representative picture of depth relations. Compare also curves in CoRDINI, 1939, p. 32 and profiles on p. 33. The approximate extension of the trophogenic layer is indicated in fig. 17 by dots. The transparency values for Lake Nahuel Huapi lie between

Fig. 1 7 .

0 100 200

m

J A. Percentage volume curve for Lake N ahuel

· . Hu�pi consideri'ng the hitherto ·sounded parts.

· B. · HY!>sographic curve for subsurface horisontal ·areas down to 3 00 m depth.

K. THOMASSON

10 to 15 m. The transparency can be used as a rather good indicator for the estimation of the thickness of the trophogenic layer.

b. Thermal relations

The thermal relations are of profound importance for aquatic life, both through their direct effect and indirect influences due to thermal stratification and its consequences. Without any doubt the thermal relations hold a key position in our knowledge of lake biota.

From the thermal point of view the majority of lakes in the National Park belongs to tropical lakes, their surface temperatures being always above 4 °C.

After a bathe in the cold waves of Lake Nahuel Huapi one is fully convinced that the term "tropical" is at times eminently inappropriate. It is often quite misleading since it carries the thoughts involuntarily to the tropics, viz. to the belt between the tropics . of Cancer and Capricorn. However, it seems to be rather difficult to · substitute this classical term, introduced by FoREL. It may · be


38 Lakes

remarked here that a change of terms and their identity would only cause confusion as have most attempts to restrict the meaning of terms which have been used in sensu lata. One needs only to consider the confusion about the littoral-concept (cf. Du RrETZ, 1940) and the more current discussion on the trophic system in limnology. For the characterization of "tropical" lakes situated outside the tropic circles, the term subtropical lakes (cf. R UTTNER, 1940, p. 34) is to be preferred in regard to thermal conditions.

There occur also temperate lakes in the Nahuel Huapi National Park, e . . g. Lago Frias, which in some Winters has a solid ice cover. Obviously the thermal conditions of this lake are influenced by the cold glacier-�ater from the glaciers of Cerro Tronador. At low .air temperatures permanent ice cover also forms on lakelets and tarns, and they show an inverse stratification characteristic of temperate lakes during the winter stagnation period.

The thermal conditions of Lake Nahuel Huapi have been investigated by CORDINI (op. cit. ) . Temperature relations during the year are shown in fig. 18 from CORDINI, 1939.

For Lake Nahuel Huapi CoRDINI gives the following temperatures:

Maximum Minimum Amplitude

Surface water Bottom water

As· has been mentioned above Lake Nahuel Huapi is a subtropical lake, its waters being disturbed throughout their entire depths but once each year. That is the so-called process of overturning and circulation which occurs in July-August, during the winter in the southern hemisphere. Lake N ahuel Huapi is to · be classified as a holomictic lake, because its period of thermal stratification is followed by a regular period of total circulation. It belongs to the so-called hypolimnic lakes of the second order, i .e. the thermal stratification is well developed and characterized by a stable hypolimnion during the summer, and by the temperature of the bottom water being above the temperature of maximum density, which is according to STR0M, 1945, variable with depth thus:


O m

50 100 1 50

3.93°

3.87 3.82 3 . 7 7

200 m 300 400 500

3 . 7 1°

3.60 3.49 3.39

The total turnover of heat in a body of water is equivalent to the difference between the highest and lowest storage of heat. The maximum amount of heat in Lake Nahuel Huapi is storaged in December, minimum in August. During the circulation period the total heat content of Lake Nahuel Huapi is according to a rough estimation about 701 ,076 x 106 ton calories. As the sun gathers strength and the accumulation of heat increases towards the summer the lake becomes stratified and remains so throughout the entire summer. It may be noted that no epilimnion of considerable duration and stability is established. The heating of deeper layers is the result of heat transfer by windgenerated turbulences and currents.

The heat content of Lake Nahuel Huapi in summer has been calculated to be about 829,619 x 106

ton calories. Hence the lake has accumulated about 128,543 x 106 ton calories during the period of heating, a quantity equivalent to the thermal- capacity of 16 .5 million tons coal (the coal import of Sweden was in 1955 about 3.2 million tons).

It may be emphasized that all figures on heat economy of Lake Nahuel Huapi are rather approximate, partly because the morphometry of the lake is not fully investigated, and partly because there exist only a few temperature series. In spite of their shortcomings these figures may be of some interest in a survey of thermal conditions in Lake Nahuel Huapi.

It may be remarked in this connection that random temperature series often result in quite misleading pictures of thermal conditions depending on the submerged depression . individuality. This fact has been demonstrated inter alia by JoHNSSON (1946) in a close study of the Swedish lake Klammingen. The general basin of this lake contains three isolated depressions with submerged bars separating the deeps, each depression has its own thermal conditions, the anual heat budget varing between 1 1 ,800 and 27,000 cal. fcm2• The same is valid for the Finnish lake-system of Paijanne which has a very complicated pattern showing

Lakes 39

A G O S T O S E P T I E M B R E � � · � · � � w � · � r � • w � Qm Om I I I

I I

100 " 100 "

-1----1---

200 • -1--

-I-

-I-

-I-300 · -1--

..__ -

NVIERNO FASE DE I Toda la mass de agua tiene

a uniform e. temperatur

N O V I E M B R E 7° 8° 9° 10° 1JO 12° 13° 14° Om

100 .

2 0 0 ·

300 ·

O m

100 .

300 ·

1r I I

" I

3Q --

-

-

-

1--1--1--1--1--1-1--

-'--

FASE Las capas s

DE PRIMA VERA uperiores siguen ea

a estratificacion ler· nza a nofarse en toda

asa de agua.

lenfandose; I mica comie

la m

E N E R O 7° 8° 9° 10° 11° 12° u·

,.

/ l/

,v

r-:- 552 r-r-

'-r-f-r-

-r-

-r--........J 1--

FA Las capas

S.E DE VER.A NO

superiores m uy calien· a tificaci6n termiea bien

en toda la mass de agua tes; la estr mareada comienza a prepara.rse la c:apa de

sa/to termlco". ..

200 "

3 0 0 "

O m

1 0 0 ..

20 0 •

'200 "

30 0 ·

r�

--

--

1--

1-'--

1-r--

1-r--

>-r-

-r-'--1--

-

PRIN El agua

CIPIO DE PRIMA VERA eomienza. a ealen tarse en

e superior; !as capas infe· es tienen tempera fura

unifor1n e.

la part rior

D I C I E M B R E 7° 8° 9° 10 11° 12 13° 14u 15u 16°

V J

1---�

1--r-

1-r

-r-'--1--

I

� l/

4 Q PR.INC/P/0 D

Las capas superio

E VER.ANO res m uy callenci6n termica, tes; la estra tifiea

bien mareada masa de

�

en toda la agua.

rJ -� 6.2 -

-

-

-

1--

1--

1--

f--

-:-

FASE DE OTONO

La cap pletam de agu

a de "sa/to term/eo", com · ente formada; u n a mssa a ca.liente, superpuesta en 6n brusca a otra de agua a estratificaci6n termica. ende a desapa recer.

fransici frfa; I

ti

400 � --Fig. 18. Temperature relations of Lake Nahuel Huapi. From CoRDINI, 1 939.

r--'---A cta Phytogeogr. Suec. 42

40 Lakes

annual heat budgets from 50,000 to 73,000 cal.fcm2

in different parts of the lake-system, accordmg to the report by SrMOJOKI ( 1956) . Also . concerning Lake Nahuel Huapi with its dendritic pattern of basin, the individuality of different parts is a factor to be considered. Of course there are no separating bars but all sinousities and fjords communicate with deeper layers. Here probably the orientation of the fjord-like arms of the lake in relation to prevailing winds is of soine importance for heat eco:Q.omy, cf. JoHNSSON (op. cit. p. 144) who has eminently elucidated the role of wind on the storage of heat in Lake KHimmingen.

It is well known that heating of the lake only by insolation is negligible, except at the surface; the warming is mainly effected by the wind, which distributes the warmer water from the surface to the deeper strata. The distribution of p.eat solely by conduction would require tremendous time because the thermal conductivity of water is at 18°0 as low as 0.0014 cal.fcm x sec x grade.

At the warming up of a lake we have to deal with two contrar.y forces, viz. on one hand the insola� tion which warms up the surface water and thus decreases its density and tends to confine the in� fluence of the wind to the surface. On the other hand the wind induced currents strive to derange the thermal stratification produced by incident radiation and to maintain a complete circulation. The result is the distribution of heat in a lake.

In the foregoing paragraphs the approximate quantity of heat which is distributed in Lake Nahuel Huapi during the heating , period by the work of wind has been calculated. In the following an attempt has been made to compute the amount of work done by the wind per unit area. This work has been performed against gravity, viz. the warmer, lighter water must be forced down against the resistance of the colder, denser water and mingled with it.

'

For computation of the work of the wind in warming lakes the following· well�known formula given by BrRGE, 1915� has been used.

W = RT X Z x ( l - Dn)

W is work expressed in gram-centimeters pe� square centimeter of surface of lake

A cta Phytogeogr . . Suec. 42

RT is reduced thickness, i.e. the thickness of a stratum if its area is made equal to that of the lake and its sides vertical, reduced thickness is expressed in cm

Vn- m R T = --

A o

A0 is the area of the lake at the surface

Vn- m is the volume of a stratum between two selected levels

Z is the distance from the surface of the lake to the middle of the stratum expressed in cm

Dn is the density of water at any selected temperature Since Lake N ahuel Huapi is a subtropical lake the

factor 1 - Dn which is valid for temperate lakes has been substituted by Dm - Dn

Dm gives the density of water at the lower limit of temperature considered.

The computation has been made for single strata and the sum of the several products amounted to about 7735.4 gcmfcm2 of work done by the wind on the water surface to bring about the thermal conditions characteristic for Lake Nahuel Huapi during the time of maximum storage of heat. It thus appears that work amounting to 409,202.66 x 106

kgm has been expended in the distribution of the heat gained by the entire lake which amounted to about 128,543 x 106 ton calories.

Heat budget of a lake is a good and often applied exponent for comparison of thermal conditions in different lakes. Hence I have tried to compute the heat budget of Lake Nahuel Huapi in spite of the many deficiencies of available material.

Since the heat budgets are computed as differ� ences between the temperature extremes they are higher in temperate lakes than in tropical and polar lakes. The heat budget is thus something quite different from the capacity of the lake for absorbing heat, which is highest in tropical lakes, and in temperate lakes of second order depending on the effect of vo1ume and depth. On the other hand, in temperate lakes of first order where the storage of heat is �ot li�ited by' an insufficient depth, the maximum storage

. of heat is a function of climate

and location. The most useful conceptions of heat budgets are

the following two.

1 . Tll;e summ�r .J?.eat �.corn� or ,the wind-distributed heat, i .e . the · amount of heat· · needed to rise the

Lakes 41

lake from an isothermal condition at 4°0 up to the highest observed summer heat content.

2. The annual heat budget, i .e . the total amount of heat that enters the lake between the time of its lowest and its highest heat content.

By means of computing the number of calories entering per unit area during the heating of the body of the water from its maximum temperature, we obtain the annual heat budget of a lake.

Annual heat budget = Dm ( Tm8 - Tmw)

Dm is the mean depth of the lake in cm which is obtained by dividing the volume of the lake by its area

Tm8 is the mean summer temperature Tmw is the mean winter temperature

For computing the volumes of horizontal strata, the following formula according to ALSTERBERG

( 1935, p. 68) has been used.

h V = 2 (al + a2)

F is the volume of stratum h is the vertical depth of stratum a1 is the area of the upper surface of a stratum in sq. m

a2 is the area of the lower surface of a stratum in sq. m

The basin of Lake Nahuel Huapi has steep slopes, therefore the following formula given by BIRGE

(1915) has been used for computing the mean temperature of the lake (for a basin with gently sloping sides the formula of ScOTT is to be preferred) .

Tm is the mean temperature of a lake T1, T2 , T3 etc . are the temperatures at respective

levels bounding the various strata of water. T n is the temperature at the lowermost level of the

last stratum P1, P2, P3 etc. are the percentages of the entire lake

volume which the respective strata constitutes P n is that percentage of the entire lake volume which

the lowermost stratum constitutes

The annual heat budget of Lake Nahuel Huapi calculated according to the formulas given above ranges from 24,300 to . 26,300 cal.fcm2• For comparison a few values on annual heat budgets have

been compiled from the literature and given in table 1 . More data on annual heat budgets is to be found in the recent book by HuTCHINSON ( 1 957). The first six lakes in the table are temperate lakes, and the last three are subtropical lakes in regard to their thermal conditions. Bodensee forms a transition between temperate and subtropical lakes.

TABLE 1 . Heat. budgets of a selected group of lakes

Lake

Baikal Inarinjarvi Pohj ois-

Kilpisjarvi

Mj0sa Klammingen Vat tern Bodensee Lac Leman Lago di Como Nahuel Huapi

Area sq. km

31,500 1 ,050

23 26 1 1 .2

1 ,900 538 582 146 529

Depth Annual heat budget

me d. max. cal.fcm2

730 1940 57,300-72,200 60 35,000

13 45 36,000 187 449 37,000-41,�00

12 .5 37 17 ,700 39 128 30,000-32,000 9 1 . 7 252 29,000

152 310 30,000-37,000 185 440 32,000

438 24,300-26,300

In this table it is evident that the amount of the annual heat budgets for various lakes situated in different latitudes is rather uniform. The subtropical lakes in this table have more in common with temperate lakes than with tropical ones, the only difference from the temperate lakes lies in the lack of a winter stagnation period. Proceeding to the lakes in the tropics they often have low budgets because the seasons differ little from one another in respect of temperature changes.

When we compare the maximum heat storage in tropical, subtropical and temperate lakes instead of annual heat budgets a quite different picture emerges. The subtropical lakes, viz. Lac Leman (60,000 cal.fcm2) , Lago di Oomo (82,000 cal.fcm2) , Nahuel Huapi (83,000 cal.fcm2), and the tropical lake Danau Manindjau, Sumatra (244,000 cal.fcm2) obviously have higher heat content than the temperate lakes. (It must be noted that to obtain comparable values the amount of heat corresponding to 4°0 has been subtracted. ) For comparison a few figures from STR0M, 1944, on maximum heat storage in temperate lakes-Mj 0sa 23,000, Hornindalsvatn 19,000, and Bessvatn 8000 cal.fcm2• Lunzer Untersee has a storage of heat of about


42 Lakes

12,300 cal.fcm2, and in another temperate lake, Baikal the storage of heat reaches 34,500-42,300

cal.fcm2•

c. Plankton

Due to strong winds and heavy waves only one plankton sample was collected, on January 25,

1954 from Brazo Campanario, a relatively shallow bay situated inside the peninsula of San Pedro. The maximum depth of Brazo Campanario is 43 m, mean depth is 21 .3 m, and area 7.9 km2• The water temperature was 12.5°0. Collection of plankton from the waters mentioned below was made with nets of fine-mesh silk, Defour bolting silk no. 25 standard having been used. All the samples from lakes in the National Park accounted for in the present . paper were collected from off-shore waters by means of boats, in Lake Guillelmo a raft was used.

Naturally this sample from Brazo Campanario is by no means characteristic of the whole lake. The plankton stocks in different parts of such a divided basin often show some degree of discrepancy in relation to each other.

The plankton sample from Brazo Campanario was dominated by Dinobryon divergens, Rhizosolenia eriensis, Gastropus· styli fer, Keratella cochlearis,

and Boeckella gracilipes. Moreover a Philodina sp. and Staurastrum spp. were abundant. The composition of the plankton is given in the following list. There is no doubt that this is an oligotrophic lake.

Oyanophyta

Ohroococcus minutus (KuTz . ) NXG.

Dactylococcopsis ellipsoideus (SCHRODER) TElLING

Gomphosphaeria lacustris CHOD . Merismopedia glauca (EHRENB.) NXG.

Pyrrophyta

Peridinium willei HurTF. -KAAs

Ohrysophyta

C H R Y S O P H Y C E AE

Dinobryon cylindricum lMH. D. cylindricum var. palustre LEMM. D. divergens lMH. D. divergens var. schauinslandii (LEMM.) BRUNNTH. D. sertularia EHRENB. Salpingoeca elegans (BACHM.) LEMM.


. B A C I L L A R I O P H Y C E AE :

Oyclotella stelligera var. elliptica FRENG.

Diatoma elongatum (LYNGB .) AG.

Melosira ambigua (GRUN.) 0. MULL. - rare M. pseudogranulata A. CL. Rhizosolenia eriensis H. L. SMITH - also broad

forms, 22.5 p, Surirella sp.

Synedra ulna var. danica (KuTz. ) GRUN.

Ohlorophyta

V o L v o c A L E S

Eudorina elegans EHRENB. Gloeococcus schroeteri (CHon. ) LEMM. Paulschulzia pseudovolvox (TElLING) SKUJA Tetraspora lacustris LEMM.

C H L O R O C O C C A L E S

Botryococcus braunii KuTz. Ooelastrum proboscideum BoHLIN - rare Orucigenia rectangularis (NA.G.) GAY Dictyosphaerium pulchellum W oo:O Hojmania lauterbornei (ScHMIDLE) WrLLE - THo -

1\-IASSON, 1957, fig. 1 : 6 Kirchneriella obesa (W. WEST) SCHMIDLE Micractinium sp. - Fig. 20: 10 Oocystis solitaria WrTTR. 0. submarina var. variabilis SKUJA Pediastntm boryanum (TURP. ) MENEGH. - rare • C O N J UGALE S

Oosmarium botrytis (BORY) MENEGH . 0. connatum BREB. 0. contractum var. ellipsoideum (ELFV.) G. S. WEST Sphaerozosma vertebratum (BRilm .) RALFS - also

broad forms, 27-28 p, Staurastrum longipes var. evolutum (WEST & WEST)

THOM.

St. planctonicum var. ornatum (GRONBLAD) TElLING

- GRONBLAD, 1 938, fig. 3 St. rotula var. smithii (G. M. SMITH) THOM. St. sebaldi var. ornatum fa. planctonica (LUTKEM.)

TElLING

St. trachytithophorum WEST & WEST Staurodesmus megacanthus (LUND.) THUNM. Std. sellatus fac. jacobsen# TElLING

Protozoa

Raphidiophrys sp. Raphidocystis tubijera PEN. Vorticella sp.

Rotatoria

Oollotheca sp.

Oonochilus unicornis Rouss. Gastropus· styli fer IMH.

Lakes 43

Keratella c. cochlearis ( GossE) M onostyla (Lecane) lunaris (EHRENB.) Philodina sp. Polyarthra vulgaris CARLIN Pompholyx sulcata HuDSON Synchaeta pectinata (EHRENB. ) S. sp. Testudinella cf.. coeca var. lermaensis AHLSTR. Trichocerca birostris (MrNKIEWrcz) T. brachyura GossE

Cladocera

Bosmina chilensis DADA Y Alonellct, sp. - a species close to A. exigua (LrLLJEB. )

Copepoda

Boeckella gracilipes DADAY

In · the foregoing paragraphs some thermal features common to Lago Nahuel Huapi and Lago di Como have been pointed out. Therefore it may be of some interest to corn pate the plankton of these two lakes. The plankton sample from Lago di Como was collected on July 7, 1955 .near the little town of Como, in northern Italy; it was dominated by Eudorina elegans and K eratella quad rata. Furthermore Fragilaria crotonensis, Staurastrum cingulum

var. obesum, Staurastrum sebaldi var. ornatum fa. planctonicum, Cyclotella comensis, and Ulothrix sp. were frequent in the sample. Of the Crustacea Me

socyclops leuckarti is the dominant.

Bacteriophyta

Pelogloea bacillifera LTB . Sphaerotilus natans KuTz. __:__ rare

Cyanophyta

Anabaena flos-aquae (LYNGB. ) BREB. - few colo-nies

Ghroococcus limneticus LEMM. Goelosphaerium kuetzingianum NAG. Dactylococcopsis irregularis G. M. SMITH Gomphosphaeria lacustris CHOD . Merismopedia major (SMITH) GEITL. Microcystis flos-aquae (WITTR. ) KrncHN. - rare

Euglenophyta

Phacus longicauda (EHRENB. ) DuJ.

Pyrrophyta

Geratium hirundinella - fa. austriacum (ZEDERB. ) BACHM.

Peridinium cinctum (0. F. M.) EHRENB.

Chrysophyta

0 HR Y S O PH Y C E AE

Bicoeca oculata ZACH.

B A C I L L A R I O PH Y C E AE

Asterionella formosa HASS. Gyclotella comensis GRUN. Fragilaria crotonensis KrTTON M elosira varians AG. Tabellaria flocculosa (ROTH) KNUDSON

Chlorophyta

V o L V O C A L E S

Garteria ovata J AKOBSEN Elakatothrix gelatinosa WrLLE Eudorina elegans EHRENB. Gloeococcus schroeteri (CHOD .) LEMM.


Botryococcus braunii KuTz. Goelastrum microporum NAG. Dictyosphaerium pulchellum W oon Kirchneriella obesa (W. WEST) ScHMIDLE Nephrocytium limneticum (G. M. SMITH) SKUJA N. lunatum W. WEST Oocystis borgei SNow 0. elliptica W. WEST 0. lacustris CHOD. Pediastrum boryanum (TURP.) MENEGH. P. duplex ME YEN P. limneticum THUNM. Scenedesmus arcuatus LEMM. S. longispina CHOD. S. quadricauda (TURP.) BREB. - rare Schroederia setigera (ScHRODER) LEMM. Tetraedron caudatum var. longispinum LEMM.

UL O T R I C HA L E S

Ulothrix sp. ster. - planktic Planktonema sp.

C O NJ U G A L E S

Gosmarium depressum var. achondrum (BoLDT) WEST & WEST

G. subtumidum var. klebsii (GuTw.) WEST & WEST Staurastrum cingulum var. obesum G. M. SMITH

radiation 3 + 3, 3 + 4, 4 + 4 S. sebaldi var. ornatum fa. planctonica (LUTKEM. )

TElLING

Protozoa

Acanthocystis sp.

Rotatoria

Branchionus quadridentatus HERMANN - incl. var. rhenanus (LAUT.) , var. cluniorbicularis (SKORIK.},


44 Lakes

and var. brevispinus (EH ENB.) which cannot be kept apart in the present population

Kellicottia longispina (KELLICOTT) Keratella c. cochlearis (GossE ) K. q. quadrata (0. F. M.) Monostyla (Lecane) lunaris (EHRENB.) Polyarthra major (BURCKH.) P. vulgaris CARLIN Pompholyx sulcata HUDSON Testudinella patina (HERMANN) Trichocerca birostris (MINKIEWicz) T. sty lata ( GossE)

Cladocera

Ohydorus sphaericus 0. F. M. Daphnia longispina 0. F. M. Diaphanosoma brachyurum (LrEVIN) Leptodora kindtii (FocKE)

Copepoda

Cyclops strenuus FISCHER Eudiaptomus graciloides var. padance BuRCKH. M esocyclops leuckarti CLA us

There is no doubt that the composition of plankton in Lago di Como reflects a higher trophic level in comparison with that of Lago Nahuel Huapi. The plankton of Lago di Como seems to

be more related to the plankton of the preandean lakes in the Chilean lake district. These often have a Melosira-Eudorina-filamentous green-algae dominant community which is comparable to the Fragilaria-Eudorina-filamentous green-algae plankton in Lago di Como. It is interesting to note that in both plankton communities planktic filamentous green-algae occur, and in both cases are sterile . Noteworthy also is the absence of Stauro

desmus species in the present sample from Lago di Como. Naturally one ought to take into consideration the fact that important geographical peculiarities in the distribution of plankters occur, e.g. some taxa common in European lakes are lacking in the lakes of southern South America (THOMASSON,

1955) . Also the trophic status of Lago di Como is higher than that of Chilean lakes, e.g. Lake Villarrica, and Rifiihue. This is partly due to considerable auxotrophication especially near the town of Como. The high epilimnic temperature is probably of some importance in regard to the similarity in plankton composition between those lakes, e.g. in Lake Villarrica about 20°, in comparison to the low temperature of Lake Nahuel Huapi.

2. L A K E G U T I E R R E Z

Lake Gutierrez is situated about 1 5 km SW from Bariloche, just north of the continental divortium aquarum. The shores of the lake are bordered by Nothofagus antarctica. The northern end of the lake where the plankton sample was collected is rather shallow. The subaquatic macrophyte vegetation consists of M yriophyllum and a plant similar to Litorella.

The plankton was collected on January 22, 1 954, the weather was cloudy, cool and windy. The sample is dominated by Dinobryon divergens,

Botryococcous braunii, and Keratella cochlearis. Also frequent are Synchaeta pectinata, Filina limnetica,

and Bosmina . chilensis. Few benthic organisms were noted among the plankters.

Cyanophyta

Aphanocapsa elachista var. planctonica G. M. SMITH Ooelosphaerium kuetzingianum NA.o. Lyngbya sp. - fig. · 20: 1


Pyrroph:yta

Gymnodinium sp. - fig. 20: 8 Peridinium willei HUITF.-KAAS P. sp.

Chrysophyta


Dinobryon cylindricum lMH. D. divergens IMH. Stichogloea doederleinii (SCHMIDLE) WILLE

B A CILLARI O P H Y C E A E

Oyclotella stelligera CL. & GRUN. Oymatopleura solea (BREm.) W. SMITH Diatoma elongatum Aa.

Epithemia muelleri FRICKE - benthic Fragilaria sp. Melosira ambigua (GRUN.) 0. MULL. - rare M. distans (EHRENB.) KuTz. - a forma with a

diam. of 1 3.5 p, M. pseudogranulata A. CL. N eidium sp. - fig. 20: 6, length 234 p,, breadth 37 .5 p,

benthic

Lakes 45

Pinnularia spp. - benthic Rhizosolenia eriensis H. L. SMITH Stauroneis phoenicenteron EHRENB. - benthic Surirella guatimalensis EHRENB. S. robusta EHRENB. S. tenera GREG. Synedra acus KuTz. S. ulna (NITZSCH. ) EHRENB.

Ohlorophyta

V O L V O C A L E S

Asterococcus super bus ScHERFF. Elakatothrix gelatinosa fa. biplex NYGAARD Eudorina elegans EHRENB. Gloeococcus schroeteri (CHOD. ) LEMM.


Botryococcus braunii KuTz. Orucigenia quadrata MoRREN 0. rectangularis (N.AG. ) GAY Nephrocytium lunatum W. "-'TEST Oocystis lacustris CHOD. 0. solitaria WITTR. Pediastrum integrum fa. granulata RACIB . P. integrum var. scutum RACIB.

C o N J U G A L E S

Olosterium aciculare T. WEST 0. kuetzingii BREB. Oosmarium botrytis (BORY) MENEGH 0. margaritatum var. subtumidum - length 125 p, 0. quadrum LUND 0. subcucumis ScHMIDLE 0. subspeciosum var. validus N ORDST. Gonatozygon pilosum W OLLE Mougeotia sp. ster. - benthic Netrium digitus (EHRENB.) !TZIGS . & RoTHE -

benthic Spirogyra sp. ster. - benthic

Staurastrum dispar BREB. fa. WEST & WEST, 1 9 12, pl . 1 27' fig. 7

St. furcigerum BREB. St. planctonicum var. ornatum (GRONBLAD) TElLING St. sebaldi var. brasiliense B0RGES. St. sebaldi var. ornatum fa. planctonica (LuTKEM.)

TElLING Staurodesmus megacanthus (LuNn.) THUNM. Std. phimus (THURNER} THOM.

Protozoa

Arcella discoides EHRENB. "Cilia to negro" Oyphoder'ia margaritacea EHRENB . - benthic Euglypha brachiata LEIDY - benthic

Rotatoria

Oollotheca mutabilis (HUDSON) Oonochilus unicornis Rouss. Euchlanis dilatata EHRENB. Filina limnetica ZACH. H exarthra fennica (LEv.) Keratella c . cochlearis (GossE) - fig. 24: 10 K. c. tecta GossE Monommata longiseta 0. F. M. Monostyla (Lecane) lunaris (EHRENB.) Mytilina ventralis (EHRENB. ) var. - fig. 24: 5 Polyarthra vulgaris CARLIN Synchaeta oblonga EHRENB. S. pectinata (EHRENB.) Trichocerca insignis HERRICK T. tigris (0. F. M.)

Gladocera

Alona sp. - a species close to A . rectangula Bosmina chilensis DADAY

Copepoda

Boeckella schwabei BREHM M esocyclops annulatus (WIERZ.)

3. L A K E MA S C A R D I

The horseshoe-shaped Lake Mascardi lies on the southern side of the continental watershed, and it is remarkable for the singular beauty of the scenery surrounding it, cf. pi. 5. The plankton sample was collected on January 23, 1954. The weather was sunny, water temperature at noon about 12°0, in the morning 6-7°0. It may be of interest to mention that, according to the information from the manager

of the Hotel Mascardi, a few weeks before our visit, as low a water temperature as 1°0 has been noted.

The quantitatively as well as qualitatively sparse plankton was dominated by V agnicola sp. and Dinobryon divergens. Subdominants were Peridi

nium willei, Eudorina elegans, Rhizosolenia eriensis,

and Keratella cochlearis. Botryococcus braunii was frequent . .

A cta Phyfogeo.ar. Suec. 42

46 Lakes

Oyanophyta

Anabaena sp. Ohroococcus limneticus LEMM.

Pyrrophyta

Peridinium willei HurTF.-KAAS P. sp.

. Ohrysophyta

0 H R Y S O P H Y C E AE

Dinobryon cylindricum lMH. D. divergens lMH. M allomonas elongata REv. Salpingoeca buetschlii LEMM.

B A C I L L AR I O P H Y C E AE

Melosira patagonica (0. MuLL. ) FRENG. M. pseudogranulata A. CL. Pinnularia sp. R,hizosolenia eriensis H. L. SMITH Synedra ulna var. danica (KuTz. ) GRUN.

Ohlorophyta

V O L V O C A L E S

Eudorina elegans EHRENB. Gloeococcus schroeteri (CHOD.) LEMM.

0 H L O R O C O C C A L E S

Botryococcus braunii KuTz. Dictyosphaerium pulchellum WOOD Oocystis crassa WITTR.

C O N JU GALE S

Olosterium leibleinii KuTz. - benthic Euastrum pinnatum RALFS Gonatozygon monotaenium BARY Hyalotheca dissiliens (SMITH) BREB. Oosmarium depressum var. achondrum (BOLDT)

WEST & WEST 0. subspeciosum var. validus NORDST. 0. tetraophthalmum BR:EB. - length 125 p, Staurastrum excavatum var. minimum BERNARD

St. jurcigerum BREB. St. longipes var. evolutum (WEST & WEST) THOM. St. manjeldtii DELP. fa. - rare Staurodesmus triangularis fac. stroemii TElLING

Protozoa

Acanthocystis aculeata HERTW. & LESS. "Ciliato negro" Vagnicola sp.

Rotatoria

Oonochilus unicornis Rouss. Gastropus stylijer IMH. Keratella c. cochlearis ( GossE) Lecane glypta HARRING & M YERS Philodina sp. Polyarthra vulgaris CARLIN Trichocerca sp.

Copepoda

Boeckella sp.

Among the zooplankters there are two interesting members - Vagnicola sp. , fig. 24: 18, which was often met in great quantities in the plankton of the South American lakes investigated by us. It is an euplankter and will be treated by Dr. H. L6FFLER

in his work on the zooplankton collected by our expedition. The second, hitherto unidentified plankter, also belongs to the Protozoa. It is a large black ciliate, noted in my lists as "Cilia to negro" , not unlike Stentor niger EHRENB. It was often frequent in the lakes on both sides of the Andean range. The taxonomic position of this pseudonym will b e evident from the above mentioned study. The present notes as regards zooplankters are brief, since they will be subjected to a throughout treatment by my travelling companion Dr. H. L6FFLER.

4. L A K E G U I L L E L M O

Lake Guillelmo is a little picturesque valley-lake situated just a few kilometres south of Lake Mascardi; it borders the eastern slopes of Mt. Padre Laguna ( 1615 m a. s.-1 . ) . Lake Guillelmo discharges through a rapid flowing stream into Lake Mascardi. Here the construction of a waterpower plant was in progress. In the cle�r shallow

A cta Phytogeogr. Suec; 42

waters of the lake the occurrence of Potamogeton

linguatus HAGSTR. , M yriophyllum and rosettes rather like Litorella were noted. On the muddy bottom stoneworts (Nitella clavata KuTz. ) were abundant (distribution in Latin America see HoRN

AF RANTZIEN, 1950 ) .

The plankton sample collected on January 23,

Lakes 47

1954, is dominated by Dinobryon divergens and the above mentioned V agnicola sp. ; furthermore Aca-n,thocystis aculeata and "Ciliato negro" are frequent. The sample contains quantities of detritus and benthic algae owing to the shallowness of the lake and partly to the punting of the raft, cf. pl. 10.

Cyanophyta

Anabaena spp. - at least 2 species Aphanocapsa roeseana BARY Aphanothece microspora (MENEGH. ) RABENH. Chroococcus limneticus LEMM. Lyngbya sp. - diam. 10-1 1 .5 p, trichomes con-

stricted at the cross wall, cells short Merismopedia tenuissima LEMM.

Nostoc sp. - fig. 20: 2 Oscillatoria spp. - at least 2 species

Euglenophyta

Euglena sp.

Pyrrophyta

Peridinium cinctum (0. F. M.) EHRENB . ·

P. willei H UITF . -KAAS P. spp. - at least 2 more species

Chrysophyta

C H R Y S O P H Y C E A E

Dinobryon divergens !MH.

D. divergens var. schauinslandii (LEMM.) BRUNNTH. Salpingoeca buetschlii LEMM.

B A CILLARI O P H Y C E AE

Diatoma elongatum (LYNGB.) AG. Fragilaria sp. M elosira pseudogranulata A. CL. Rhizosolenia eriensis H. L. SMITH Sur�rella sp. Synedra ulna var. danica (KuTz.) GRUN.

Chlorophyta

V O L V O C A L E S

Gemellicystis neglecta TElLING Pandorina morum BoRY Paulschulzia pseudovolvox (SCHULZ & TElLING)

SKUJA Schizochlamys sp.

C H L O R O C O C C AL E S

Ankistrodesmus falcatus (CORDA) RALFS Botryococcus braunii

·KuTz.

Nephrocytium limneticum (G. M. SMITH) SKUJA Scenedesmus quadricauda (TURP. ) BREB.

U L O T R I C HALE S

Geminella mutabilis (BREB.) WILLE

0 E D O G O NIALE S

Oedogonium spp. - at least 2 species

C O N J U GALE S

Cosmarium quadrijarium LUND. C. subspeciosum var. validu_s N ORDST. C. sp. Gonatozygon aculeatum HAST. M ougeotia elegantula WITTR. - benthic M. gracilima (HAss.) WITTR. - benthic M. kerguelensis KRIEGER - diam. 1 2 p, benthic M. parvula HAss. - benthic M. sp. - zygospore globose, diam. 19 p,, vegetative

cells 12 .5 p in diameter Netrium digitus (EHRENB.) lTZIGS. & RoTHE -

benthic Staurastrum excavatum var. minimum BERN ARD St. furcigerum BREB.

St. longipes var. evolutum (WEsT & WEsT) THOM. St. pilosum (NA.G.) ARcH. St. sebaldi var. orientale (WEST & WEST) THOM. fa. St. sp . - fig. 23: 8 Staurodesmus dickiei RALFS Std. triangularis fac. stroemii TElLING

Protozoa

Acanthocystis aculeata HERTW. & LEss. Arcella discoides EHRENB.

"Cilia to negro" . Euglypha brachiata LEIDY Vagnicola sp.

.· Vorticella sp.

·Rotatoria

Cephalodella sp. Collotheca atrochoides WrERZ. 0. clava HUDSON Conochilus unicornis Rouss. Filina limnetica ZACH. Gastropus stylifer lMH. Keratella c. cochlearis (GossE) K. c. tecta GossE M onostyla (Lecane) hamata STOKES Philodina sp. Synchaeta sp. Trichocerca cylindrica (IMH.)

Trichotria tetractis EHRENB.

Copepoda

Boeckella schwabei BREHM Cyclops sp.


48 Lakes

5 . LA K E F R I A S

Lake Frias, appropriately also called Lake Frio, is a precious gem, hidden in the solitude of huge mountains and surrounded by thick woods, cf. pi. 1 1 . In the east it is framed by craggy mountains and in the west, behind the heights of Perez Rosales hill, one can see the white massive cupola of Mt. Tronador towering in the sky, looking round superciliously with the majesty of an emperor. Here · the ageold, almost impenetrable temperate rainforest creeps up the mountainside and extends along the lakeshore, cf. pl . 13. The colour of the lake is suggestive of the colour of Norwegian mountain lakes fed by glacier waters. The water of Lake Frias is coloured by glacier-ooze carried in by Rio Frias, which receives its water from the glaciers of Mt. Tronador. The capacity of Rio Frias for transporting gravel and sand is shown by the beautiful unilobate delta of this river, cf. pi. 12.

The macrophytes are represented in Lake Frias by sparse growth of Scirpus californicus at Puer.to Alegre and Hotel Lago Frias.

On February 1 , 1954, Diatoma elongatum was the dominant algae in the sparse plankton of Lake Frias; the subdominant species was Synedra ulna

var. danica. The most frequent zooplankter was Keratella cochlearis fa. tecta. There is no doubt that according to the composition of the plankton Lake Frias is ·the most oligotrophic among the lakes ·

in the National Park which have been studied by us.

Cyanophyta

Lyngbya putealis MoNT. - benthic

Euglenophyta

Euglena sp.

Chrysophyta

0 H R YS O P H Y C E AE

Dinobryon cylindricum lMH. M allomonas tonsurata TElLING

B A C I L L A R I O P H Y C E A E

Diatoma elongatum (LYNGB. ) AG. D. hiemale var. mesodon (EHRENB. ) GRUN.


D. vulgare var. producta GRUN. Fragilaria sp. Gomphonema acuminatum var. coronata (EHRENB. )

RALFS - benthic Melosira italica (EHRENB.) KuTz. Neidium hitchcockii (EHRENB.) CLEVE - benthic Nitzschia vermicularis (KuTz.) GRUN. Surirella guatimalensis EHRENB. S. robusta H UST. S. spp. Synedra ulna var. danica (KuTz. ) GRUN. Tetracyclus ellipticus var. lancea fa. chilensis

KRASSKE

Chlorophyta

0 H L O R O C O C CA L E S

Ankistrodesmus falcatus (CORDA) RALFS A. falcatus var. spiralis WEST

C O N J U G A L E S

Olosterium kuetzingii BR1Jm. Oosmarium botrytis (BORY) MENEGH. 0. speciosum var. validus NORDST. 0. subtumidum var. pachydermum PRESCOTT &

SCOTT Staurastrum laeve RALFS S. pseudonanum GRONBLAD fa. Xanthidium antilopaeum (BREB. ) KuTz.

Protozoa

Vagnicola sp.

Rotatoria

Oolurella tesselata (GLASCOTT) G. uncinata (0. F. M.) G. sp. Oonochilus un,icornis Ro uss. Euchlanis incisa 0ARLIN Keratella c. cochlearis (GossE) K. c. tecta GossE Lecane flexilis (GossE) L. luna (0. F. M.) Monostyla (Lecane) lunaris EHRENB. Polyarthra vulgaris CARLIN Trichocerca longiseta SCHRANK T. tigris (0. F. M.)

Lakes 49

Fig. 1 9. Lake Hess. Dense reeds of Scirpus calijornicus; in the foreground Eucryphia cordijolia (right) and Spartium junceum (left) .

K. THO�IASSON

6. L A K E H E S S

This small and shallow lake is . situated about 10

kilometres west of Lake Mascardi, which discharges

its waters through Rio Manso into Lake Hess.

Furthermore it receives considerable inflow from

Lake Fonck. Rio Manso forms also the outflow

from Lake Hess after passing through this lake.

Lake Hess holds an exceptional position among the

lakes visited by us in Nahuel Huapi National Park

in respect to the aquatic vascular plants. It has a

very rich vegetation, e.g. in the shallow south

western end of the lake a dense vegetation of

Scirpus californicus (C. A. MEY) BRITTON grows.

Through these reeds the arms of Rio Manso pene

trate in the form of channels. The water depth in

this part of the lake varies around 3-5 m. Areas not

overgrown with reeds are characterized by dense

growth of pondweeds (Potamogeton linguatus HAG

STR.) and the bottom is covered with M yriophyllum.

According to information from residents, the lake

is on the whole shallow, its maximum depth being

about 20-25 m. Of course in the deeper parts of the


lake the macrophytic vegetation is absent, e.g.

on the muddy bottom at 17 m depth, only a rich

occurrence of mussels was noted. The mussels were

abundant also in the shallow areas of this lake.

The Ca content of the water of Lake Hess was

4.9 p. p. m. , Na 1 .3 p. p. m. and K 0.4 p. p. m.

For comparison some low values from the lakes

situated west of the Andean range may be quoted

Lake Llanquihue ( 10.2 . 1954) Ca 4.9 p. p. m. , Na

9 . 1 p. p. m. , K 1 .2 p. p. m . , and Lake Villarrica

( 19 .2 . 1954) Ca 5.0 p. p. m. , Na 4.4 p. p. m. , K 1 .3

p. p. m. The relatively high ·value for Na in Lake

Llanquihue may be due to the influence of the sea;

the lake is separated from the Bay of Reloncavi by

a strip of land few kilometres broad. The deepest

part of Lake Llanquihue is below sea level, but there

is no evidence of a .sea water layer at this depth.

The plankton sample was collected on January

26, 1954. Unlike the sparse plankton of Lake Frias

and other lakes the plankton of Lake Hess is

relatively rich. Lake Hess is the most eutrophic


50 Lakes

among the lakes studied by us in the National Park. Remarkable is the richness of desmids, but one must not draw precipitate conclusions from this, since a great portion of these desmids is of tychoplanktic and benthic origins. The abundant occurrence of these desmids in the plankton is due to the unique hydrographic conditions in this lake. As has been mentioned before, Rio Manso flows through the southern part of the lake, and here we also find delta formations. Furthermore, there is a considerable inflow from Lake Fonck. These currents produce mixing of the rather small and shallow body of water and this brings numerous benthic organisms and organisms living on and among submerged macrophytes into the plankton. Also the presence of great quantities of detritus in the seston is due to the same factors.

The plankton of Lake Hess is dominated by the same Vagnicola sp. which is frequent in many of the lakes in Nahuel Huapi National Park. The most interesting group of algae in the present sample is desmids. The composition of plankton is evident from the following list.

Cyanophyta

Anabaena scheremetievi var. recta fa. ovalispora ELENKIN

A . spp - at least 2 more species Aphanocapsa pulchra (KuTz. ) RABENH. Gomphospaeria lacustris CHOD. Merismopedia glauca (EHRENB.) N.AG. Nostoc sp. Oscillatoria lacustris (KLEB . ) GEITL. - cf. KUKK,

1 958. 0 . tenuis AG. 0. sp.

Py'rrophyta

Glenodinium sp. Peridinium cinctum ( 0. F. M.) P. willei HUITF.-KAAS

Chrysophyta

H E TE R O K O NTAE

Botrydium granulatum (L.) GREY. - washed in by the inflows


Ohrysosphaerella sp. Dinobryon divergens IMH. D. sertularia EHRENB.


M allomonas acaroides PERTY M. caudata IwANOFF M. elliptica (KISSELEW) CoNRAD Salpingoeca buetschlii LEMM.

B AC I L L A R I O P H Y C E A E

Diatoma elongatum (LYNGB. ) AG. Fragilaria sp. Melosira ambigua (GRUN.) 0. MuLL .

M. pseudogranulatum A. CL. Pinnularia spp.

, Rhizosolenia eriensis H. L. SMITH Surirella guatimalensis EHRENB . S. sp .. Synedra ulna (NITZSCH. ) EHRENB .

Chlorophyta


Ankistrodesmus jalcatus (CoRDA) RALFS Botryococcus braunii KuTz. Dictyosphaerium pulchellum W oon Nephrocytium lunatum W. WEST Oocystis elliptica W. WEST 0. solitaria WITTR. Pediastrum araneosum var. rugulosum (G. S . WEST)

G. M. SMITH P. boryanum (TURP.) MENEGH. P. boryanum var. longicorne RACIB. P. duplex ME YEN

C o NJ U GA L E S

Olosterium acerosum (ScHRANCK) EHRENB. 0. aciculare T. WEST 0. kuetzingii BR11m. 0. leibleinii KuTz. Oosmarium abbreviatum var. planctonicum WEST &

WEST 0. botrytis (BORY) MENEGH. 0. brebissonii MENEGH. - length 1 12.5-135 p, 0. contractum var. ellipsoideum (ELFV.} G. S. "-'EST 0. depressum var. achondrum (BOLDT) WEST & WEST 0. monilijorme (TURP.) RALFS 0. pseudoconnatum var. ellipsoideum WEST & WEST 0. quadrijarium fa. THOMASSON, 1 955, fig. 1 : 2, cf.

also BORGE, 1 906, pl . 2, fig. 6. 0. subdanicum WEST 0. subspeciosum var. validus NoRDST. Oylindrocystis brebissonii MENEGH. Desmidium cylindricum GREV. Gonatozygon aculeatum HASTINGS G. monotaenium BARY Gymnozyga monilijormis EHRENB. Hyalotheca dissiliens (SMITH) BR:Em. Micrasterias denticulata BR:Em. - fig. 2 1 : 20. M. radians TURN. M. sol var. elegantior G. S. WEST - fig. 2 1 : 13 .

Tarns, ponds, and temporary waters 51

M . truncata (CORDA) BREB. Pleurotaenium coronatum var. nodulosum (BREB .)

w. WEST P. coronatum var. nodulosum fa. constricta - KRIE-

GER, 1937, pl. 49, fig. 1, length 682 P, P. ehrenbergii (BREB . ) BARY Sphaerozosma aubertianum WEST Staurastrum avicula var. subarcuatum (WOLLE)

WEST - !RENEE-MARIE, 1 939, pl. 50, fig. 5 & 8 St. dilatatum EHRENB . St. excavatum var. minimum BERNARD St. furcigerum BREB. St. laeve RALFS - a form with long processes St. longipes var. evolutum (WEST & WEST) THOM. St. margaritaceum (EHRENB.) MENEGH. St. pseudosebaldi WILLE fa. - fig. 22: 5 St. punctulatum var. ellipticum LEWIN St. rotula var. smithii (G. M. SMiTH) THOM. St. tetracerum var. biverruciferum GRONBLAD St. tohopekaligense var. brevispinum G. M. SMITH St. sp. - fig. 2 1 : 3 St. sp. - fig. 23: 17 Staurodesmus megacanthus (LUND.) THUNM. fa. Std. mucronatus var. subtriangulare (WEST & WEST)

THOM. Std. triangularis fac. stroemii TElLING Xanthidium antilopaeum (BREB .) KuTz.

Protozoa

Acanthocystis aculeata HERTW. & LESS. Arcella discoides EHRENB. A. hemisphaerica PERTY Euglypha brachiata LEIDY Vagnicola sp.

Rotatoria

Collotheca sp. Oolurella adriatica EHRENB. - fig. 24: 13 C. oxycauda CARLIN Conochilus unicornis Rouss. Euchlanis cf. meneta MYERS Keratella c. cochlearis (GossE) Lecane flexilis (GossE) L. luna (0. F. M. ) Lepadella patella (0. F . M.) Monostyla (Lecane) constricta MURRAY M. lunaris EHRENB. M. sp. Polyarthra dolichoptera (!nELSON) P. vulgaris CARLIN Ploeosoma sp.

Oladocera

Alona sp. Bosmina chilensis DADAY

V I I I . T A R N S , P O N D S, A N D T E M P O RA RY WAT E R S

During our visit to Nahuel Huapi National Park a few collections were made from minor bodies of lentic water. These waters are not expected to be representative, but they were situated along our

excursion routes. There are many interesting lakelets in the western part of the National Park, but because of lack of time, unfortunately there was no opportunity to visit them.

1. S T R A N D P O O L

In a temporary pool on the beach of Lake N ahuel Huapi near the military camp, the algal vegetation consisted of sterile threads of Zygnema intermingled with the threads of M ougeotia and Spirogyra, the latter present as at least two species. Among these plants was the multiplicity of algae listed below, the most frequent species being Synedra ulna.

Minute benthic diatoms were abundant, they will be included in a special study of the diatom flora. The following algae were noted:

4* - 588829

Oyanophyta

Dactylococcopsis rhaphidioides HG. Merismopedia glauca (EHRENB. ) N.AG. Oscillatoria cf. borneti ZUKAL. - diam. 9 p, 0. sancta (KuTz. ) GoM. 0. sp.

Ohrysophyta

B A C I L L A R I O P H Y T A

Oeratoneis arcus K UTZ. Diatoma elongatum AG. Synedra ulna (N ITZSCH.) EHRENB .


52 Tarns, ponds, and temporary waters

Chlorophyta


Ankistrodesmus cf. convolutus CoRDA A. falcatus (CoRDA) RALFS A. falcatus var. spiralis (TURNER) G. S. WEST Quadrigula closteroides (BOHLIN) BaRGE Scenedesmus bijugatus (TURP.) KuTz. S. ecornis (RALFS) CHOD . S. obliquus (TURP. ) KuTz. S. ovalternus var. graevenitzii (BERNARD ) CHOD . -

cf. p. 66

C O N J U GALE S

Closterium venus KuTz. Cosmarium botrytis (BORY) MENEGH. C. inconspicuum WEST & WEST C. laeve RABENH. C. lundellii var. ellipticum WEST C. ochthodes var. amoebum WEST - length 75 p C. pachydermum LuND .

C. vexatum WEST fa. WEST & WEsT, 1 908, pi. 92, fig. 5

Euastrum turneri W. WEST Staurastrum brebissonii ARcH. - length 60 p Staurastrum punctulatum BREB. St. suborbiculare WEST & WEST fa.

Phycomycetes

Tetracladium setigerum (GROVE) !NGOLD

Rotatoria

Cephalodella sp. Colurella hindenburgi STEINECKE - fig. 24: 1 C. tesselata (GLASCOTT) - fig. 24: 2 Keratella c. cochlearis (GossE) Lecane flexilis (GossE) Lepadella amphitropis BARRING - fig. 24: 12 L. patella (0 . F. M.) M onostyla lunaris EHRENB.

Cladocera

Chydorus sphaericus (0. F. M.)

2 . P O N D N E A R M I L I T A R Y C A M P

This small and shallow pond is situated just by the road to Llao-Llao, outside of the camp area, which was our head-quarter during our stay in the National Park.

The plankton sample was collected on January 25, 1954. The whole plankton is to be considered as heleoplankton; it is a mixture of plankters, littoral forms, and bottom organisms . Desmids are frequent, especially Cosmarium botrytis, which is dominant together with Lecane flexilis. Subdominants are H yalotheca dissiliens, Pleurotaenium

ehrenbergii, Cosmarium connatum, Ankistrodesmus

falcatus, and Lepadella patella. Minute diatoms of benthic origin were also abundant.

Euglenophyta

Euglena sp. Phacus orbicularis HuBNER

Pyrrophyta

Peridinium cinctum (0. F. M.) EHRENB . P. sp .

Chrysophyta


Dinobryon cylindricum lMH.


D. divergens lMH. Salpingoeca buetschlii LEMM.

B A C ILLARIO P H Y C E AE

Diatoma elongatum AG. Gomphonema acuminatum var. coronata (EHRENB. )

w . SMITH G. constrictum EHRENB . G. sp . Melosira sp.

Neidium hitchcockii (EHRENB.) CLEVE Pinnularia sp. Synedra acus KuTz.

Chlorophyta

V o L v o c A L E S

Eudorina elegans EHRENB.

C H L O R O C O C A L E S

Ankistrodesmus falcatus (CORDA) RA:CFS Pediastrum araneosum RACIB. p. duplex ME YEN Scenedesmus abundans (KIRCHNER) CHOD . S. bijuga var. alternans (REINSCH) BaRGE S. bijuga var. alternans (REINSCH) BaRGE fa. S. quadricauda EHRENB.

C O NJ U GA L E S

Closterium abruptum W . "''"EST C. acerosum (SCHRANK) EHRENB.


0. kuetzingii BREm. 0. venus KuTz. Oosmarium botrytis (BORY) MENEGH. 0. connatum BREB . 0. cucumis (CORDA) RALFS 0. diplosporum (LUND. ) LUTK. 0. granatum BREB. 0. perincisum GRONBLAD fa. 0. punctulatum var. subpunctulatum (NoRDST. )

B0RG. 0. quadrijarium fa . octasticha NoRDST. 0. subcrenatum HANTZSCH - length 41 p, 0. subspeciosum var. validus N ORDST. Euastrum insulare var. silesiacum GRONBLAD E. turneri W. WEST - length 28 p, Hyalotheca dissiliens (SMITH) BREB. Micrasterias truncata var. pusilla G. S. WEST -

fig. 22: 4 Penium margaritaceum (EHRENB.) BREB. - length

90- 1 84 ft P. margaritaceum var. elongatum KLEBS - length

225 ft P. sp. Pleurotaenium ehrenbergii (BREB. ) BARY P. ehrenbergii var. undulatum SCHAARSCHM. Sphaerozosma granulatum RoY & BISS. Staurastrum alternans BREB. St. erasum BREB. fa. St. excavatum var. planctonicum KRIEGER St. furcigerum BREB . St. orbiculare RALFS

St. perundulatum GRONBLAD St. sebaldi var. brasiliensis B0RGES. St. sebaldi var. orientale (WEST & WEST) THOM. fa. St. sebaldi var. ornatum NoRDST. fa. St. spongiosum BREB. - rare St. subpolymorphum BoRGE fa.

St. tetracerum RALFS-TAFT, 1 945, pi. 5, fig. 17 St. tohopekaligense W OLLE Staurodesmus convergens (EHRENB .) LILLIER. Std. dejectus (BREB.) TElLING - G. M. SMITH, 1 924,

pl. 68, fig. 1 8-24 Std. triangularis var. convergens THOM.

Protozoa

0 entropyxis sp. Euglypha brachiata LEIDY

Rotatoria

Keratella c. cochlearis (GossE) Lecane brundinii THOM. L. flexilis ( GossE) Lepadella patella ( 0. F. M.) L. triptera EHRENB. M onostyla hamata STOKES Polyarthra vulgaris CARLIN

Cladocera

Ohydorus sphaericus (0. F. M.) lliocryptus agilis KURz Simocephalus sp.

3. I N U N D A T I O N P O O L

Along the route "258" on the road between Lake Gutierrez and Lake Mascardi, and situated about l 0 m from the road, a large but shallow body of water was visited. The maximum depth of water was about l m and the whole bottom was covered with grasses. My impression was that it was a temporary body of water, a piece of submerged valley bottom. The sample collected on January 23, 1954, contained a rich population of Cladocera and beautiful red-colored Boeckellids, which could be observed on the spot. The main part of the sample was composed of filamentous algae. Amongst the threads there were abundant minute diatoms and a varied flora of desmids. The most frequent desmid was Cosmarium subspeciosum

var. validus. Geminella mutabilis may be designated as frequent in the sample and the following organisms were noted:

Cyanophyta

Anabaena lapponica BoRGE - Diam. veget. cells 6 .3 ft, hetercysts 1 0 x 10 p,, spores 61 x 15 p,, thick wall.

A . sp.

Aphanothece castagnei (BREB.) RABENH. Ohroococcus dispersus (KEISSL. ) LEMM. 0. turgidus (Kfuz. ) N.AG. Gomphosphaeria lacustris CHOD. Nostoc coeruleum LYNGB. Oscillatoria agardhii GoM. 0. sp.

Euglenophyta

Phacus anacoelus STOKES P. caudatus HuBN. Trachelomonas hispida (PERTY) STEIN em. DEFL. T. rugulosa var. dangeardii DEFL.

Pyrrophyta

Gymnodinium sp.

A cta Phytogeogr. Szzec. 42

54 Tarns, ponds, and temporary waters

Peridinium umbonatum STEIN P. umbonatum var. inaequale (LEMM.) LEF. P. willei HurTF.-KAAs

Ghrysophyta

H E TE R O K O NT A E

Ophiocytium cochleare A. BRAUN 0. parvulum A. BRAUN Tribonema sp.


Bicoeca lacustris .J. CL. Ohrysopyxis paludosa FoTT Dinobryon sertularia EHRENB. M allomonas sp.

B A C IL L A R I O P H Y C E AE

Diatoma elongatum AG.

Ghlorophyta

V o L v o cALE S

Eudorina elegans EHRENB. Pandorina morum BoRY Paulschulzia pseudovolvox (TElLING) SKUJA Schizochlamys sp. Tetraspora lacustris LEMM.


Ankistrodesmus falcatus (CoRDA) RALFS Botryococcus braunii KuTz. Dictyosphaerium ehrenbergianum NA.G. D. pulchellum WooD Nephrocytium limneticum (G. M. SM!TH) SKUJA Pediastrum tetras var. tetraodon (CORDA) HANSG. Quadrigula closterioides (BOHLIN) PRINTZ Scenedesmus bijugatus (TURP.) KuTz. S. ecornis (RALFS) CHOD, - G. M. SMITH, 1920, pl.

37, fig. 1 9-20 S. sp.

UL O T R I C HALE S

Geminella mutabilis (BREB.) WILLE Radiojilum flavescens G. S . ��EST

0 E D O G O N IALE S

Bulbochaete sp.

C O N J U G ALE S

Closterium dianae KuTz. C. kuetzingii BRll:B. Cosmarium binum NoRDST. C. boeckii WILLE C. botrytis (BORY) MENEGH. 0. depressum var. planctonicum REv. 0. quadrijarium LUND . fa. THOMASSON, 1 955, fig. 1 : 2 0. quadrifarium fa. octasticha No:a.nsT.


0. regnellii WILLE - length 13 p,, breath 1 0.5 p, 0. regnesi REINS CH

.

0. subdanicum WEST 0. subnotabile WILLE 0. subspeciosum var. validus N ORDST. 0. viride (CORDA) .JOSHUA - ]ength 47.5 p, Oylindrocystis sp. Euastrum bidentatum var. speciosum (BoLDT)

SCHMIDLE fa. - fig. 22: 9 E. dubium NA.G. E. turneri W. WEST Gonatozygon aculeatum HAST. G. monotaenium BARY Hyalotheca dissiliens (SMITH) BREB. M ougeotia parvula HASSALL M. spp. - almost 2 more species Penium margaritaceum (EHRENB.) BREB. Spirogyra sp. Staurastrum alternans BREB. St. brebissonii ARcH. St. furcigerum BREB. St. margaritaceum (EHRENB.) MENEGH. fa. St. pilosum (NA.G. ) ARCH. St. polymorphum BREB. St. punctulatum var. kjellmani WILLE St. sexcostatum var. productum WEST __; length

37 .5 "'

St. spongiosum BREB . - length 4 1 .2 p, St. suborbiculare WEST & WEST fa. Staurodesmus dejectus var. patens NoRDST. - WEST

& CARTER, 1 923, pl . 1 30, fig. 1, 2, a little form, length only 20 p,

Std. glabrus fac. glabrus (EHRENB.) TEILiNG Std. megacanthus var. scotticum (WEsT & WEST)

FLORIN Std. triangularis fa. curvispina THOM. - THOMAS

SON, 1957, fig. 2: 8 Zygnema fanicum LI

Protozoa

Arcella discoides EHRENB. A. vulgaris EHRENB. Difflugia acuminata EHRENB. Euglypha alveolata DuJ. E. brachiata LEIDY E. compressa CARTER Heliozoa Trinema enchelys (EHRENB . )

Rotatoria

Oephalodella cf. forficata (EHRENB.) 0. cf. sterea fa. mutata DoNNER Oolurella obtusa ( GossE) Dicranophorus grandis (EHRENB.) Euchlanis dilatata EHRENB.


E. cf. meneta M YERS Keratella c . cochlearis (GossE ) Lecane luna (0. F. M.) - fig. 24: 1 6 Lepadella patella ( 0 . F. M. ) L. rhomboides (GossE) Monommata longiseta 0. F. M. Monostyla (Lecane) hamata STOKES N otholca caudata CARLIN Scaridium longicaudum (0. F. M.) - fig. 24: 8 Trichocerca tigris 0. F. M. T. sulcata (JENN. ) T . weberi J ENN. Trichotria tetractis (EHRENB .)

Oladocera

Alona rectangula SARS - DADAY, 1 905, close to A. intermedia SARS

A. sp. Ceriodaphnia spp. - almost 2 species Chydorus sphaericus (0. F. M.) Bosmina chilensis DADA Y Diaphanosoma sp. Scapholeberis spinifera NICOLET

Copepoda

Boeckella schwabei BREHM Microcyclops anceps (RICHARD )

4. T A R N I N P A S O V U R I L O C H E I

On January 30, 1954, an attempt was made to visit some bog tarns situated south of Mt. Tronador in Paso Vuriloche. According to information from the late Prof. E. LJUNGNER these tarns are characterized by brown water and are surrounded by Sphagnum-bog. They are situated immediately below the timber-line. Due to the time-consuming ascent of this pass the plan of study was never carried out. One of these tarns I did reach, but our pack-horse with equipment was far behind and there was no time to wait. I was forced to turn on my heel and be satisfied with a fistful of Spirogyra sp. ster. Amongst these filaments were intermingled some Mougeotia sp. ster. and many minute diatoms. The following plants were recorded:

Oyanophyta

Cylindrospermum majus KuTz. Gloeotrichia pisum THURET N ostoc kihlmani LEMM. Oscillatoria cf. hamellii FREMY - cells 6 p, broad,

8-1 0 p, long Tolypothrix sp.

Euglenophyta

Trachelomonas hispida var. punctata LEMM.

Ohrysophyta

H E T E R O K O NTAE

Ophiocytium parvulum (PERTY) A . BRAUN

0 H R Y S O P H Y C E AE

Dinobryon cylindricum IMH. Salpingoeca sp.

B A C IL L A R I O P H Y C E AE

Melosira distans (EHRENB .) KuTz .

Ohlorophyta

V O L V O C A L E S

Gloeocystis gigas (KuTz. ) LAGERH.

C H L O R O C O C C ALE S

Dictyosphaerium pulchellum W oon Pediastrum tetras (EHRENB.) RALFS Scenedesmus ecornis (RALFS) CHOD . S. brasiliensis BoHLIN Selenastrum bibraianum REINSCH

C O N J U GA L E S

Closterium dianae EHRENB . C. kuetzingii BREB. C. sp. Cosmarium amoenum var. intumescens NoRDST. C. controversum fa. major THOM. C. punctulatum BREB. fa. C. pyramidatum -BREB. C. subspeciosum var. validus NoRDST. C. vexatum WEST. fa. - length 44 p,, breadth 44 p, Euastrum dubium var. latum KRIEGER E. insulare var. silesiacum GRONBLAD Netrium digitus (EHRENB.) ITZIGS. & RoTHE Staurastrum alternans BREB . fa. WEST & WEST, . 1 9 1 2, pl. 1 26, fig. 9 & 1 0 St. dilatatum EHRENB.

Protozoa

Opercularia sp. Vagnicola sp.

Rotatoria

Lecane flexilis GossE L. glypta HARRING & MYERS Trichocerca tigris ( 0 . F. M.)


56 Running waters

5. A L P I N E T A R N S

While studying South American lakes, our special interest was directed to alpine waters with the intention of making a comparison with European alpine waters. There are no suitable waters of alpine character within the National Park of Nahuel Huapi. Only three tarns north of Mt. Tronador, near the field hut at Rigi, can be considered alpine waters, cf . pi. 16 . They are all small and only knee-deep, and practically lacking in planktic life. The altitude is about 1600 m a. s .-1. The few chemical factors analysed for these waters are as follows : Tun � Na K

I 0 . 5 p.p.m. 0.8 p.p.m. 0.4 p.p.m. II 0.5 0. 2 .

0.2

Ill 0 .9 15.2 1 . 3

The bottoms are of darkish sand of volcanic origin. The samples from these alpine tarns are of

quite uniform content, mainly organisms of benthic origin. The following were noted: A phanothece

castagnei (BR:Ifm.) RABENH. , Chroococcus tenax

(KIRCH) HIERON., C. turgidus (Kti"Tz.) N.A.G. , Eudo

rina elegans EHRENB . , Surirella sp. , Closterium

venus Kti"TZ. , Cosmarium quadrifarium LuND fa. THOMASSON, 1955, fig. 1 : 2, length of plants 52.5 fl, frequent, Euastrum lapponicum ScHMIDLE fa. The genus M esotaenium is richly represented in our samples, but these plants should be studied in a living state. The following taxa have been identified with some doubt: M esotaenium de greyi var. borgei

(BORGE) KRIEGER and M. chlamydosporum BARY.

The genus Penium is represented by Penium spiro

striolatum BARKER and P. margaritaceum (EHRENB . )

BREB. Moreover a few specimens of M onostyla (Le

cane) lunaris (EHRENB. ) were recorded.

I X . R U N N I N G WA T E R S

I . S M A L L B R O O K A T L A K E G U T I E R R E Z

This brook runs down from mountains more than 2000 m high on the eastern side of Lake Gutierrez. It crosses the route "258" and enters Lake Gutierrez. During a stoppage caused by engine failure the fresh green colour of the stones in the brook caught our eyes. It was found that the stones were covered with a dense vegetation of Prasiola mexicana LrEBM.

apud J. AG. cf. JESSEN, 1 848, pi. 1, fig. 17-20. The alga seems to thrive in this clear and cold water. The characteristic clavate basal-cells in this species which are a very conspicuous feature was first pointed out by LAGERSTEDT, 1869. The frequent

occurrence of Prasiola mexicana in Andean brooks in Venezuela has been recorded by GESSNER, 1955,

p. 338. In GESSNER, 1956, p . 19, the favourite habitat for Prasiola mexicana in the Venezuelan Andes is given as small cataracts; that is the same as our locality in the Nahuel Huapi National Park. Among the "leaves" of Prasiola the occurrence of band-shaped colonies of Diatoma

hiemale var. mesodon (EHRENB. ) GRUN, and a Fra

gilaria sp. was noted. Furthermore, there were a few specimens of Arcella discoides EHRENB. present in my samples.

2 . B R O O K A T L A K E M A S C A R D I

On January 26, i954, a sample containing epilithic tufts of a sterile Spirogyra was collected from a brook at Lake Mascardi. Among the filamentous green algae the following organisms were noted:


Cyanophyta

Merismopedia glauca (EHRENB. ) NA.G. N ostoc sphaericum V A ucH.

Oscillatoria sp.

Running waters 57

Euglenophyta

Euglena cf. clavata SKUJA E. sp. Phacus pleuronectes (0. F. M.) DuJ.

Ohrysophyta


Diatoma hiemale var. mesodon (EHRENB. ) GRuN. Gomphonema acuminatum var. coronata (EHRENB. )

w . SMITH H antzschia cf. subandina FRENG. H. sp. N eidium hitchcockii (EHRENB . ) CLEVE Nitzschia acicularis W. SMITH Melosira varians AG. Stauroneis sp. Synedra acus KuTz. S. ulna (NITZSCH) EHRENB .

S. ulna (NITZSCH) EHRENB. var. S. ulna var. amphirhynchus (EHRENB.) GRUN.

Ohlorophyta

C O N J U GALE S

Olosterium venus KuTz. Cosmarium botrytis (BORY) MENEGH. 0. subspeciosum var. validus N ORDST. Staurastrum punctulatum BREB . - length 41 p. St. trapezicum BoLDT

Phycomycetes

Tetracladium setigerum (GROVE) INGOLD

Protozoa

Euglypha brachiata LEIDY

Rotatoria

Lepadella patella ( 0. F. M. )

3. R I VE R B E T W E E N L. G U T I E R R E Z AN D L. NA H U E L H UA P I

Through this river Lake Gutierrez discharges its waters into Lake Nahuel Huapi. In the slower water of lentic character Potamogeton grows. The benthic vegetation is composed of Fissidens rigidulus, and on stones a dense mat of Spirogyra occurs. Intermingled with the filaments of Spirogyra grow M ougeotia and Bulbochaete. Besides the huge number of minute benthic diatoms the following organisms have been noted amongst the tufts of Spiro

gyra:

Oyanophyta

M erismopedia glauca (EHRENB . ) N A. G. Nostoc kihlmani LEMM. Oscillatoria sp.

Ohrysophyta


Dinobryon cylindricum IMH.

B A C I L L A R I O P H Y C E A E

Cyclotella stelligera CL. & GRUN. a. stelligera var. elliptica FRENG. Cymatopleura solea (BREB. ) W. SMITH Diatoma elongatum AG. Epithemia muelleri FRICKE Eunotia monodon var. maior (W. SMITH) HusT.

length 1 0 2 . 5 p. Fragilaria pinnata EHRENB.

Gomphonema acuminatum var. coronata (EHRENB . ) w. SMITH

M elosira pseudogranulata A. CL. M. distans (EHRENB. ) KuTz. M. italica (EHRENB.) KuTz. Pinnularia legumen EHRENB. P. spp. Surirella guatimalensis EHRENB. S. cf. engleri fa. sublaevis 0. MuLL. Stauroneis sp. Synedra acus KuTz. S. ulna (N ITZSCH) EHRENB.

Ohlorophyta


Pediastrum boryanum (TURP.) MENEGH. P. tetras (EHRENB.) RALFS Scenedesmus quadricauda (TURP.) BREB.

C O N J U G ALE S

alosterium venus K UTZ. aosmarium arnellii fa. compressa WEST - length

44 p., breath 40 p., isthmus 1 2 . 5 p. 0. botrytis (BORY) MENEGH. a. subspeciosum var. validus NoRDST. Euastrum turneri var. bohemicum L UTKEM. Gonatozygon kinahani (ARcH.) RABENH. Penium margaritaceum (EHRENB.) BREB. Staurastrum brebissonii ARcH. St. punctulatum BREB.


58 Benthos

Phycomycetes

Tetracladium setigerum (GROVE) !NGOLD

Protozoa

Vorticella sp.

Rotatoria

aolurella adriatica EHRENB . Keratella c. cochlearis (GossE ) Lepadella patella ( 0. F . . M.) Monostyla (Lecane) lunaris EHRENB.

X. B E N T H O S

In spite of the fact that the benthos is not considered in any detail in the present paper, the following list may be of some interest. The sample was collected from the littoral of Lake Guillelmo on January 23, 1954. It consisted mainly of detritus and small diatoms. The following organisms were recorded:

Oyanophyta

Anabaena spp. Aphanothece stagnina (SPRENG.) A. BRAUN ahroococcus turgidus (KuTz.) NA.G. a. turgidus var. maximus NYGAARD Gomphosphaeria aponina KuTz. G. aponina var. cordiformis WILLE G. lacustris CHOD. Merismopedia glauca (EHRENB. ) NA.G: N ostoc kihlmani LEMM. Oscillatoria limosa AG. 0. splendida GREV. 0. spp.

Pyrrophyta

Gymnodinium rotundatum KLEBS Peridinium cinctum (0. F. M.) EHRENB. P. inconspicuum tab. contactum fa. armatum (LEMM.)

LEF. P. willei H UITF. -KAAs

Ohrysophyta


Dinobryon divergens !MH.


ayclotella stelligera CL. & GRUN. a. stelligera var. elliptica FRENG. Diatoma elongatum AG. Fragilaria pinnata var. elliptica (ScHUMANN)

CARLSS. M elosira distans var. alpigena GRUN.


M. pseudogranulata A. CL. Pinnularia macrocephala A. CL. P. polyonca (BREB. ) 0. MULL. P. spp. Rhizosolenia eriensis H. L. SMITH Surirella guatimalensis EHRENB. S. spp. Synedra acus KuTz.

Ohlorophyta


Botryococcus braunii KuTz. Pediastrum boryanum (TURP.) MENEGH. P. integrum var. scutum RACIB. p. tetras (EHRENB.) RALFS

U L O T R I C HALE S

Geminella mutabilis (BREB.) WILLE

CoN JUG ALES

Euastrum attenuatum var. lithuanicum WoLosz. E. bidentatum NA.G. aosmarium controversum WEST - length 1 1 2 . 5 p,,

breadth 8 6 p,, isthmus 2 7 . 5 p, a. granatum BREB. a. laeve RA:BENH. fa.

a. pseudoexiguum RACIB. - length 27.5 p, a. subspeciosum var. validus NORDST. aylindrospermum sp. Staurastrum excavatum var. minimum BERNARD St. furcigerum BREB. St. longipes var. evolutum (WEST & WEsT) THOM. Staurodesmus triangularis fasc. stroemii TElLING -

length 1 9 p,

Protozoa

Euglypha brachiata LEIDY V agnicola sp.

Rotatoria

aollotheca sp.

Retrospective survey 59

Euchlanis dilatata EHRENB. Keratella c. cochlearis (GossE ) Monostyla (Lecane) cf. acanthinula HAUER Polyarthra dolichoptera (IDELSON) P. sp. 'Prichocerca insignis (HERRICK) T. weberi J ENN.

Copepoda

Cyclops sp.

A number of other samples contains benthos. These are to be the object of investigation in connection with the studies on the diatom flora, since the diatoms are the most important part of the benthic population. In this connection it should be empha-

.

sized that the only hitherto published report on the algal flora of the Nahuel Huapi National Park deals with the benthos. It is a paper by CLEVE -EULER

( 1943) on a bottom sample from Lake Frey, based on a sample collected by the late Prof. E . LJUNGNER.

XI. R E T R O S P E C T I V E S U R V E Y·

One of the purposes of our journey was to investigate the characteristic features of the plankton in southern South American lakes and check on the fl6ristic features pointed out by the author ( 1955) . In that paper written before our journey I directed attention to the absence of some widely distributed and common plankters in Valdivian and Tierra Fuegian lakes, e.g. Tabella1·ia spp. , Asterio

nella spp. , Oeratium spp . , Fragilaria crotonensis,

Melosira islandica, M. italica, Kellicottia longispina,

and K eratella quadrata.

Of these planlders M elosira italica has been recorded in the present material from Lake Frias and from a stream between Lake Gutierrez and Lake Nahuel Huapi. It seems to occur here and there along the Andean c;hain, e.g. in Lake Frey, situated about 1 km SW from the end of Brazo de la Tristeza in the Nahuel Huapi National Park {CLEVE-EULER,

1943) . Furthermore, it has been reported by FRENGUELLI ( 1942) from the National Reserve Copahue (Prov. Neuquen) north of the Nahuel Huapi National Park. KRASSKE ( 1939) has found it in South Chile. The occurrence of M elosira italica seems to be the only certain modification of the above mentioned statement.

Tabellaria flocculosa has been reported by KRASSKE (op. cit . ) from South Chile, but only as a few specimens and thus further investigations re-

garding this species are of importance. It has hitherto not been found in our collections from Valdivian lakes. OLIVER (1955) reports the occurrence of Tabellaria in the plankton of Salada Grande Lagoon, Province of Buenos Aires.

Asterionella gracillima has been reported by FRENGUELLI (op. cit . ) but in spite of the appended figures I am not convinced of the identification. Even the habitat is quite atypicaL Compare the figures under consideration (pl. 10, fig. 48, 49) and the figure of Diatoma elongatum var. fuegensis in CLEVE-EULER, 1948, fig. 5. Diatoma elongatum is a plant of frequent occurrence, but it is not noted in FRENGUELLI (op . cit . ) . I have studied this species from the Chilean lake district and from the N ahuel Huapi National Park; it resembles in shape the above mentioned figures, but the striae could often be observed only with difficulty. The Asterionella

gracillima figured in THOMASSON, 1955, is a doubtfull specimen; it may be a large Diatoma elongatum?

In lakes of the Chilean lake district I have hitherto not found any Asterionella. These diatoms seem to be absent in the an dine and preandine lakes of southern South. America.

From the lakes of the Altiplano down to the lakes of Tierra del Fuego there is no evidence of the occurrence of any Oeratium species, no Fragilaria

crotonensis, nor Kellicottia longispina. The southern


60 Retrospective survey

ljmit for the distribution of Keratella quadrata

seems to be the lakes on the Altiplano in Peru (THOM.ASSON, 1956) . Also in the waters in the Province of Buenos Aires there is no evidence of its occurrence, cf. 0LIVIER, 1955. Proceeding northwards along the Andean range it is interesting to note that Keratella quadrata has not been reported from Venezuela or Columbia by H.AUER (1956) . The same is the case with Kellicottia longispina. Both

TABLE 2. Distribution of phytoplankters.

Anabaena scheremetievi v. recta + Aphanocapsa elachista v. planctonica - + -

Ghroococcus limneticus + + - -

of these rotifers have somewhat erratic distributions C. minutus + -- + + -+ -+ -

and have been reported from Mexico. According to OsoRIO T.AF.ALL ( 1 942) Kellicottia longispina

seems to be rare, and for Keratella quadrata AHL

STROM ( 1 943) reports its occurrence near Mexico City. OsoRIO T.AF.ALL (op. cit . ) has not recorded this species.

Concerning the distributjon of Fragilaria cro

tonensis HuBER-PEST.ALOZZI ( 1942) is quite right in saying that "die geographische Verbreitung von Fragilaria crotonensis diirfte den Limnologen noch manches Ratsel zu lOsen geben", e.g. it has not been observed in South- and East-Africa, Sumatra or Java.

BREHM (1935) has pointed out a curious feature in the distribution of the genus Diaptomus in South America and has suggested the existence of W .AL

L.ACE's line as the limit for the range of this genus. Already in 1955 I had recorded Diaptomus in the Chilean lake district, and according to our rec�nt studies Diaptomidae are frequent in the lakes of this area. Probably due to the paltry material there are no records for the Nahuel Huapi area.

There are some more plankters common in the corresponding lake types of the northern hemisphere which seem to be absent in plankton samples from South America. I have mentioned here only some of the most striking ones. Further comments may be possible when the material from the Chilean collections is analysed. It should also be borne in mind that the conclusions based on rather random samples can never give an unalterable and convincing picture of the distribution of plankters.

In the following Table 2. I have indicated the distribution of phytoplankters in the lakes of Nahuel Huapi National Park. The dominants are indicated with - e , the subdominants with - () , abundant occurrence with - �-


Goelosphaerium kuetzingianum

Dactylococcopsis ellipsoideus

Gomphospaeria lacustris

M erismopedia glauca

M. tenuissima

Oscillatoria lacustris

0. tenuis

Peridinium cinctum

P. willei

Ghrysosphaerella sp. Dinobryon cylindricum

D. cylindricum v. palustre

D. divergens

D. divergens v. schauinslandii

D. sertularia

M allomonas acaroides

M. caudata

M. elliptica

M. elongata

M. tonsurata

Salpingoeca buetschlii

S. elegans

Stichogloea doederleinii

Gyclotella stelligera

G. stelligera v. elliptica

Gymatopleura solea

Diatoma elongatum

D. vulgare v. producta

Fragilaria sp. M elosira ambigua

M. distans

M. ita.lica

M. patagonica

M. pseudogranulata

Rhizosolenia eriensis

Surirella guatimalensis

S. robusta

S. tenera

Synedra acus

S. ulna

S. ulna v. danica

+ + + � +

- - - -+ • + -+

• + • • + + �

- - - -

+ -

--

+

-

+

+ +

+ +

+ +

+

+

+ + + +

+ + - + + -- + - -

- + - -+ - + -

+ + - + • + + -

+ + + + + + - - - +

+ + -

+ -

+ + + + + () + � + · - + - + - + + - + - - + - + -- + -

- + - + + - + + () -

Retrospect,ive survey 61

·a ·a ce ce ::I ::I 0 � � ;.a 0 � � ;.a Q;) .§ Q;) .§ - � � § � � Q;)

� � Q;) ::I .� 0 <ll <ll .� 0 s <ll 1'JJ

...c +" Cll ·s .� <ll � +" <ll .� <ll ce ::I ce � <l:l ::I ce ::I � <l:l z C:> � C:> � � z C:> � 0 � �

Asterococcus superbus + C. subspeciosum v. valid us + + + + +

Elakatothrix gelatinosa f. biplex + C. tetraophthalmum +

Eudorina elegans + + � Euastrum pinnatum +

Gemellicystis neglecta + H yalotheca dissiliens + +

Gloeococcus schroeteri + + + M icrasterias radians +

Pandorina morum + M. truncata +

Paulschulzia pseudovolvox + + Sphaerozosma vertebratum +

Schizochlamys sp. + Staurastrum avicula v. subarcuatum +

Tetraspora lacustris + St. dilatatum + St. dispar + St. excavatum v. minimum + + +

Ankistrodesmus falcatus + + + St. furcigerum + + + + A . falcatus v. spiralis + St. laeve + Botryococcus brauni-i + () � + + St. longipes v. evolutum + + + + Ooelastrum proboscideum + St. manfeldtii f. + Crucigenia quadrata + St. margaritaceum + 0. rectangularis + + St. pilosum + Dictyosphaerium pulchellum + + + St. planctonicum v. ornatum + + Hofmania lauterbornii + St. pseudosebaldi + Kirchneriella obesa + St. punctulatum v. elUpticum + N ephrocytium limneticum - + St. rotu.la v. smithii + + N. lunatum + + St. sebaldi v. brasiliense f. + Oocystis crassa + St. seb. v. orientale f. + 0. elliptica + St. seb. v. ornatum f. planctonica + + 0. lacustris + St. tetracerum v. biverruciferum + 0. solitaria + + + St. tohopekaligense v. bre1;ispinum + 0. submarina v. variabilis + St. trachytithophorum + Pediastrum araneosum v. rugulosum + Staurodesmus dickiei + P. boryanum + + Std. phimus + P. boryanum v. longicorne + Std. megacanthus + + P. duplex + Std. sellatus f. J"acobsenii + P. integrum f. granulata + Std. triangularis f. stroemii + + + P. integrum v. scutum + Xanthidium antilopaeum -i- + Scenedesmus quadricauda + Olosterium acerosum + 0. aciculare + +

0. kuetzingii + + + The species lists for phytoplankton communities

Oosmarium abbreviatum v. show that all these lakes are of oligotrophic char-planctonicum + acter. The most oligotrophic seems to be Lake

0. botrytis -i- + + + Frias as far as can be judged from a random sample. 0. brebissonii + The most eutrophic is the Lake Hess, which on C. connatum + the whole shows a divergent character due to the 0. contractu m v. ellipsoideum + + C. depressum v. achondrum + + exceptional hydrographic conditions. These circum-0. margaritatum v. subtumidum + stances are reflected also in the composition of the C. moniliformis + present plankton-community. C. pseudoconnatum v. ellipsoideum + The Chrysophyceae are dominant in the phyto-C. quadrifarium + +

plankton of four out of six lakes. In all cases the 0. quadrum + C. subcucumis + dominant species belongs to the genus Dinobryon.

0. subdanicum + The plankton of Lake Nahuel Huapi, Lake


62 Retrospective survey

Mascardi, and Lake Guillelmo is dominated by Dinobryon divergens. Only in Lake Gutierrez is this species surpassed by Dinobryon cylindricum. The remaining genera of Ohrysophyceae are of no importance in the plankton of these lakes.

The Oyanophyta are qualitatively as well as quantitatively sparsely represented in the plankton of these lakes. The same could be said about the Pyrrophyta, only Peridinium willei is of regular occurrence. There are only a few diatoms which are of importance from the quantitative point of view. These are Rhizosolenia eriensis, M elosira pseudo

granulata, Synedra ulna, and Diatoma elongatum.

The two last named diatoms are present in considerable quantities only in Lake Frias. Among the V olvocales only Eudorina elegans and Gloeococcus

schroeteri are common. Many species of Ohloro

coccales are tabulated above, b�t they all occur only sparsely in the plankton of these lakes. The list of desmids is long, but there are f�w constantly occurring species. The greater part of the desmids tabulated above are quite rare in the plankton of the lakes and of practically no influence on the character of the present plankton communities. The few taxa among planktic desmids which occur in noteworthy quantities are Staurastrum

longipes var. evolutum, Staurastrum sebaldi varieties, and especially in Lake Nahuel Huapi Staurastrum

rotula var. smithii . It is remarkable that species of Euastrum, Micrasterias, and Xanthidium are rarely to be found. The absence of plants belonging to the Staurastrum paradoxum and Staurastrum anatinum

groups, and many of common Staurodesmus species, e.g. cuspidatus, curvatus, and dejectus, is difficult to explain. There is no evidence that these plants avoid oligotrophic habitats and they have . been recorded from Tierra del Fuego as well as from the Chilean lake district (THOMASSON, 1955) . The above mentioned Staurodesmus species are, judging from

· Scandinavian mountain lakes , quite rare in pronounced oligotrophic lakes. However, at least taxa belonging to Staurastrum paradoxum and Stauras

trum anatinum groups were expected in the lakes of Nahuel Huapi National Park.

In the following paragraphs an attempt will be made to outline some comparative features of the phytoplankton composition of the lakes in southern


South America. The material which could be used for such a comparison is rather scanty.

Between the lakes in the Nahuel Huapi National Park and those in Tierra del Fuego there are some resemblances. The lakes in the Nahuel Huapi National Park and also La go Fagnano and Ca be cera Lago, cf. THOMASSON, 1955, are of oligotrophic character and have their plankton dominated by Dinobryon. The other components of the plankton show many similarities.

Nothing seems to be known about the large Andean lakes situated between Tierra del Fuego and the Nahuel Huapi area, e.g. L. Argentino, L. Viedma, L. San Martin, L. Cochrane, and L.

Buenos Aires. The only note from this area is by KRASSKE ( 1939) who reports Dinobryon-plankton from the oligotrophic Lake Riso Patron ( 130 m a. s . -1. ) , situated near Puerto Puyuhuapi, Prov. Aysen, 72°36'W longitude and 44 °25'S latitude.

On the western side of the Andean range the Chilean lake district corresponds to the Argentine lakes in Nahuel Huapi and Lanin1 national parks. On the lakes of the Chilean lake district there are some notes published by KRASSKE ( 1939) and THOMASSON ( 1955) . These lakes were the main object of the Swedish Limnological Expedition to South America in 1953-54. The collections of this expeQition are still being analysed. Therefore I must confine my remarks to some preliminary notes. about the phytoplankton of these lakes. From Lake Mansa, situated near Puerto Montt, KRASSKE (op . cit . ) has recorded Dinobryon-plankton in December 1935. Dinobryon-plankton seems to be characteristic . of the following lakes visited by us: Pellaifa, Quillehue, Pichilafquen, and Huilipilun. They are all small and have a limited catchment area. In the plankton of large lakes situated in the longitudinal valley, e.g. Villarrica, Calafquen, Panguipulli, Riiiihue, Ranco, Puyehue, Rupanco, and Llanquihue, Melos�ra plays an important role. It is also found in the beautiful Lake Todos los Santos, situated between Lake Llanquihue and Lake Nahuel Huapi, and surrounded by volcanoes. In the plankton of the five last mentioned lakes Dino-

1 Lanin National Park connects in the north with the Nahuel Huapi National Park.

Retrospective survey 63

bryon is frequent and it may be that the plankton of these lakes in early summer is dominated by Dinobryon. This idea is borne out of the observations on the plankton of Lake Llanquihue, which on December 25, 1953, had a Dinobryon plankton . O n February 10 , 1954, this dominance o f Dinobryon

was repla?ed by M elosira. But that supposition is only tentative since there were no year-round observations on the succession of the different facies of plankton-communities.

Summing up, we can say that there are many common features between the plankton communities of Nahuel Huapi area and the Chilean lake district. Nevertheless it seems that the plankton of the Chilean lakes represents a somewhat higher trophic level, and in some small lakes it is of a quite strange character. A good indicator in the plankton of the Chilean lake district seems to be M elosira hustedti, common to most of the lakes. It is endemic in the Valdivian area and does not seem to cross the Andean range. It has not been recorded from the lakes of Nahuel Huapi National Park.

Only a few data on the chemical composition of the waters in the Nahuel Huapi area are available. If we compare these with the results of water analyses from the Chilean lake district (LoFFLER,

1959) we find that similar low values are common to both regions. But to our astonishment these low values are often accompanied by rather high quantities of plankton. This may be partly due to considerably higher epilimnic temperatures in the lakes under consideration. It is also probable that the results of chemical analyses on the water from these lakes, characterized by high plankton production do not reflect the real resources of nutrient

substances. In these lakes of the Chilean lake district, e.g. Lake Villarrica, throughout the year large quantities of nutrients are bound in living organisms and will immediately after decay be incorporated by other plankters. It thus seems that the plankton is a much better indicator of the trophic level, than the chemical analyses, since it also reflects climatic factors; e.g. the above mentioned Lake Villarrica is according

. to chemical

analyses a decidedly oligotrophic lake, but according to the quality and quantity of the plankt.on this lake is closest to the mesotrophic lakes. This seems to be partly due to high water temperatures which promotes reduction processes in the deeper layers. The nutrative substances liberated are immediately incorporated into the biomass. The very opposite conditions prevail in cold water lakes where considerable quantities of nutrients are deposited in sediments and not involved in further production. The effective decay is rendered difficult by

. low

bottom temperatures. A factor of some importance for the trophic level of the lakes in the Chilean lake district seems to be the violent winds resulting in tutbulences influencing water layers down to considerable depths. This turbulence contributes to the redistribution of nutrative substances released in deeper layers.

In the lakes of the Nahuel Huapi National Park . the low temperatures of the water together with the low supply of nutrients seem to prevent higher production of planldon and the occurrence of plankters of a demanding character. All the lakes studied by us in the National Park of Nahuel Huapi are oligotrophic as distinguished from many of the lakes on the western side of the Andean range which are on a somewhat higher trophic level.


64 Taxonomical comments

X I I. TAX O N O M I C AL C O J\II M E N T S

During my work on the samples collected from the National Park of Nahuel Huapi many notes of taxonomical character were accumulated and a great number of drawings of microscopical freshwater organisms were made. A part of these notes and drawings are presented on the following pages. I · hope they will contribute to the knowledge of the microorganisms in the waters of the Nahuel Huapi National Park, and call forth discussion on the systematic position of some doubtful taxa mentioned below.

I have included a large number of drawings partly because I hope that they will be of some use for the further investigation of the plankters in this area, and partly because there are . many troublesome algae whose names are meaningless without an illustration. This often makes it difficult to evaluate the lists of freshwater plankters; there are many vague conceptions about numerous common plankters. Therefore it is of great importance to depict as many as possible or to give a reference to an already published identical figure, cf. also LuND & TALLING, 1957, p. 491 . In many

cases such records are useful for further taxonomical, geographical, and ecological work. The uncertainty of a great many records makes it very difficult to make a statement on the ecology of numerous planktic algae. The matter is still more complicated through the existence of ecotypes which as a rule cannot be identified morphologically.

The often rather inadequate knowledge of the taxonomic position of the algae used in culture experiments sometimes makes one hestitate in drawing conclusions about the ecology of these algae. There are many species of quite different ecological valence which are labeled in cultures under the names of collective species.

In consequence of the importance of taxonomic work for ecology I would like to give prominence to the very suggestive chapter on relations of systematics to ecology and limnology published recently by HusTEDT (cf. HusTEDT, 1957, p. 200) . Summing up he says: " Ohne Okologie gibt es Imine Limnologie, und keine Okologie ohne Systematik! " . I

would only like to say limnobiology instead of limnology.

1. M I C R O P H YT A

Lyngbya sp.

Fig. 20: l . This plant was collected from Lake Gutierrez . It seems to be related to Lyngbya

ceylanica WILLE .

Nostoc sp. Fig. 20: 2. Filaments irregularly interwoven in gelatinous sphaerical colonies, 58 fl in diameter. The surface of colonies distinct and smooth. Cells 1 .5-2 f.1, broad and 6 p long. Occurs in Lake Guillelmo. Compare N ostoc paludosum fa. longius

KosSINSKAJA, 1936, fig. 2, which, however, is a larger species.

Phacus orbicularis HUBN.

Fig. 20: 4. Length 42 fk, breadth 30 fl· Pond near military camp in Bariloche .


Gymnodinium sp .

Fig. 20: 8. Length 18 f.J,, breadth 18 fl· Rare in the plankton of Lake Gutierrez . This plant somewhat resembles Gymnodinium neglectum (SCHILLING)

LINDEM. No platelets have been discerned in the periplast of this specimen.

Salpingoeca elegans (BACHM. ) LEMM.

Fig. 20: 7. This plant occurs in the colonies of Gloeococcus schroeteri in the plankton of Lake Nahuel Huapi. As a rule the lorica has one thin filamentous stalk, only in few cases were two observed. Compare Salpingoeca buetschlii LEMM.

in SKUJA, 1948, pl. 34, fig. 12-15, and Salpingoeca

convolvulus SKuJA, 1939, pi. 2, fig. 14.

Taxonomical comments 65

�11 Fig. 20: 1. Lyngbya sp. 5. M elosira pseudogranulata 9. Scapholeberis spinifera

2. Nostoc sp. 6 . Neidium sp. 10. Micractinium sp. 3 . Scenedesmus bijugatus 7. Salpingoeca elegans 1 1 . Ohrysopyxis paludosa

4. Phacus o·rbicularis 8. Gymnodinium sp.

Diatoma elongatum (LYNGB. ) AG. Already in 1909 0. MuLLER has recorded the

Frequent plankter in the lakes of the N ahuel

Huapi area. The valve of my specimens resembles

the one pictured in MAYER, 1919, pl. 5, fig. 33

under the name of Diatoma tenue var. elongatum

(AG.) GRUN.

M elosira pseudogranulata A. CL.

This species . was described by AsTRID CLEVE

EuLER in 1948, p. 6, pl. 1, fig. 3, from Tierra del

Fuego. All the specimens of Melosira granulata

type occurring in my samples from the National

Park of Nahuel Huapi are identical with Melosira

pseudogranulata. This species is characterized by

its pores occurring in striae running parallel with

pervalvar axis. The proximal ends in all cells of

the filament bear powerful spines, just like those

in M elosira granulata RALFS. The taxonomic position and rank of this

taxon ought to be cleared up. Fig. 20: 5 shows an

anomalous arrangement of pores, diameter of cell

is l l p;.

occurrence of Melosira granulata (EHRENB. )

RALFS var. characterized b y straight lines of

pores from South-Patagonia.

. Pinnularia macrocephala A. CL.

This diatom has been recorded in a bottom

sample collected from Lake Guillelmo. It was

described by AsTRID CLEVE-EULER, 1943, p. 224,

fig. 7. Compare also Pinnularia sp . in KRASSKE,

1939, pl. 1 1 , fig. 39, which is according A.

CLEVE-EULER synonymous with her species. The

length of my specimen is 109 p;, and it seems to

form a transgression to Pinnularia polygona

(BREB. ) 0. MULLER pictured in FRENGUELLI,

1942, pl. 4, fig. 32. Also this species is represented

in the present sample, its length is 107.6 p;. Both

the above mentioned diatoms seem to be closely

related and may likely be united.

Synedra ulna (NITZSCH) EHRENB.

I n a brook a t Lake Mascardi a curious form of

this species was observed. Seen from the valve



side it · was undulated in its whole length, just

like an Indonesian dagger with its flame-like

blade.

Ghrysopyxis paludosa FoTT.

Fig. 20: 1 1 . Length of lorica 9 p,. Found in the

submersion pool between Lake Gutierrez and

Lake Mascardi. Compare pl. I, fig. 2, in FoTT,

1957.

Botryococcus braunii ](uTz.

The systematic position of this plant is not

definitely established. It has wavered between

Heterokontae and Ghlorococcales, cf. SKUJA, 1948,

p. 339. My impression, mainly based on speci

mens from Mrican lakes, has been that this plant

belongs to the ii eterokontae. It is included here

amongst the Chlorococcales as a result of the

recent investigations by BLECHER & FoGG, 1955.

M icractinium sp.

Fig. 20: 10. In the plankton of Lake Nahuel

Huapi, a hitherto unidentified Micractiniaceae

was observed. · This plant was of frequent oc

currence, usually 2 or 4 plants occurred to

gether. The diameter of cells is 6.25 p,, the colour

of chromatophore lustrous golden; cells covered

by delicate long spines. For the identification an

investigation of living specimens is necessary.

Dr. BouRRELLY has kindly examined this plant,

he writes: "Apres action de la solution iodo

ioduree, on observe de l'amidon et 1 pyrenoide

par cellule. La membrane, les aiguillons et la

gelee se colorent en rose par le Rouge de Ruthe

nium. Avec, l 'encre de Chine on ·voit une large

aureole gelatineuse autour des colonies. "

Scenedesmus bijugatus (TURP. ) KuTz.

Fig. 20: 3. Length of coenobia 37 .5 fl· Collected

from the submersion pool between Lake Gutierrez

and Lake Mascardi. Compare pl. 28, fig. 29-32 in

SKUJA, 1956. My specimens have marked thick

nings of cell wall at the poles. According to the

nomenclature introduced by CHODAT (1926) this

form corresponds with Scenedesmus ovalternus

CHODAT.

Acta Phy togeogr. Suec. 42

Scenedesmus ovalternus var. gravenitzii {BERNARD)

CHODAT.

This plant has been frequently noted in a sample

collected from a strand-pool on the beach of

Lake Nahuel Huapi. It is the impression of the

present writer that this plant is just a benthic

life-form of Scenedesmus bijugatus. The colonies

are held together with the remains of mother-cell

wall, the result is large conglomerations of cells.

See also GRONBLAD, 1 934, p. 270.

Gosmarium amoenum var. intumescens NoRDST.

Fig. 23: 5, a specimen from a tarn in Paso Vuri

loche. Compare also Gosmarium amoenum var.

mediolaeve N ORDST. and Cosmarium tumens

· N ORDST. : the last one is considerably smaller.

Cosmarium botrytis (BoRY) MENEGH.

Fig. 21 : 1 6 shows an anomalous course of divi

sion observed in a sample from the pond near

military camp.

Gosmarium controversum WEST fa. major n . fa.

This form which occurs in the bog tarn in Paso

Vuriloche is characterized by its large dimensions:

length 106-116 p,, breadth 81-91 p,, some of the

specimens observed have a smooth apex.

Cosmarium depressum var. pla;nctonicum REV.

Fig. 23: 20-21 . Length 1 2 p,, breadth 13 .5 p,. From submersion pool between Lake Gutierrez

and Lake Mascardi. Compare also Cosmarium

subarctoum fa. punctata WEST & WEST.

Cosmarium laeve RABENH. fa.

The form noted in a bottom sample from Lake

Guillelmo is an intermediate form between the

main species and its variety septentrionale WILLE.

Cosmarium margaritatum var. subtumidum.

This species was observed in the plankton of

Lake Gutierrez. Length of my specimen is 125 p,, the punctulation between granulas is very

marked, compare WEsT's Monograph, vol. .4,

p . 19.

Cosmarium phaseolus BREB.

Fig. 21 : 19 . Length 36 p,, breadth 30 fl· My spe

cimen is very close to the specimen figured in


CROASDALE, 1956, pl. 4, fig. 6. Compare also

Oosmarium subtumidum var. pachydermum PRES

COTT & SCOTT, 1952, pl. 1, fig. 18 .

Cosmarium reniforme var. alaskanum CROASDALE fa.

Fig. 2 1 : 8. Length 62.5 fl, breadth 27 .5 p,. From the

pond near military camp. I have regarded my

specimens as related to Oosmarium reniforme var.

alaskanum CROASDALE, 1956, pl. 13 , fig. 4. They

should be compared with some forms of Oosma

rium portianum ARCH. , e.g. Cosmarium portianum

fa. in KRIEGER, 1932, pl. 1 1 , fig. 16, and Cosma

rium portianum in BoRGE, 1918, pl. 2, fig. 19.

Moreover there is some resemblance to Cosmarium

margaritiferum TURP . b. incisum KIRCH. in

BoLDT, 1888, pl. 2, fig. 28, and to Oosmarium

perincisum GRONBLAD, 1921 , pl. 6, fig. 24.

Cosmarium tetraophthalmum BREB.

Fig. 23: 6. Length 125 fl, from the plankton of

Lake Mascardi. It resembles in many respects

Cosmarium baccatum ScoTT & GRONBLAD, 1957,

pl. 6, fig. 2 & 3, which is, however, considerably

smaller.

Euastrum attenuatum var. lithuanicum W OLOSZ.

Fig. 22: 8. �ength 54 p,, breadth 37.5 p,. Drawn

from a specimen from a bottom sample from

Lake Guillelmo. This specimen is obviously more

closely related to var. lithuanicum WoLosz. than .

to the main species. The main species has hitherto

been reported from North- and South-America,

and var. lithuanicum only from Poland; the var.

splendens (FRITSCH & RICH) ScoTT & GRoNBLAD

has been reported from Africa.

Euastrum bidentatum var. speciosum (BoLDT)

ScHM. fa.

Fig. 22: 9. Length '57 .5 p,, breadth 40 p,. From the

submersion pool between the lakes Mascardi and

Gutierrez. Compare pl. 1 1 , fig. 1 6, in TURNER,

1892, and figure of Euastrum rostratum RALFS fa.

in WITTROCK & NORDSTEDT, 1893, pl. 195.

$uastrum dubium var. latum KRIEGER.

Fig. 2 1 : 1 1 . Length 25 p,, breadth 23.7 p,. From a

. tarn in Paso Vuriloche. Compare pl . 79, fig. 7, in

KRIEGER, 1937.

Euastrum pinnatum RALFS.

Fig. 2 1 : 2 1 . Length 141 p,, breadth 75 p,. Lake

Mascardi.

Euastrum turneri var. bohemicum LiJTKEM.

Fig. 22: 7 . Length 43 p,, breadth 34 p,. Occurring

among the filamentous algae in the little river

running from Lake Gutierrez into Lake N ahuel

Huapi.

M icrasterias denticulata BREB.

Fig. 21: 20. Length 279 p,, breadth 252 p,. Lake

Hess. Compare pl. 3, fig. 7 in TAYLOR, 1935.

Micrasterias sol var. elegantior G. S. WEST.

Fig. 2 1 : 13. Length 173 fl, breadth 1 68 fl· Lake

Hess.

Staurastrum brebissonii ARCH.

Fig . 2 1 : 5.' Length 44ft; breadth 42.5-45p,. Sampled

from amongst the tufts of filamentous algae in.

the brook between Lake Gutierrez and Lake

Nahuel Huapi.

Staurastrum dilatatum EHRENB.

Fig. 23: 2. Length 59 fl, breadth 64 fl· Tarn in Paso

Vuriloche. Compare pl. 126, fig. 14, in WEST &

WEST, 1912. The alternans-dilatatum-striolatum

group seems to be in need of revision.

Staurastrum erasum BREB.

Fig. 23: 19. Length 34 fl, breadth 36 p,. From the

artificial pond situated west of military camp.

For the distinction between Staurastrum brebis

sonii ARCH . and Staurastrum erasum BREB . , see

GRONBLAD, 1942, p. 40. My specimen should

be compared with Staurastrum gladiosum, it

resembles also Staurastrum erasum in BoRGE , 1913, p. 32, .. pl. 3, fig. 35, except in size.

Staurastrum excavatum var. planctonicum KRIEGER.

Fig. 23: 12 . Length with processes 34 fl, without

processes 13 .5 fl, breadth 42 p, . Also tri-radiate

forms have been observed. The plant depicted

originates from the pond near the military camp .

It ought to be noted that there are many forms

among the plants of Staurastrum tetracerum

RALFS which in many respects resemble the

algae belonging to Staur�strum excavatum-group.



Compare also Staurastrum americanum . var.

excavatum ScoTT & GRONBLAD, 1957, pi. 26, fig.

10, and Staurastrum excavatum WEST & WEST fa.

in FRITSCH & RICH, 1 937, fig. 22e.

Staurastrum· laeve RALFS.

Fig. 23: 14. Length 23.2 fl, breadth 41 .2 fl, Lake

Frias. Compare Staurastrum fissum TURNER,

1892, pi. 14, fig. 24, . Staurastrum fissum var.

perfissum WEsT & WEST, 1895, pi. 8, fig. 23,

KosSINSKAJA, 1936, pi. 2, fig. 10, and GRONBLAD ,

1920, pi. 3, fig. 70 & 7 1 . It seems that all these

records on Staurastrum fissum var. perfissum had

better to be included within the rather variable

Staurastrum laeve.

Staurastrum longipes var. evolutum (WEST & WEsT)

THOM.

Fig. 23: 10-1 1 . Length without processes 12 .5 fl, with processes 50 fl, breadth 50 fl· Frequent

plankter in the lakes of Nahuel Huapi National

Park. Compare ·THOMASSON, 1955, fig. 3: 5 .

Staurastrum manfeldtii DELP. fa.

Fig. 23: 7. Length 40 fl, breadth 49 #· Lake

Mascardi. With great dubiety I have assigned

this specimen to the Staurastrum manfeldtii

group. �t should be compared with Staurastrum sebaldi var. ornatum fa. novizelandicum in N Y

GAARD, 1926, pi. 5, fig. 46, which is, however,

somewhat larger. The present specimen resembles

in many respects the specimen of Staurastrum sebaldi var. ornatum N ORDST. figured in the pres

ent paper, fig. 2 1 : 2, which is, however, somewhat

larger and has more pr<;mounced apical orna

mentation .

The occurrence of intermediate forms and the

lack of distinctive characters between Stauras-

trum menfeldtii-group and plants grouped under

Staurastrum sebaldi var. ornatum N ORDST. make

their identification rather a matter of personal

taste. Perhaps a more strict consideration of the

shape of the basal part of semicells is to be de

sired. In· this case the plants with cylindrical basal

half of semicells belong to Staurastrum manfeldtii

group, and plants with conical, cup-shaped basal

part belong to Staurastrum sebaldi-group, as has

been pointed out by TElLING, 194 7, p. 223. Such

a procedure would lead to that many taxa which

are hitherto regarded as belonging to Staurastrum

sebaldi var. ornatum NoRDST. are to be transferred

to Staurastrum manfeldtii. In a few cases the

plants labelled as Staurastrum manfeldtii would be

transferred to Staurastrum sebaldi-group, e.g. the

Staurastrum manfeldtii in FLORIN, 1957. But I

am afraid that the occurrence of intermediate

forms would place a stumbling-block in the way

of distinction.

The problem of Staurastrum manfeldtii DELP. ,

as well a s that o f Staurastrum. luetkemuelleri

DoNAT, v. i . , is a good illustration of the necessity

for every description to be accompanied by a

good drawing, for any verbal diagnosis alone is of

minor value. Probably the only way out of this

dilemma is to accept only those species which

have been established thr�ugh a good figure

which gives an undisputable interpretation of

the plant in question. The relations between

Staurastrum manfeldtii DELP.- and Staurastrum

luetkemuelleri DoNAT-groups to the Staurastrum

sebaldi-group have been the object of·

many

discussions. However, the result is rather poor.

Recently CROASDALE, 1957, published an

interesting picture (fig. 121) of Staurastrum

manfeldtii var. parvum MESSIK. with a reference

Fig. 21 : I . Staurastrum sebaldi var. or- 9. Staurastrum margaritaceum fa. 1 6 . Oosmarium botrytis

natum fa. planctonica

2. St. sebaldi var. ·ornatum fa. 3. St. sp. 4. St. subpolymorphum fa. 5 .. St. brebissonii

6. St. subpolymorphum fa. 7. Staurodesmus defectus

8. Oosmarium reniforme · var. alaskanum fa.

A.cta Phytogeogr. Suec. 42

1 0. St. sebaldi var. ornatum fa. planc

tonica

. 1 1 . Euastrum dubium var. latum

1 2. Staurastrum planctonicum var. ornatum �a.

1 3. Micrasterias sol va.r. elegantior

1 4. Staurodesmus mucronatus var. subtriangulare

1 5 . Staurastrum suborbiculare

17. Staurodesmus convergens

1 8. Std. glabrus fac. glabrus

1 9. Oosmarium phaseolus

20. M icrasterias denticulata

2 1 . . Euastrum pinnatum

22. Staurastrum sebaldi var. ornatum

fa. planctonica

23. Staurodesmus megacanthus fa.

Fig. 1 - 10, 14-18, 2 1

0

5 - 58882 9 Kuno Th

-

omasson

50Jl Fig. 1 1, 1 9

Taxonomical eo mments 69

� �Q 2 1 �a · CO) 23 b

F i g . 22,23 0 50 �

A cta Ph t y ogeogr. Sue.c. 42


to fig. 140 & 141 in GRONBLAD, 1956. However,

-this reference would have been in its proper

place according her fig. 122 of Staurastrum

luetkemuelleri fa. which has much more in

common with the specimen figured by GRON

BLAD. Both of them are to be considered as be

longing to Staurastrum luetkemuelleri-group.

Staurastrum margaritaceum (EHRENB. ) MENEGH. fa.

Fig. 21 : 9. Length 40 fl, breadth 40 fl· This 6-radi

ate specimen belongs to the rather variable group

of algae brought together under the name of

Staurastrum margaritaceum (EHRENB.) MENEGH.

Compare Staurastrum margaritaceum var. hirtum

NoRDST. in FuKUSHIMA & FuJISAWA, 1954, fig.

5 J . .

Staurastrum perundulatum GRONBLAD fa.

Fig. 23: 9. Length with processes 28 fl, without

processes 9.3 fl, breadth 24 fl· Collected from the

pond near military camp. Compare Staurastrum

pseudoiotanum GRONBLAD, 1 921 , pl. 5, fig. 36 &

37, Staurastrum irregulare WEST in WEST &

CARTER, 1923, pl. 149, fig. 7, and Staurastrum

perundulatum GRONBLAD in SKUJA, 1949, pi. 34,

fig. 16 & 17.

Staurastrum planctonicum var. ornatum (GRON

BLAD) TElLING .

I just wonder how many algologists have puzzled

their brains over the distinction between the

array of forms grouped around Staurastrum

sebaldi var. ornatum fa. planctonica (LUTKEM. )

TEILING, and Staurastrum planctonicum var.

ornatum (GRONBLAD) TEILINq. These forms are

common plankters and often noted in plankton

samples, yet in spite of this one is always as

nonplussed as to the precise distinctions between

these taxa.

TElLING, 1 94 7, has built up an evolutional

series with Staurastrum sebaldi var. ornatum

NORDST. as starting point. In a recently published

paper (THOMASSON, 1957) another possible direc

tion of evolution has been suggested, though

such speculations seem to be of little value.

However, sound advice of the eminent planktolo

gist TElLING (op . cit.) should be borne in mind:

"It is however a good thing to remember that

our present taxonomy is a more stock-taking

and denomination of the material, but it is not

founded upon genetic facts" .

For the future rearrangement of the above

mentioned forms it is of great importance that as

many forms as possible will be depicted and the

figures published for the future establishing the

width of variation in different populations and

between these populations.

As representative figures of Staurastrum planc

tonicum-group I consider the following ones:

TElLING, I946, fig. 30 & 32 SKUJA, I948, pl. I 6, fig. 9 NYGAARD, I 949, fig. 56 PRESCOTT, I952, pi. 5, fig. I FLORIN, I 957, figs. 27: 7, 28: I , 34: IO, 35: 2

Amongst these plants the var. ornatum (GR6N

BLAD) TEILING with pronounced ornamentation

on the basal part of semicells could be singled

out. Moreover we have a good variety namely

the var. bullatum TElLING. It will be evident

from the pictures listed above, that the main

criterion has been the shape of the semi cells. The

apical ornamentation is variable, and is not

applicable for the strict distinction between the

algae belonging to Staurastrum planctonicum

group and to Staurastrum sebaldi var. ornatum

group. It seems to be impossible to restrict in

Staurastrum planctonicum the number of verrucae

on each edge of the vertex to two. Also the

character of endspines of the processes, which

has been considered as main characteristicum by

some authors, is of less value. The prominent

Fig. 22: 1-3 . Staurastrum rotula var. smithii

6. St. sebaldi var. ornatum 10. Staurastrum sebaldi var. brasili-

4. M icrasterias truncata var. pusilla

5. Staurastrum pseudosebaldi

fa.

· A cta Phytogeogr. Suec. 42

7. Euastrum turneri var. bohemicum

8. Euastrum attenuatum var. lithua

nicum

9. E. bidentatum var. speciosum fa.

ense fa.

1 1 . St. planctonicum var. ornatum

1 2-13. St. sebaldi var. brasiliense fa. Scale cf. fig. 2 1 : 1 1 , 1 9.

Taxonomical comments 7 1



ventral spine which is usually directed in the

direction of process occurs also in Staurastrum

sebaldi var. ornatum-group.

Amongst the plants collected from the N ahuel

Huapi National Park the specimen from Lake

Nahuel Huapi, fig. 22: 1 1 , length 67 .5 p,, breadth

1 10 p,, strongly resembles Staurastrum planctoni

cum and could well be considered as belonging

to this group. The specimens from Lake Gutierrez,

figs. 21 : 1 and 21 : 10, length 64 p,, breadth 106 p,, are to be considered as belonging to Staurastrum

sebaldi var. ornatum fa. planctonica (LUTKEM.)

TElLING, cf. TElLING, 1947, fig. 20. To this taxon

should also be included the specimen figured in

FLORIN, 1957, fig. 27 : 1 . Probably is it better to

consider the fa. planctonica as a taxon of higher

rank, at least as a variety of its own.

The plant from Lake Gutierrez, fig. 2 1 : 22, length

52.5 p,, breadth 106 p,, is Staurastrum sebaldi var.

ornatum fa. planctonica, but a form with rela

tively longer processes. It is rather difficult to

find a proper place for the plants from Lake

Gutierrez, figs. 22: 10, 12, and 13, breadth 77.5-

100 fl· Obviously they belong to Staurastrum

sebaldi -group , though the apical ornamentation

is reduced and processes are converging. As a pre

liminary assessment I have labelled these plants

as Staurastrum ·sebaldi var. brasiliense B0RGES. fa.

Compare also fig. 3 in LuTKEMULLER, 1910.

Staurastrum pseudonam.tm GRONBLAD fa.

Fig. 23: 13 . Length 32 p,, breadth 41 p,. Rare in

the plankton of Lake Frias. This species was de

scribed by GRONBLAD, 1920, p. 74, and should be

compared with Staurastrum aciculiferum (W.

WEsT) BoRGE in RuzicKA, 1954, fig. 19, which is

a stouter plant of somewhat minor size.

Fig. 23: 1 . Staurastrum trapezicum 8. St. sp.

Staurastrum pseudosebaldi WILLE fa.

Fig. 22 : 5. Length 50 p,, breadth 68 fl· Lake Hess.

Compare THOMASSON, 1955, fig. 3 : 8, Staurastrum

pseudosebaldi var. planctonicum TElLING, 1947,

figs. 18 & 19, and Staurastrum pseudosebaldi var.

elongatum MESSIKOMMER, 1957, fig. 27 .

Staurastrum punctulatum var. kjellmani WILLE.

Fig. 23 : 4. Length 37-44 p,, breadth 26-28.5 p,. Noted from the submersion pool between Lake

Gutierrez and Lake Mascardi. Compare WEST &

WEST, 1912, pl. 127, fig. 18 . Also specimens of

more normal appearance were observed.

Staurastrum rotula var. smithii (G. M. SMITH)

THOM.

Figs. 22 : 1 and 3. A common plankter in Lake

N ahuel Huapi, furthermore it was noted in

Lake Hess. In the lakes of Chilean lake district,

e.g. in Lake Villarrica a form with pronounced

upward bent to its processes occurs.

I have mentioned these specimens already in

an earlier paper (THOMASSON, 1957 ) . This algae

seems to be rather variable in shape. The popula

tion of Lake Nahuel Huapi has shorter processes,

and moreover the spines tipping the processes are

also considerably shorter than in the specimens

from Lake Bangweulu (THOMASSON, op. cit. ) .

In spite of this I a m not inclined to make a new

forma or variety, because it seems to be impos

sible to make a clear distinction between the

different types. In fig. 22: 2 .I have depicted an

extreme type from Lake Nahuel Huapi. When

comparing this with those occurring in Lake

Bangweulu one would be inclined to separate

them as distinct varieties, but having studied a

large amount of material containing similar

1 7. St. sp. 2. St. dilatatum 9. St. perundulatum fa. 1 8. St. subpolymorphum fa. 3 . St. sebaldi var. oriental·is fa. 4. St. punctulatum var. kjell

mani

5. Oosmarium amoenum var. intumescens

6. 0. teraophthalmum

7. Straurastrum manfeldtii

.Acta Phytogeogr. Suec. 42

1 0- 1 1 . St. longipes var. evolutum

1 2 . St. excavatum var. planctonicum

1 3 . Staurastrum pseudonanum fa. 1 4. St. laeve

15. St. sebaldi var. brasiliense

1 6 . St. sebaldi var. ornatum fa. planc

tonica

1 9. St. erasum

20-2 1 . Oosmarium depressum var. planctonicum

22. Staurodesmus phimus

23. Std. tr·iangularis var. convergens

24. Std. dejectus

Scale cf. fig. 2 1 : l l 19.

74 I Taxonomical comments

specimens it will be evident that the forms like

those figured in figs. 22 : 1-3 are related ones,

and that all of these · forms are connected to

each other by intermediate forms. Describing

still more varieties and forms will only add further

complications for future taxonomy.

Staurastrum sebaldi var. brasiliense B0RGES.

Fig. 23: 15. Length 44fl, breadth 62.5ft. From the

pond near military camp. Compare Staurastrum

sebaldi var. brasiliense in GR6NBLAD, 1945, fig.

265, Staurastrum sebaldi var. brasiliense in THO

MASSON, 1955, fig. 2: 3, and Staurastrum sebaldi

var. corpulentum ScoTT & GRONBLAD, 1 957, pl.

32, fig. 13.

Staurastrum sebaldi var. orientalis (WEST & WEST)

n. comb . fa.

Fig. 23: 3. Length 41 ft , breadth 56 11-· Noted from

Lake Guillelmo and from the pond near the mili

tary camp . I have considered this plant as related

to the forma described by WEST, 1907 , pl. 16,

fig. 9 . My plant differs from WEsT's by its some

what converging processes. It should be compared

also with Staurastrum sp. in RuzrcKA, 1957, fig .

50. The taxonomic position relating to all these

plants needs a special study, as do the many

forms and varieties within Staurastrum sebaldi

group as a whole.

Staurastrum sebaldi var. ornatum NoRDST. fa.

Figs. 21 : 2 and 22 : 6. Length 56-60 fl, breadth

72.5-80 11-· From the pond near the military

camp. The semicells are cylindrical in the lower

part and form in this respect a transition to fa.

planctonica (Li:TTKEM. ) TElLING. The figure in

WEST & CARTER, 1 923, pl. 148, fig. 7, is not a

typical one, as much I can judge from my studies

on the Norwegian specimens (THOMASSON, 1957) .

The drawing given by NoRDSTEDT, 1873, is very

characteristic, though perhaps a little too ex

tended, whilst the prominent apical ornament

should be especially noted. A good figure is also

to be found in TElLING, 1 94 7. Cf. also the dis

cussion under Staurastrum manfeldtii.

Staurastrum spongiosum BREB.

Length of my specimen is 41 .3 fl· A good picture

of this species is given in IRENEE-MARIE, 1938, pl.

A cta Phylogeogr. Suec. 42

51 , fig. 8. There is some resemblance between the

present species and Staurastrum mayori G. S .

WEST, 1914, pl. 2 1 , fig. 32 , but the latter is

much smaller. Moreover it should be compared

with Staurastrum monticulosum var. bifurcatum

NoRDST. in CEDERGREN, 1932, pl. 4, fig. 50.

Staurastrum suborbiculare WEST & WEST fa.

Fig. 21 : 15 . Length 35-42.5 fl, breadth 30-35 fl·

Noted from the strand-pool on the. beach of

Lake Nahuel Huapi at Bariloche, and from the

submersion pool between Lake Gutierrez and

Lake Mascardi. The apex of my specimens is

slightly impressed.

Staurastrum subpolymorphum BoRGE fa.

Figs. 21 : 4, 6 and 23: 18. Length 31-34 ft, breadth

37 .5 11-· I have considered my plant as related

to the species described by BoRGE in 1903, pl.

107 , pl. 7, fig. 13. The dimensions of my plant

agree well with those given by BoRGE, but in

top view there are some differences. However,

considering the apical ornamentation in BoRGE's

figure one is somewhat doubtful about the

correctness of contour-lines. My specimens should

be compared also with Staurastrum distentum

WoLLE, e.g. in IRENEE-MARIE, 1951, pl. 2, fig. 3,

or in WEST & WEsT, 1897, fig. 6, though the

present species is somewhat smaller. There are

also superficial similarity to the plants of Stau

rastrum asteroideum and Staurastrum subgernu

latum var. gracilis.

Staurastrum trapezicum BOLDT.

Fig. 23: 1. Length 53 fl, breadth 79 fl · Occurring

among the filamentous algae in a brook at Lake

Mascardi. Compare BoLDT, 1888, pl. 2, fig. 46,

and MESSIKOMMER, 1927, pl. 2, fig. 20. This

species somewhat resembles Staurastrum pyra

midatum W. WEST which, however, has relatively

higher pyramidate semicells. There is also some

resemblance to Staurastrum brebissonii var.

truncatum GR6NBLAD in HoMFELD, 1929, pl. 8,

fig. 93.

Staurastrum sp. ...

Fig. 23: 17 . Length 22.5-25.5 fl, breadth 31 .3 11-· Lake Hess. Due to the only few specimens noted,


I have not been able to make an interpreta

tion of this plant. It should be compared with

Staurastrum oxyacanthum fa. in BoRGE, 1895 ,

fig. 16, and with some forms of Staurastrum

crenulatum var. britannicum MESSIK, fa. in

NYGAARD, 1 949.

Staurastrum sp.

Fig. 2 1 : 3 . Length 45 ,u, breadth 35.5 ,u. Lake Hess .

Probably this specimen belongs to the Stauras

trum sebaldi-group. There is some resemblance

to the forms grouped around var. productum and

var. altum, also B0RGESEN's figure of var.

brasiliense is quite similar. From all these varie

ties my specimen differs in its smaller size and

divergent apical ornamentation.

Staurastrum sp.

Fig. 23: 8. This specimen was noted in the plank

ton of Lake Guillelmo. It somewhat resembles in

shape the specimen designated as Staurastrum

manfeldtii, fig. 23: 7, from Lake Mascardi, but

differs in its rich ornamentation. There is also

some slight reminiscences to Staurastrum java

nicum (NoRDST. ) TURN. pictured in SKUJA, 1949,

pi. 34, fig. 2 1 , which is, however, larger. My plant

ought to be compared with some forms of Stau

rastrum cyclacanthum WEST & WEST . Of. also

Staurastrum borgeanum ScHMIDLE in THOMASSON,

1 957 .

Staurodesmus convergens (EHRENB. ) LrLLIER.

Fig. 2 1 : 17 . Length 40 ,u breadth without spines

77 ,u From the pond situated west of the military

camp. The figure shows a dichotypical specimen.

Compare pl. 13, fig. 1 1 of Arthrodesmus convergens

EHRENB. fa. in ScOTT & GRONBLAD, 1957.

Staurodesmus dejectus (BREB. ) .

Figs. 2 1 : 7 and 23: 24. Length 26.3 ,u, breadth

24.7 ,u. From the pond situated west of the

military camp.

Staurodesmus . glabrus fac. glabrus (EHRENB.) TEl

LING.

Fig. 2 1 : 18. Length 24 ,u, breadth with spines

32.5 ,u, without spines 24 ,u. Collected from the

submersion pool between the lakes Gutierrez

and Mascardi.

Staurodesmus megacanthus (LUND. ) THUNMARK fa.

Fig. 2 1 : 23. Length 31 ,u, breadth with spines

50 ,u, without spines 32 ,u. From the plankton

of Lake Hess. This taxon has also been met

in the lakes of Tierra del Fuego, and noted under

the name of Staurodesmus megacanthus, though

I am not quite satisfied about the identification

for there is great resemblances to some forms of

Staurodesmus dickiei, e.g. the plant figured in

TAYLOR, 1935, pl. 33, fig. 12, and also to some

forms of Staurodesmus convergens (EHRENB . )

KuTz.

Staurodesmus mucronatus var. subtriangulare (WEST

& WEST) n. comb.

Fig. 2 1 : 14. Length 33 ,u. Noted in the plankton of

Lake Hess. Compare WEST & CARTER, 1923, p1 .

130, fig. 13 & 14.

Staurodesmus phimus (TURNER) n. comb.

Fig. 23: 22. Cells 15 ,u long without spines, 26 ,u long with spines, breadth without spines 15 ,u, with spines 22.5 ,u. Rare in the plankton of Lake

Gutierrez . Compare TURNER, 1892, pl. 12, fig. 9 .

To this species belongs also Staurastrum curva

tum fa. biradiata KRIEGER, 1932, pl. 14, fig. 14.

It is the author's opinion that the specimens

figured in WEsT's Monograph represent a variety,

not the main species.

Staurodesmus triangularis var. convergens n. var.

Fig. 23: 23. Length 24 ,u, breadth without spines

22 ,u, breadth of isthmus 7 .5 ,u. Differs from the

main species through its short and converging

spines. Noted in a sample collected from the

pond situated west of the military camp in

Bariloche. The specimen of Arthrodesmus trian

gularis LAGERH. with converging spines figured

in B0RGESEN, 1890, pi. 4, fig. 41 , is probably better

to include to Staurodesmus cuspidatus. Likewise

the specimen with parallel spines in SKUJA, 1949,

pi. 33, fig. 4. On the other hand the alga pictured

as Staurastrum triangularis LAGERH. by CEDER

GREN, 1932, pl. 3, fig. 46, obviously represents

Staurodesmus subtriangularis var. inflatus (WEST

& WEsT) TElLING, likewise also Arthrodesmus

granulatus LARSEN, 1907, pi. 7, fig. 7 .



2. M I CR O Z O A

r agnicola spp.

Fig. 24: 18. The common planktic species. Usually

there is one animal, seldom two, in every lorica.

Fig. 24: 19. A benthic species occurring attached

on filamentous algae. Dimensions of lorica are:

height 152 fl, breadth 77 .[:, fl, breadth of the

mouth of lorica 102.5 fl· From a tarn in Paso

Vuriloche, v.s.

Cephalodella cf. sterea fa. mutata DoNNER.

Fig. 24: 9. Length of body 94 fl, length of toes

62 fl· Occurs in the submersion pool between

Lake Gutierrez and Lake Mascardi. This identi

fication is made with some doubt. The eyespot

has not been observed. The toes are twice as

long as in Cephalodella sterea. In this respect the

present specimen resembles somewhat Cephalo

della intuta MYERS which has no eyespot, but

the toes of Cephalodella intuta have a separating

wall in their distal end. · Probably the present

specimen from the Nahuel Huapi National Park

represents a new species, but studies of living · animals are necessary for the settling of this

question.

Colurella adriatica EHRENB .

Fig. 24: 13 . Length 94 fl, height 44 fl, Lake Hess.

Colurella hindenburgi STEINECKE.

Fig. 24: I . Length 67 .5 fl, height 41 fl, toes 24 fl· Pool on the beach of Lake Nahuel Huapi.

Colurella tesselata (GLASCOTT) .

Fig. 24: 2 . Length 60 p, , height 3 5 p, , toes 1 8 p,: Pool on the beach of Lake Nahuel Huapi.

Dicranophorus grand is (EHRENB . ) .

Fig. 24: 15. Length o f body 315 fl , breadth o f body

158 p,, toes 87 .5 fl· Submersion pool between Lake

Gutierrez and Lake Mascardi.

Euchlanis cf. meneta M YERS . ·

In some samples, collected in the Nahuel Huapi

area, e.g. Lake Hess, a submersion pool between

Lake Gutierrez and Lake Mascardi, a few speci

mens of an Euchlanis sp. were observed which

due to the state of contraction of the individuals

could not be identified with any certainity. They

seem to be related to Euchlanis meneta MYERS.

Length of the body is 150-160 fl, breadth 150-

155 p,, length of toes about 80 p,. The dorsal plate

is considerably wider than the ventral plate ,

under the posterior portion of the dorsal . pia te

there is a small cuticular shield-like process. The

toes are long and slender, and nearly parallel

sided. Probably my specimen ought to be

compared with Euchlanis cristata, described by

DADAY in 1 854 from South America, but the

drawing of the last mentioned species tells one

almost nothing.

Lecane brundinii n. sp.

Fig. 24: 7. Total length 85 jt, breadth of dorsal

plate 61 fl, breadth of ventral plate 50 fl· The

markings of the ventral plate are fairly prominent

and rather intricate. The dorsal plate is facetted.

The anterior dorsal and ventral margins are

coincident, no anterior spines are present. The

toes terminate in an acutely pointed recurved

claw, which has dorsally a distinct basal spicule.

This specimen was collected from the pond near

the military camp.

Lecane rhomboides (GossE) fa.

Fig. 24: 1 1 . Length only 64 fl, breadth 47 .5 fl, toe

Fig. 24: I. Colurella hindenburgi 9. Cephalodella cf. sterea fa. mutata 1 7. M icrocyclops anceps �

2. a. tessellata

3. Notholca caudata

4. M onostyla (Lecane) hamata

, 5. M ytilina ventralis

6. Monostyla (Lecane) hamata

7. Lecane brundinii

8. Scaridium longicaudum


1 0. Keratella cochlearis

1 1 . Lecane rhomboides fa.

1 2. Lepadella amphitropis

1 3 . Colurella adriatica

14. T·richocerca insignis

1 5. Dicranophorus grandis

1 6 . Lecane luna

1 8 . Vagnicola sp.

19. Vagnicola sp. 20. Boeckella schwabei �

2 1 . Boeckella titicaca, fifth legs of

� & d' 22. Boeckella gracilipes, fifth legs of

� & d'


A cta Pllytogeogr. Suec. 42


1 5 p,. A forma with robust dorsal wedge. Collected

from submersion pool between Lake Gutierrez

and Lake Mascardi.

Lepadella amphitropis HARRING.

Fig. 24: 12. Length lOO p,, breadth 57 .5 p,. Pool

on the beach of Lake Nahuel Huapi.

Monostyla (Lecane) cf. acanthinula HAUER.

Length 73 p,, breadth 61 p,, toe 30 p,. The identi-

fication is not conclusive.

Monostyla (Lecane) hamata STOKES.

Figs. 24: 4 and 24: 6. Length 90-91 p,, width of

dorsal plate 62.5 p,, width of ventral plate 50 p,, length of toe 32.5 fl· This species should be

compared with Monostyla 'descipiens MuRRAY and

Monostyla asinuata HAUER.

Mytilina ventralis (EHRENB. ) var.

Fig. 24: 5. A somewhat deformed specimen, prob

ably close to var. brevispina EHRENB.

N otholca caudata CARLIN.

Fig. 24: 3 . Length 200 p,, breadth 90 p,. Submersion

pool between Lake Gutierrez and Lake Mascardi.

Remarkable is the marked sculptural pattern

on the anterior dorsal margin. In this respect

our specimens resemble Argonotholca foliacea

(EHRENB . ) .

Scaridium longicaudum (0 . F. M.) .

Fig. 24: 8. Length o f body 106 p,, length o f the

second phalanx of foot 68 p,, length of toes 135 p,. Submersion · pool between Lake Gutierrez and

Lake Mascardi.

Trichocerca insignis (HERRIOK) .

Fig. 24: 14. Length 21 5-220.5 p,. Recorded in a

bottom sample from Lake Guillelmo, and in the

plankton of Lake Gutierrez. It is nearly related to

Trichocerca myersi (HAUER) which is, however,

somewhat smaller.

Boeckella gracilipes DADAY.

The distribution of Boeckellidae. round the Antarc

tic continent is one of the most fascinating prob

lems in the biogeography of the southern hemi-


sphere. The distribution of Boeckellfdae corre

sponds on the whole to the distribution of south

ern hemispheric plant genus Nothofagus, cf. Du

RIETZ, 1940. The latest survey of the distribu

tion of Boeckellidae in South America is given in

LOFFLER, 1955. One of the most widely spread

species is Boeckella gracilipes DADAY. Within

its area it seems to have differentiated into

several races which are morphologically separ

able. We have here obviously a "Formenkreis"

with many geographically isolated populations.

One of these populations has recently been

separated out by HARDING (1955) , cf. also

KIEFER, 1957. It is Boeckella titicaca HAR

DING occurring in the lakes of Altiplano, Peru.

It is the opinion of the present author that

in this case probably the application of trinary

nomenclature will be the most suitable method

for naming such taxa of Boeckella gracilipes. If so

the name for the animals from the lakes of Alti

plano would be Boeckella gracilipes titicacae

(HARDING). Also the population occurring in

Lake Nahuel Huapi is somewhat different from

DADAY's species. The fig. 24: 22 show the fifth

legg of the male and female specimens from Lake

N ahuel Huapi. The animals are somewhat differ

ent from the hitherto studied populations. There

fore I am of the opinion, that there are good rea

sons for regarding the specimens from Lake

Nahuel Huapi as representives of a taxon of its

own, and to designate this taxon preliminary as

Boeckella gracilipes nahuelhuapiensis. The most

conspicuous character is the extraordinary long

first segment of male left exopod. It is about

twice as long as the first segment of the male

right exopod, and it is also more slender than

in specimen described hitherto. The se·cond seg

ment of the fifth leg in the female is longer than

the first segment. In this respect my specimens

resemble more closely the animals described by

DADAY than Boeckella titicacae HARDING. For

comparison I have included fig. 24 : 21 represent

ing fifth legs from the specimens collected by me

from Lake Titicaca. When comparing this figure

with fig. 1 6 in HARDING (op. cit.) it is evident,

that the female fifth leg is subject to a consider

able variation, cf. also LOFFLER (op. cit.) , p . 745.

Fishes and waterfowl 79

It is probable that our large collections from

Chilean lakes are of great importance for the

taxonomy of Boeckella gracilipes-group. The

studies by Dr. L6FFLER on these collections will

certainly elucidate the taxonomy and phylogeny

of these animals. It is very plausible that owing

to the great variability of this species, the

distinguishing of minor taxa is not practicable.

X I I I . F I S H E S AN D WA T E R FOWL

Concluding m y notes from the Nahuel Huapi

National Park a few lines about the fishes and

waterfowl are of some interest. Both groups are

an integral part of the food-chain of the lakes and

streams. Fishes are wholly confined to water, but

when it comes to the birds it is often difficult to

decide which kinds qualify for inclusion. I have . considered here only the commonest of waterfowl

which have their source of food resources located

in the waters ·o f the National Park.

The lakes and rivers are well-stocked. Among

the native fishes one ought to mention first of all

Percichtys trucha Cuv. et V AL. and Patagonia

hatcheri Era ENM. , and the small fishes from the

genus Calaxias. For the delight of anglers numerous

foreign salmon-fishes have been introduced into the

waters of the Nahuel Huapi National Park. Due to

favourable conditions of life: clear and pure waters

of rather low temperature, ample supply of food and

lack of competition of any importance from native

fishes, these introduced fish thrive magnificently.

The lakes and streams of the National Park are

world famous for the excellence of their salmon

and trout and for their extraordinary sportiness. The

most important of the introduced fishes are Salmo

sebago GIRARD, Salmo irideus GIBBONS, Salmo fario

L. and Salvelinus fontinalis MrcH. Keeping up the

stock is attended to by a hatchery for salmonfishes

situated about 12 km from Bariloche.

There are many species of birds which are closely

connected with the productivity and food-chains of

the waters but here I have included only those

which feed extensively on the products of fresh

waters. Cormorants are represented by bigua cor

morant (Phalacrocorax o. olivaceus HuMBOLDT) and

magellanic blue-eyed shag (Phalacrocorax a. atri

ceps KING) ; the most common of the coots is the

white-winged coot (Fulica leucoptera VIEILLOT) . The

following ducks were noted: cinnamon teal (Anas c.

cyanoptera VIEILLOT), yellow-billed teal (Anas f.

flavirostris VIEILLOT), chilean widgeon (M areca

sibilatrix POEPPIG) , and flying steamer duck

( Tachyeres patachonicus KING) . In the Chilean lake

district the great grebe (Aechmophorus major

BoDDAERT) is frequent and occurs also on the lakes

in the N ational Park, reminding me in many respects

of the red-necked grebe. Moreover the southern

pied-billed grebe (Podilymbus podiceps antarcticus

LESSON), a species known also from the western side

of Cordillera, was met in the National Park. One

ought also to name the kelp gull (Larus dominica

nus LICHTENSTEIN) which was often noted. This is

a colony breeder.

Among mammals associated closely with fresh

water the nutria or coypu (M yocastor) feeds on

aquatic vegetation; in contrast is the Lutra provocax

THOMAS a carnivore. Both of them have a fur which

is highly prized by man.

The bottom fauna will be treated by Prof. L.

BRUNDIN, I note only the crayfishes. They are

represented in the National Park by the endemic

genus Aegla (Crustacea, Anomura) . These crayfishes

cove:red by a triangular and spinous carapax are

also common in the Chilean lake district. Moreover

from Lake Nahuel Huapi has been noted the

occurrence of Parastacus spinifrons (PHILIPPI) .

A cta Ph ytogeogr. Suec. 4-2

80 Acknowledgements

A C'K N OWLE D G E M E N T S

Concluding the present report on our journey

and studies in the National Park of Nahuel Huapi

we wish to express our most sincere gratitude to

all who have supported the Swedish Limnological

Expedition to South America 1953-54.

This expedition was made possible by means of

grants-in-aid from the Swedish Academy of Sciences,

the Swedish Natural Research Council, the Uni

versity of Uppsala, the Rockefeller Foundation,

and the Philosophical Society of Philadelphia. We

are most grateful for their assistance.

Swedish legations in Buenos Aires and in Santiago

de Chile have facilitated the work of th� expedition

in various ways. Their profund knowledge of the lab

yrinths of bureaucracy has been a great help to us.


My special thanks are due to Prof. LARS BRUN

DIN, my companion during the excursions in the

National Park of Nahuel Huapi for his good fel

lowship and for his never failing good spirits.

The working up of the material treated in the

. present paper has been carried out at the Vaxtbio

logiska Institutionen, Uppsala. To the Head of this

institute Prof. G. EINAR Du RrETZ I wish to

express my most sincere gr�titude for working

facilities and for his never failing ecouragement.

He has also read the manuscript and offered valu

able suggestions and criticisms, and determined my

collections of Euphrasia.

Vaxtbiologiska Institutionen,

Uppsala, March 15, 1958.

R E F E R E N C E S

Abbreviations according to �>World List of Scientific Periodicals�>, 3rd ed., 1 952.

AGOSTINI, A. M. DE, 1 949: Nahuel-Huapi. - Buenos Aires. AHLSTROM, E. H. , 1 943: A revision of the rotatorian genus Keratella, with descriptions of three new species

and five new varieties. - Bun. Amer. Mus. nat. Hist. , 80. A.LSTERBERG, G., 1935: Die Dynamik des Stoffwechsels der Seen im Sommer. - Lund. AUER, V. , 1958: The pleistocene of Fuego-Patagonia. 2. - Ann. Acad. Sci. fenn. , ser. A, Ill Geol.-geogr. , 50. BEETLE, A. A., 1 943: Phytogeography of Patagonia. - Bot. Rev., 9. BIRGE, E . A., 1 9 1 5: The heat budgets of American and European Lakes. - Trans. Wise. Acad; Sci. Arts Lett. ; 1 8 . BIRGE, E . A. , 1916 : The work o f the wind i n warming a lake. - Trans. Wise. Acad. Sci . Arts Lett., 1 8 . · BLACK, C. S . , 1 929: Chemical analyses of lake deposits. - Trans. V\"isc. Aacd. Sci. Arts Lett., 24. BLECHER, J. H. & FoGG, G. E., 1 955: Biochemical evidence of the affinities of Botryococcus. - New Phytol. , 54. BOELCKE, 0., 1957: Comunidades herbaceas del norte de Patagonia y sus relaciones con la .ganaderia. - Rev.

Inv. Agr., 1 1 . BoLDT, R., 1888: Desmidieer fran Gronland. - Bih. svenska Vetensk.-Akad. Handl. , Afd. 3 , 1 3 . BoRGE, 0 . , 1 895: Bidrag till kannedomen om Sveriges chlorophyllophyceer. 2. - Bih. svenska Vetensk.-Akad.

Handl. , Afd. 3, 2 1 . BoRGE, 0., 190 1 : Siisswasseralgen aus Siid-Patagonien. - Bih. svenska Vetensk.-Akad. Handl., Afd. 3 , 27 . BoRGE, 0. , 1 903: Die Algen der ersten Regnellschen Expedition. - Ark. Bot. , 1 . BoRGE, 0., 1 906: Siisswasser-Chlorophyceen von Feuerland und Isla Desolacion . - Bot. Stud. F. R. KJELLMAN.

Uppsala. BoRGE, 0 . , 1913 : Beitrage zur Algenflora von Schweden. 2. - Bot. Notiser, 1913 . BoRGE, 0. , 19 18 : Die von Dr. A. LoFGREN in Sao Paulo gesammelten Siisswasseralgen. - Ark. Bot. , 15 . BREHM, V. , 1 935: Mitteilungen von den Forschungsreisen Prof. RAHMS. 2. Gibt es in chilenischen Region Diapto-

miden? Diaptomus diabolicus nov. sp. - Zool . Anz., 1 12 . BRUNDIN, L . , 1 956: Die Bodenfaunistischen Seetypen und ihre Anwendbarkeit auf die Siidhalbkugel . Zugleich

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CABRERA, A. L. , 1 939: Las Compuestas del Parque Nacional del Nahuel Huapi. - Rev. Mus. La Plata, N.S. ,

Bot. , 2. CABRERA, A. L. , 1 953: Esquema fitogeografico de la Republica Argentina. - Rev. Mus. La Plata, N. S . , Bot. , 8. CABRERA, A. L., 1 954: Origen y evolucion de la flora del Parque Nacional Nahuel Huapi. - Natura, 1. CEDERGREN, G. R., 1 932: Die Algenflora der Provinz Harjedalen. - Ark. Bot . , 25. CHODAT, R. 1 926: Scenedesmus. Etude genetique de systematjque expedmEntale et d'hydrobiologie. - Z.

Hydrol. , 3 . CLEVE -EULER, ASTRID, 1 943 : List of diatoms from Lago Frey, with some critical remarks. - Bull. geol. Instn

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2. S. THUNl\:LABK, Der See Fiolen und seine Vegetation. vn + 198 s. (22 textfig.). 1931. Pris kr. 12: - (5: -).

3: 1. G. E. Du RIETZ, Life-forms of Terrestrial Flowering Plants. I. 95 s. (18 textfig.) ooh 10 pl. 1931. Pris kr. 8: - (4: -).

4. B. LINDQUIST, Om den vildvaxande skogsalmens raser och deras utbredning i Nordvasteuropa. (English Summary.) 56 s. ( 17 textfig.). 1932. Pris -kr. 6: (2: -).

5. H. OsvALD, Vegetation of the Pacific Coast Bogs of North America. 33 s. (5 textfig.) ooh 4 pl. 1933. Pris kr. 5: - (2: -).

6. G. SAMUELSSON, Die Verbreitung der hoheren Wasserpflanzen in Nordeuropa. 2 1 1 s. (50 kartor i texten). 1934. Pris kr. 12: - (7: -).

7. G. DEGELIUS, Das ozeanische Element der Strauchund Laubflechtenflora von Skandinavien. XII + 411 s. (88 textfig. varav 78 kartor) + 2 utvikn.-kartor och 4 pl. 1935. Pris kr. 15: - (8: -) .

8. R. SEBNANDER, Granskar och Fiby urskog. En studie over stormluckornas och marbuskamas betydelse i den svenska granskogens regeneration. (English Summary.) 232 s. (87 textfig.) och 2 pi. 1936. Pris kr. 12: - (6: -). . -

9. R. STERNER, Flora der Insel Oland. Die Areale der Gefasspflanzen Olands nebst Bemerkungen zu ihrer Oekologie und Soziologie. 169 s. (8 kartor i texten) + 64 s. med 288 kartor. 1938. Pris kr. 12: - (7: -).

10. B. LINDQUIST, Dalby Soderskog. En skansk lovskog i fomtid och nutid. (Deutsche Zusammenf.) 273 s. (99 textfig. varav 70 kartor). 1938. Pris kr. 12: -(7: -).

l l . N. ST.!i.BERG, Lake Vattern. Outlines of its Natural History, especially its Vegetation. 52 s. (2 kartor i texten) och 8 pi. 1939. Pris kr. 5: - (2: -) .

12. G . E. D u RIETZ, A. G . HANNERZ, G . LoH.AMMAB, R . SANTESSON und M. W&RN, Zur Kenntnis der Vegetation des Sees Takern. 65 s. (4 textfig. varav 2 kartor) och 7 pi. 1939. Pris kr. 5: - (2: -).

13. Vaxtgeografiska Studier tillagnade Carl Skottsberg pit sextioarsdagen 1/12 1940. x + 296 s. (49 textfig. varav 26 kartor) + 1 portratt och 30 pl., darav tva i farg. 1940. Pris kr. 15: - (8: -), for numrerade upplagan ( 150 ex.) kr. 30: - (25: -) . Contents:

AHLNER, S., Alectoria altaica und ihre V erbreitung in Fennoskandia.

ALBERTSON, N., Soorpidium turgesoens. En senglacial relikt i nordisk alvarvegetation.

ANREP-NORDIN, B., Till Carl Skottsberg.

ARNBORG, T., Der Vallsjo-Wald, ein nordschwedischer Urwald. (27 S., 10 Taf.)

CHRISTOPHERSEN, E., Ranunculus Caroli n. sp. Tristan da Cunha.

DAHLBEOK, N., Arenaria humifusa och skyddet av sallsynta vaxter.

Du RIETZ, G. E., Problems of Bipolar Plant Distribution. (68 pp. with 13 maps and 10 pp. bibliography.)

HoLMBoE, J . , Osmunda regalia. N orge. H.!.YBEN, E., Meeresalgen� ID.sel Hogland. voN KBusENSTJEBNA, E., Mossamhallen och moss

arter i Vasterbotten. LINDQUIST, B., Juncus alpinus var. Marshallii.

Scotland. Race Differentiation in the Species J. alpinus.

NoRDH.AGEN, R., Cladium mariscus. Norge. OsvALD, H., Sphagnum flavicomans. Taxonomy,

Distribution, and Ecology. PETTERSSON, BENGT, Orchis Spitzelii var. gotlan

dica n. var. PETTERSSON, BRoR, A Case of Long Distance Dis-

persal of Plants through the Import of Timber. SANDBERG, G., Gasteromycetstudier. SANTESSON, R., Valdiviansk regnskog. SELANDER, S., Till Carl Skottsberg. SEBNANDER, R., Prunus spinosa x P. insititia och

Prunus-artemas vaxtgeografiska stallning i den svenska vaxtvarlden.

SMITH, H., Carex arctogena n. sp. . W llllRN, M., Cladophora pygmaea und Leptonema

luoifugum an der schwed. Westkiiste. 14. N. HYLANDER, De svenska formerna av Mentha gen

tilis L. coll. (Deutsche Zusammenf.) 49 s. (8 textfig. ) och 4 pi. 1941. Pris kr. 4: - (2: -) .

15. T. E. HAssELBOT, Till kannedomen om nagra nordiska umbilicariaceers utbredning. (Deutsche Zusammenf.) 75 s. (6 textfig. varav 4 kartor) och 4 pl. 1941. Pris kr. 6: - (2: 50).

16. G. SAMUELSSON, Die Verbreitung der AlohemillaArten aus der Vulgaris-Gruppe in Nordeuropa. l59 s. (24 kartor). 1943. Pris kr. 10: - (6: -).

17 TH. ARWIDSSON, Studien iiber die Gefasspflanzen in den Hochgebirgen der Pite La;ppmark. 274 s. (53 textfig. varav 24 kartor) och I + 16 pl. 1943. Pris kr. 12: - (7: -).

18. N. DAHLBECK, Strandwiesen am siidostlichen Oresund. (English Summary.) 168 s. (29 textfig.). 1945. Pris kr. 10: - (6: -).

19. E. VON KBUSENSTJERNA, Bladmossvegetation och bladmossflora i Uppsalatrakten. (English Summary.) IV + 250 s. ( 13 textfig. varav 9 ka.rtor) och I + 4 pl. samt 1 utvikn.-karta. 1945. Pris kr. 12: - (7: -).

ACTA PHYTOGEOGRAPHICA SUECICA 20. N. ALBERTSON, Osterplana hed. Ett alvaromrade pa

Kinnekulle. (Deutsche Zusammenf.) xn + 267 s. (18 textfig. varav 13 kartor) + 16 pi. 1946. Pris kr. 14: - (8: -).

2 1 . H. SJORs, Myrvegetation i Bergslagen. (English Sum-· mary: Mire Vegetation in Bergslagen, Sweden. ) 300 s. (59 textfig. varav 16 kartor) + 16 s. tab. + 24 s. med 1 1 7 kartor + 32 pi. och 2 utvikn.-kartor. 1948. Pris kr. 15: - (8: -).

22. S. AHLNER, Utbredningstyper bland nordiska barrtradslavar. (Deutsche Zusammenf.: Verbreitungstypen unter fennoskandischen Nadelbaumflechten.)

· IX.+ 257 s. (22 textfig. varav 13 kartor) + 13 utvikn.kartor och 16 pl. 1948. Pris kr. 14: - (7: -).

23. E. JULIN, Vessers udde. Mark och vegetation i en igenvaxande lovang vid Bjarka-Saby. (Deutsche Zusammenf.: Vessers udde. Boden und Vegetation in einer verwachsenden Laubwiese bei Bjarka-Saby in Ostergotland, Siidschweden.) xv + 186 s. (73 textfig.) + 40 s. med 65 tab. + 48 s. med 80 kartor + 1 6 pl. 1948. Pris kr. 15: - (8: -) .

24. M. FRIEs, Den nordiska utbredningen av Lactuca alpina, Aconitum septentrionale, Ranunculus platanifolius och Polygonatum verticillatum. (Deutsche Zusammenf. : Die nordische Verbreitung von Lactuoa alpina . • . ) 80 s. (15 textfig.) + 1 pl. + 5 utvikn.-kartor. 1949. Pris kr. 7: - (4: -) .

25. 0. GJ&REVOLL, Sneleievegetasjonen i Oviksfjellene. (English Summary: The Snow-Bed Vegetation of Mts Oviksfjallen, Jamtland, Sweden.) 106 s. ( 17 textfig. + 14 tab. + 24 kartor och diagr.). 1949. Pris kr. 6: -(4: -).

26. H. OsvALD, Notes on the Vegetation of British and Irish Mosses. 62 pp., 13 Figs, IO Tables, and 16 Plates. 1949. Pris kr. 8: - (4: -).

27. S. SELANDER, Floristio Phytogeography of SouthWestern Lule Lappmark (Swedish Lapland) I. 200 pp., 33 Figs (12 maps), and 12 Plates. 1950. Pris kr. 18: ( 12: -) .

28. S. SELANDER, Floristio Phytogeography of SouthW astern Lule Lappmark (Swedish Lapland) II. Karlvaxtfloran i sydvastra Lule Lappmark. (English Summary.) 154 s. (6 textfig.) + 51 s. med 302 kartor. 1950. Pris kr. 18: - (12: -).

29. M. FRIES, Pollenanalytiska vittnesbtird om senkvartir vegetationsutveckling, sarskilt skogshistoria, i nordvastra Gotaland. (Deutsche Zusammenf.: Pollenanalytische Zeugnisse der spatquartaren Vegetationsentwicklnng, hauptsachlich der Waldgeschichte, im nordwestlichen Gotaland, Siidschweden. 85 s.) 220 s. { 43 text-fig.) + 1 + I-VIII pi. (kartor och diagram). 1951. Pris kr. 18: - ( 12: -).

30. M. W&RN, Rocky-Shore Algae in the Oregrund Archi. · pelago. XVI + 298 pp., 106 Figs, and 32 Plates. 1952. Pris kr. 25: - ( 16: -).

31. 0. RuNE, Plant Life on Serpentines and Related Rocks in the North of Sweden. 139 pp., 54 Figs, and 8 Plates. 1953. Pris kr. 18: - (12: -).

32. P. KAARET, Wasservegetation der Seen Orllmgen und Trehorningen. 66 s. (5 Fig., 3 Karten, 8 Veg.-profile) + 16 Tafeln (32 Fig.). 1953. Pris kr. 1'2: - (8: -).

33. T. E. fussELROT, Nordliga lavar i Syd- och Mellansverige. (Nordliche · Flechten in Siid- und Mittelschweden.) VIII + 200 s. (2 kartor) + 30 s. med 29 kartor. 1953. Pris kr. 22: - ( 14: -).

34. H. SJoRs, Slatterangar i Grangarde finnmark. (English Summary : Meadows in Grangarde Finnmark, SW. Dalarna, Sweden.) 4 pi. + 135 s. ( 57 fig., 14 tab.). 1954. Pris kr. 18: - (12: -).

35. S. KILANDER, Karlvaxternas ovre granser pa fjall i sydvastra Jamtland samt angransande delar av Harjedalen och Norge. [English Summary: Upper Limits of V asctilar Plants on Mountains in Southwestern Jamtland and Adjacent Parts of Harjedalen (Sweden) and Norway.] 200 s. (41 fig., 1 1 tab.) + 1 utvikn.-blad med 61 diagram. 1955, Pris kr. 22: - (14: -).

36. N. QUENNERSTEDT, Diatomeerna i Langans sjovegetation. (English Summary: Diatoms in the Lake Vegetation of the Langan Drainage Area, Jii.mtland, Sweden.) 208 s. (39 fig., 15 tab.) + 1 utvikn.-blad med diagram. 1 955. Pris kr. 22: - ( 14: -).

37. M.-B. FLORIN, Plankton of Fresh and Brackish Waters in the Sodertalje Area. 144 pp. (35 Figs and 35 Tables) and 1 (coloured) + 20 Pla�es. 1957. Pris kr. 20: - (14:-).

38. M.-B. FLoRIN, Insjostudier i Mellansverige. Mikro. vegetation och pollenregn i vikar av Ostersjobackenet

och insjoar fran preboreal tid till nutid. (Summary : Lake Studies in Central Sweden. Microvegetation and Pollen Rain in Inlets of the Baltic Basin and in Lakes from Pre-boreal Time to the Present Day.) 29 s. (10 fig., 1 tab.). 1957. Pris kr. 10: - (6:-).

39. M. FRIES, V egetationsutveckling och odlingshistoria i V arnhemstrakten. En pollenanalytisk undersokning i Vastergotland. [Zusammenf. : Vegetationsentwicklung und Siedlungsgeschichte im Gebiet von Varnhem. Eine pollenanalytische U ntersuchung aus Vastergotland (Sti.dschweden).] 64 s. ( 18 fig.) inkl. Abstract + I-V pl. {diagram). 1958. Pris kr. 16: - ( 10: -).

40. BENGT PETTERSSON, Dynamik och konstans i Gotlands flora och vegetation. (Zusammenf.: Dynamik und Konstanz in der Flora und Vegetation von Gotland, Schwaden.) 288 s. ( 124 fig.) + PI. I-XVI (kartor) + PI. XVII (kolor.). 1958. Pris kr. 40: -.

41. E. UGGLA, Skogsbrandfa.It i Muddus nationalpark. (Summary : Forest Fire Areas in Muddus National Park, Northern Sweden.) 104 sid. (8 pi., 66 fig. , 16 tab.) + 12 sid. med 22 kartor. 1958. Pris kr. 2 1 : -.

42. K. THOMASSON, Nahuel Huapi. Plankton of some Lakes in an Argentine National Park with Notes on Terrestrial Vegetation. 83 pp. (24 Figs. ) + 16 PI. 1959. Pris kr. 21: -.

RABATTE R . • D I S C OUNT • E RM A S S I G U N G E N · RABAI S

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Printed in Sweden, April 1959

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