214
CONTRIBUTIONS TO THE ECOLOGY OF SPITSBERGEN
AND BEAR ISLANDi
BY V. S. SUMMERHAYES AND C. S. ELTON.
and sevenFiguresin theText.)
(WithPlates I-1V
CONTENTS.
I.
INTRODUCTION
[I.
BEAR ISLAND
..216
Introduction
.
(a) Land Communities
(b) FreshwaterCommunities .
Food and Enemies ("NitrogenCycle")
III.
SPITSBERGEN
PAGE
..214
..216
.
.
.
.
.218
.228
.
231
..233
..233
Introduction
..236
Special Regions
.
.
A. PrinceCharlesForeland
B. Cape Boheman Region.
(a) Land Communities.
(b) FreshwaterCommunities.255
.
C. Klaas Billen Bay (Bruce City)
(a) Land Communities .259
(b) Aquatic Communities.264
D. Gips Valley .274
.
.
E. Temple Bay (Sassendale)
.
.
F. Lesser EdinburghIsland
.
.
G. HermansenIsland
..282
H. Advent Bay
IV.
SUMMARY AND CoNCLUsIoNs
V.
BIBLIOGRAPHY
.
.
.236
.
.
249
250
258
.276
.280
.281
.282
..284
I. INTRODUCTION.
The account given below is the resultof investigationscarriedout by us
while membersof the OxfordUniversityExpedition to Spitsbergenduring
the summerof 1921. A fairlycompleteidea of the biologywas obtained,as
duringthe restof the year nearlyall the plants and animals remaindormant
underthicksnow,or have migratedsouthwards.
Owingto the varied requirementsof the membersof the Expedition,it
was in many cases impossibleto completethe workon the areas visited. In
some cases a visit of a fewhoursonly was possible. Ten days werespent on
each of two areas-on Bear Island (June 13th-23rd)and on Prince Charles
I
"Results of the OxfordUniversityExpeditionto Spitsbergen,1921." No. 29.
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V. S. SUMMERHAYESAND C.
S. ELTON
215
Foreland, Spitsbergen(June 30th-July10th)-while one of us stayed at the
head of Klaas Billen Bay fromJuly19thto August16th.
As the Rev. F. C. R. Jourdainis publishingin the futurea manual on the
birdsof Spitsbergen,detailsofthisgroupare onlygivenin so faras theymake
clearthe animal communities.The materialcollectedwas identifiedby various
specialists,and we followtheirnomenclaturein most instances. We should
like to expressour thanksto themfor identifyingmembersof the following
Mr A. J. Wilmott;Mosses,
groups: PLANTS:Phanerogamsand Pteridophytes,
Mr H. N. Dixon; Liverworts,Dr W. Watson; Lichens,MrR. Paulson; Marine
Tunicata
ANIMALS:
Algae,MrA. Gepp; and FreshwaterAlgae,MrB. M. Griffiths.
and Polyzoa, Mr R. Kirkpatrick;Mollusca (and much otherhelp), Mr G. C.
Robson; Spiders,Dr A. R. Jackson; Land mites,Rev. J. E. Hull; Water
mites,Mr C. D. Soar; Diptera (Nematocera),Mr F. W. Edwards; Diptera
etc.),MrJ. E. Collin;Hymenoptera(Sawflies),MrF. D. Morice;
(Orthorrhapha,
Parasitic Hymenopteraand Mallophaga, Rev. J. Waterston; Fleas, Mr K.
Jordan; Collembola,Dr G. H. Carpenterand Miss K. C. Joyce Phillips;
Mysidacea,Dr W. M. Tattersall; help with Amphipoda,Dr W. T. Calman;
Leeches, Dr H. A. Baylis; Polychaeta,Mons. P. Fauvel; Oligochaeta,Dr J.
Stephenson;Rotifera,Mr D. Bryce; Hydroida,Mr K. Totten; and Protozoa
and Bacteria,Mr H. Sandon.
We also thankMr J. F. Manleyfortwo wateranalysesand Mr H. C. Mills
forthe chloridedeterminations.We are deeply indebtedto ProfessorE. B.
Poulton and to ProfessorF. W. Oliver formuch help in gettingthe animal
and in otherways. We shouldliketo express
and plantcollectionsdetermined,
our gratitudeto the membersof the Expedition,whose help is much appreciated. In particularwe owe our thanksto the Rev. F. C. R. Jourdain(32),
Dr T. G. Longstafi,MessrsJ. S. Huxley, A. M. Carr-Saundersand J. Walton.
We are indebtedto Mr J. Mathiesonofthe ScottishSpitsbergenSyndicatefor
permissionto reproducethe map of Prince CharlesForeland,and to Messrs
A. M. Carr-Saundersand J. Walton forthe use of various photographs.
Althougha greatmanypapershave been publishedon theFlora and Fauna
and practically
of the region,yet fewhave dealt withthe plant communities,
nonewiththoseofthe animals. Ofthe latterwe have foundOlofsson'spapers
communities.The first
(47, 48) a greathelp in connectionwiththe freshwater
and almost the only importantwork on the plant communitiesis that of
Nathorst(44) who divides the countryinto (1) Strand, (2) Bogs, (3) Slopes,
and pointsout the mainplantsin each. Otherwritershave noticedprominent
suchas "slopes underbirdcliffs,"etc.,or have dealt withgeneral
communities
lifeconditionsin the Arctic.
The seasonal successionin the animals is not so markedas in temperate
countries,and does not lead to manyerrorsin collecting,the only exceptions
beingthe Diptera and Hymenoptera.A shortvisit to a locality,if the ground
is worked carefully,gives a true idea of most of the animal communities.
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216 Contributions
to Ecology of Spitsbergen
and Bear Island
Owingto thelabourofcollectinganimalstypicalareas wereselected,and these
wereworkedout in detail.
Detailed experimentalwork was only possible at Klaas Billen Bay, and
the divisioninto communitiestherefore
restsusually on moreeasily obtained
superficial
evidence. The accountis in no way to be consideredan attemptat
a, finalclassificationof communitiesin the region.
II. BEAR ISLAND.
INTRODUCTION.
We propose to deal with Bear Island firstbecause it is much smaller,
has fewerspecies of plants and animals,and is thereforesimplerthan Spitsbergen. Also, in some ways, it is transitionalbetween Arctic Europe and
Spitsbergen.
Bear Island lies 240 miles northof Scandinavia, on the westernedge of
the shallowsubmarinebank of the BarentsSea, whileto the westis the deep
GreenlandSea. The island (Fig. 1) consistsof two distinctportions-a flat
northernarea of sedimentaryrocks,and a southernmountainouspart composed mostlyof the faulted and metamorphosedlayers of the Hecla-Hook
system. MountMisery(1800 feet)is the highestpoint on the island.
Although120 miles south of Spitsbergenthe climateof Bear Island is in
manyrespectssomewhatmoresevere. The mean temperaturein Februaryis
-
12.00 C. while that of August is
4.50
C. The Gulf Stream drift here meets
the cold polar currentfromeasternSpitsbergen,producingmany fogswhich
give the island a bad reputationamong seamen. This fogginessaffectsthe
plant lifeespecially,since it reducesthe amount of directsunlightavailable.
This is ofmuchmoreimportancein theArcticthanis themeanair temperature
(68, etc.).
Anotherimportantfactoraffecting
lifeis thefrequencyofstormsin winter.
The resultof these is the completeremovalof the snow frommany places,
and theirexposureto the low temperaturesthen occurring.This is well seen
in the extremefrostweatheringvisiblein the mountainouspart of the island.
The sea does not usually freezein winter,but at timesthereis much drifted
ice fromthe northaroundthe island and this adverselyaffectsthe climate.
We wereonlyable to studythe regionsouthofthe brokenline on the map
(Fig. 1). Thisarea consistsofveryvariedrocks-limestones,slates,sandstones,
etc., but the vegetation,on the whole,seems to be the same on the different
rocks. The followingclassificationof communitieswas arrivedat:
(a) Land Communities.
1. Bird Cliffs.
2. "Fjaeldmark" as definedby Warming(68).
3. Herb-mat(with "Skua hummocks").
4. Moss Heath.
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216 Contributions
to Ecology of Spitsbergen
and Bear Island
Owingto thelabourofcollectinganimalstypicalareas wereselected,and these
wereworkedout in detail.
Detailed experimentalwork was only possible at Klaas Billen Bay, and
the divisioninto communitiestherefore
restsusually on moreeasily obtained
superficial
evidence. The accountis in no way to be consideredan attemptat
a, finalclassificationof communitiesin the region.
II. BEAR ISLAND.
INTRODUCTION.
We propose to deal with Bear Island firstbecause it is much smaller,
has fewerspecies of plants and animals,and is thereforesimplerthan Spitsbergen. Also, in some ways, it is transitionalbetween Arctic Europe and
Spitsbergen.
Bear Island lies 240 miles northof Scandinavia, on the westernedge of
the shallowsubmarinebank of the BarentsSea, whileto the westis the deep
GreenlandSea. The island (Fig. 1) consistsof two distinctportions-a flat
northernarea of sedimentaryrocks,and a southernmountainouspart composed mostlyof the faulted and metamorphosedlayers of the Hecla-Hook
system. MountMisery(1800 feet)is the highestpoint on the island.
Although120 miles south of Spitsbergenthe climateof Bear Island is in
manyrespectssomewhatmoresevere. The mean temperaturein Februaryis
-
12.00 C. while that of August is
4.50
C. The Gulf Stream drift here meets
the cold polar currentfromeasternSpitsbergen,producingmany fogswhich
give the island a bad reputationamong seamen. This fogginessaffectsthe
plant lifeespecially,since it reducesthe amount of directsunlightavailable.
This is ofmuchmoreimportancein theArcticthanis themeanair temperature
(68, etc.).
Anotherimportantfactoraffecting
lifeis thefrequencyofstormsin winter.
The resultof these is the completeremovalof the snow frommany places,
and theirexposureto the low temperaturesthen occurring.This is well seen
in the extremefrostweatheringvisiblein the mountainouspart of the island.
The sea does not usually freezein winter,but at timesthereis much drifted
ice fromthe northaroundthe island and this adverselyaffectsthe climate.
We wereonlyable to studythe regionsouthofthe brokenline on the map
(Fig. 1). Thisarea consistsofveryvariedrocks-limestones,slates,sandstones,
etc., but the vegetation,on the whole,seems to be the same on the different
rocks. The followingclassificationof communitieswas arrivedat:
(a) Land Communities.
1. Bird Cliffs.
2. "Fjaeldmark" as definedby Warming(68).
3. Herb-mat(with "Skua hummocks").
4. Moss Heath.
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V. S. SUMMERHAYESAND(J. S. ELTON
lXoJSs
H
>
LOW FLAT LAN>
In
C....-;.J.
<~~~~~~~~~~~~~~~~~~~.
CAPE
~ ~ ~ ~
...L
KCAPR
o
217
m
~ ~~~~
COAL
o-;-
C>
-
o~~~~~~~~~~~'
FIG. 1.
notcontours.
Map ofBearIsland. Dottedlinesareform-lines,
---S-Boundary ofareainvestigated.
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and Bear Island
to Ecology of Spitsbergen
218 Contributions
5. Rock and Boulder Community.
(a) Rock surfaces.
(b) Crevices.
6. Moss-mat.
7. Wet Tundra.
(a) Pond marginalcommunity.
(b) Streammarginalcommunity.
(c) Moss-bog.
(b) FreshwaterCommunities.
1. Still water.
(a) Lakes.
(b) Ponds.
2. Runningwater.
that all transitionsoccurbetweenthesegroupings.
It mustbe remembered
(a) Land Communities.
1. BIRD CLIFFS.
High, almost perpendicularcliffs,sometimesseveral hundredfeet,occur
all along the southerncoast. They are inhabitedby vast coloniesofsea birds.
Thereare threekindsof habitatswhichare not always clearlyseparable.
(a) Ledges on the sides ofthe cliffs.The followingbirdsnestherein large
numbers:
foreach speciesis givenwhenthatspeciesis mentioned
forthefirsttime.
(The authority
a., abundant;
c.d.,co-dominant;
d.,dominant;
areusedthroughout
as follows:
symbols
Frequency
butfrequency
notdetermined.)
o., occasional;r.,rare;1.,local; + present,
f.,frequent;
a.
Common
Guillemot
(UriatroilletroilleL.)
,,
a.
(U. troillevar.ringviaBrin.)
Ringed
a.
Briinnich's ,,
(U. lomvialomviaL.)
,,
(U. gryllemandtii
o.
Mandt's
Mandt)
a.
Kittiwake
(Rissatridactyla
tridactyla
L.)
a.
FulmarPetrel
(Fulmarus
glacialisglacialisL.)
Gieb. occurredon the Fulmar.
The bird-louseEsthiopterusnigrolimbatus
In a fewplaces thesebirdswerenestingon the cliffsofinlandvalleys. All these
speciesfeedat sea.
(b) In crevicesof the rock the Little Auk (Plotus alle L.) and the NorwegianPuffin(FraterculaarcticaarcticaL.) occurin small numbers.The former
also nestsin the screesbelow. These two speciesfeedalso at sea.
(c) At the tops of the cliffson the mainland,and on small skerries,one
Gunn.)and the GreatBlack-backed
findsthe GlaucousGull (Larus hyperboreus
Gull (L. marinusL.). The formerare abundant; of the latteronlya fewpairs
upon which
wereseen. The Glaucous Gull usually nestsabove the cliff-birds
it preys,eatingthe eggs and youngof such birdsas Guillemots. It will also
devour anythingin the nature of carrionor garbage. This gull is powerful
enough to defendits nest against the Arctic Fox (Vulpes lagopus) which
explainsits choice of nestingplace. The Fulmar also sometimesnests in the
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V. S. SUMMERHAYES AND C.
S. ELTON
219
same places; this is probably due to the presentscarcityof foxes. A few
NorthernEider Ducks (SomateriamollissimaborealisBrehm.)probablybreed
in this kind of place.
Kittiwakes use mosses chieflyfor building their nests, but also algae,
lichens,grasses,Salix, Cerastiumalpinum,Saxifraga opposittfolia,
etc. (62).
The GlaucousGullalso uses moss. Thereis quite a variedfaunain thesenests:
viaticusTullb.
Collembola:Achorutes
armatusTulib.var.aroticus
Tullb.
Onychiurus
0. neglectus
Schiaf.
Tardigrades:
Echiniscus
testridi
Doyere
Ehrb.
E. arctomys
Macrobiotes
hufelandi
C. Schultze
M. ornatusRichters
Leriaseptentrionalis
Collin
Diptera:
The Collembola were found by Wahlgren(65) in Glaucous Gull nests.
The Tardigradesare someofthe commonmossspecies. The flyLeria was found
both as pupae and adults in a Glaucous Gull's nest, and has been found
elsewhereon St Kilda only. This suggeststhat the flymay be associated with
gulls. The warmthof the sittingbirdswould speed up the flies'development.
Much bird dung is droppedon the cliffs,whichenables the plants present
to growveryluxuriantly.On the ledges around the nestsare large plants of
L. togetherwith much grass(18). Cochleariais indeed
Cochleariaofficinalis
a well-knownplant in such places, and has been recordedfromnearlyall
arctic and sub-arcticcountries. It also occurs in the BritishIsles at Great
Orme's Head, Abbotsbury,etc., where droppingsaccumulate. The plants
observedon the cliffsnear Mount Miserywere 7 to 8 ins. in heightand very
bushy, while on the flat "fjaeldmark" above the cliffthey were adpressed
closelyto the groundand about 1 inch in diameter.
2.
"FJAELDMARK."
Most ofthe regioncan be includedunderthisheading. It consistsofareas
on whichthe vegetationis open. As a resultofseverefrostweatheringin this
districtthe groundis coveredwithrock detritus,mostlyof small size, which
variesin shape, etc., withthe type of the parentrock.
The plants are almost all herbaceous,the only dwarf-shrubs
being the
threeDwarf Willows(Salix polaris Wahl., S. herbaceaL. and S. reticulataL.).
The firstis the only one of these at all common. This "Herb fjaeldmark"
passes over in favourableplaces into "Herb-" or "Grass-mat" and not into
" Dwarf-shrub
Heath" as in Greenland(28, 67). The plantsare almosteverywhereverysmalland stunted,beingrarelyover2 or 3 ins. high. Manyofthem
are cushionforms(e.g. Papaver, Saxifraga caespitosa),or else they formlow
creepingmats (S. oppositifolia).
The amount of animal lifeis verysmall,as is also the numberof species.
The local distributionof the invertebratesseems to depend a great deal on
chance. The intenseweatheringmust be constantlyexposingfreshsurfaces
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and(Bear-I4las d
220 Contribatiousto Ecology of Spitsbergen
emptyhabitats,whichare occupiedbythefirstcomers.As with
and forming
thelargest
thingis thesmallsizeoftheinvertebrates,
theplants,onestriking
long.Another
whichis lessthanhalf-an-inch
Arnauronematus,
beingthesawfly
or moths,
butterflies
ofmolluscs,
absenice
featureofthefaunais thecomplete
beetles,ants,beesor wasps,etc.
andwiththistheanimals,variesconsiderably,
Thedegreeofplantcovering,
On theridges
froinbaregroundto the" herb-mat."
therebeingall gradations
of the hills,especiallyin the area of the TetradiumLimestone the oldest
oftheHecla-Hooksystem-arelargeareasofrubblewhicharealmost
stratum
to 2 ins.in diameter
are usuallyfromone-half
devoidofsoil. Thefragments
similarto scree.Herelifeis verysparse.
a veryunstablesubstratum
forming
L. anid
The onlyhigherplantsare isolatedtuftsof Saxifragaoppositifolia
few
lichens
The
few
crustaceous
occur.
L., whilehereand therea
S. caespitosa
understones(e.g. Lauterborina
animalslivinghereare somefliessheltering
lineatus
urstnusHolmg.),a mite(Scutovertex
caracinaZett.,Metrtocnemus
on the undersidesof flatstones,and the spiderCoryThor.)congregated
Thor.
phaeusholmgrenii
in similarplaces,probablyowingto lower
On the slatesand sandstones
are
is a littlericher.Here,besidesthe Saxifrages,
altitude,the vegetation
The
polaris.
anid
Salix
Rottb.,
radicatum
L.
var.
nudicaule
Papaver
found
be
to
lichens,
is morestable,witha littlesoil,andso therearenumerous
substratum
commonest:
are the
on thesmallstones.Thefollowing
crustaceous,
mnainly
Acarosporasp.
AlectoriabicolorNyl.
A. nigricansNyl.
GyrophorahyperboreaAch.
G. proboscideaAch.
Lecidea confluensAch.
L. pantherinaTh.Fr.
RhizocarpongeographicumDC.
VerrucarianigrescensPers.
f.
lessscantyfaunathanthebareareas. Thefollowing
hasa rather
Thisregion
werecollectedon the flatshatytop of a hillamongsttheplantsmentioned
above:
Collembola: Onychiurusarmatusvar. arcticus
Diptera:
SQiarasp.
Coryphaeusholmgrenii
Spiders:
Mites:
L.Koch
Cyta brevirostris
Rhagidia gelida Thor.
One or two otherspecies
the typical"fjaeldmark"on
A largepartofthe area, whichconstitutes
a numberof
thanthatalreadydescribed,
BearIsland,has a richervegetation
mossesandlichensbeingcommon.In placesSalixpolarisforms
phanerogams,
no grasseson thisarea. The folquitelargepatches.Thereare practically
lowingis a generallistofplantsforthetypical"fjaeldmark":
Phanerogams,etc.:
Arabis alpina L.
CerastiumalpinumL.
Cochleariaofficinalis
Draba alpina L.
D. hirtaL. var. aretica Vahl.
Equisetum variegatumSchl.
Hill
Oxyriadigyfna
var. radicatuni
Papaver nu(dicaiule
RanunculuspygmaeusL.
r.
f.
1.
o.
r.
I'.
f.
f.
1.
Salix herbacea
S. polaris
S. reticulata
Saxifragacaespitosa
S. cernuaL.
S. nivalis L.
S. oppositifolia
rhodiolaDC.
Sedumn
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1.
a.
o.
a.
1.
o.
a.
1.
V. S. SUMIERHAYES AND C.
S.
ELTON
221
Bryophytes:
BryumglobosumLindb.
*HypnumuncinatumHedw.
B. pseudo-triquetrum
Schw.
Lophozia floerkiiSchiffn.
var. cavifoliumBerggr.
PolytrichumjuniperinumWilid.
*CynodontiumvirensHedw.
Preissia commutataNees.
var. arcticumBerggr.
Timmia austriaca Hedw.
DicranoweisiacrispulaHedw.
var. atrata Schimp.
Lichens:
Cladonia rangiferina
Web.
Peltigeracanina Willd.
*CetrariaislandicaAch.
P. rufescensHoffm.
C. aculeata Fr.
Of lowerplants those withan asteriskare the mostimportant.
Sedumand Cochleariaoccurnearthe sea only. In slighthollows,especially
where the general surfaceis ratherexposed, a society of Salix polaris and
Cetrariaislandicais very common. The mossesand lichensare usuallyrather
scattered. All the lichensmentionedabove, togetherwitha fewothers,occur
on "fjaeldmark" (stonydesert)in Iceland (20).
The followingis a typical animal communityfromthe "fjaeldmark":
Isotoma multisetisCarp. and Phillips
Exechia frigidaBoh.
CamptocladiuslongicostaEdw.
Hymenoptera: Pontania birulaeKonow.
Spiders:
Coryphaeusholmgrenii
Mites:
Cyta brevirostris
Scutovertexlineatus
SphaerozetesnotatusThor.
Oligochaeta: Enchytraeusalbidus Henle.
Birds:
Purple Sandpiper(Erolia maritimamaritimaBriinn.) nestingC
Collembola:
Diptera:
The ArcticFox, formerly
common,has been almostexterminatedby man,
as in Spitsbergen.Noniewas seen by us. In summerit feedson birdsand their
eggs and young. In winter,in other countriesit partly makes caches of
lemmings,etc. (26), and partlygoes out on to the sea-ice,whereit eats mostly
the leavings of bears, e.g. remainsof seals, of which the bears oftenonly
eat the fat (59, 38), and theirexcreta(39). Theredoes not seemto be enough
foodon land in Spitsbergenand Bear Island formakingwintercaches (e.g. no
lemmings)and the foxesprobablylive then mostlyon the sea-ice (cf. 41).
Thus the numberof foxes depends not only upon the amount of direct
destructionby man, but also indirectlyon the numberof bears killed. The
Polar Bear still visits Bear Island in small numbersin winter,fifteenbeing
shot in 1919-20 and threein 1920-21.
The Purple Sandpiper,the commonest"fjaeldmark" bird,will eat almost
anythingit can get, judgingfromits habits in Spitsbergen.The stomachof
one bird containedCollembolaand flies(June 15th). The followingbird-lice
occur on this species: PhilopterusfusiformisDenny, Degeertellazonaria
Nitzschand D. arctophilusKell. and Chap.
The Collembolaor Springtailsappear to feed on decayingplants,and the
and Sphaerozetes,
are vegetarian. Coryphaeus
mites,at any rate Scutovertex
is knownto eat Collembolain Jan Mayen(7), and probablyfeedsalso on flies
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222 Contributions
to Ecology of Spitsbergenand Bear Island
and mites. It seems to make no web. The lifehistoriesof the fliesare very
littleknown. Camptocladiuslongicostaand the mite Sphaerozetesoccur sometimes in flowersof Saxifraga oppositifolia.The sawflyPontania btrulaewas
seen walkingabout on Salix polaris, and probablylays eggs in it. The food
of the adult is unknown. The Snow Buntingeats this sawfly.
The Enchytraid worms of the Arctic seem to replace ecologicallythe
of lowerlatitudes,owingto theirpowersof withstandingfreezing
earthworms
(see 60).
Otherspecies occurringon the "fjaeldmark" are:
Collembola:
Achorutesviaticus
Isotoma viridisBourlet
Xenylla humicolaFab.
Folsomia quadrioculataTeb.
Sminthurinus
in fungi
nigerLubb.
Tetracanthellapilosa Schoett undermoss
Diptera:
CamptocladiuseltoniEdw.
Diamesa hyperboreaH.
D. septimaEdw.
D. ursusKieff.
Lauterborinacaracina
Metriocnemus
ursinus
OrthocladiusconformisHolmg.
Sciara praecox Mg.
Trichoceralutea Becker
Hymenoptera: AmauronematusvillosusThoms.
PristiphorafrigidaBohem.
Mites:
Bdella groenlandicaTrag.
Hypoaspis ovalis L. Koch
Birds:
Golden Plover (CharadriusapricariusapricariusL.) r. nesting
Snow Bunting(Plectrophenaxnivalis nivalis L.)
feeding
The latterthreeCollembolaare recordedby Wahlgren(65).
Of the Collembola Onychiurusis never presentin exposed habitats, but
lives under stones and among plants. Isotomaviridtsseemsable to stand air
ofa greaterevaporatingpowerthan the otherspecies; it oftenwalks about on
open ground. The closely allied I. multtsetis,
however,seems more oftento
inhabit plants. Achorutes,
Xenylla and AgrentiabidenticulataTullb. (another
collembolan)occur typicallyon or near water. Xenylla was once found in
flowersof the Purple Saxifrage. The two sawfliesprobablylive on Saltx.
3. HERB-MAT(withGrass).
This is developed in shelteredplaces, and appears to be a climax to the
"fjaeldmark." It seems to agree with the various herb-matsdescribedfrom
Greenland(50, 67), etc., but differs
in havinga good deal of grass,probably
Catabrosaalgida R.Br. The surfacein these areas is comparativelystable
owingto the closed vegetationbindingthe soil. On a shelteredrockyslope
in the TetradiumLimestoneregionwere:
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V. S. SUMMERHAYES AND
C. S. ELTON
Phanerogams:
Catabrosa algida
Cochleariaofficinalis
Cerastiumalpinum
f.
Draba alpina
Oxyriadigyna
Saxifragacaespitosa f.
f.
S. cernua
f.
S. oppositifolia
Bryophytes:
Bryumsp.
ClimaciumdendroidesW. & N.
HypnumvaucheriLesq.
H. uncinatumand var. plumulosumB. & S.
Lindb.
Mniumaffinevar. integrifolium
Pseudoleskeacatenulata B. & S.
P. tectorumB. & S.
Tortula ruralisEhrh.
Lichen:
Cladonia sylvaticaHoffm.
Collembola:
Onychiurusarmatusvar. arcticus
223
plantsare theusual " fjaeldmark" species. Sax4fragacernuais
The flowering
thanin "fjaeldmark." A fewmigrantPink-footedGeese
luxuriant
muchmore
Baill.) wereseen on the smallareas ofgrassyland. Their
(Anserbrachyrhyncus
dung consistedof remainsof moss and grass. Brent Geese (Branta bernicla
berniclaL.) also occur on migration. Neitherspecies breedshere. Skuas and
Purple Sandpipersnest.
" Skua Hummocks." We have giventhisnameto whatis perhapsthe most
strikingtype of herb-matoccurringin the region. The Skua hummocksare
small grassypatches scatteredover the "fjaeldmark." They are the resultof
constant manuringby the Arctic Skua (Stercorariusparasiticus L.), which
nestson the tops of hillockswhichare the firstplaces to be clear of snow in
spring. The male stands on neighbouringhummocksduringthe breeding
season, watchingforenemies,chieflyfoxes,whichthe Skuas are able to drive
(40)
off. Thus the hummocksbecomewell manuredby the birds. Middendorf
has describedsimilarhummocksin Siberia.
Thereis a considerablelayerof peaty soil (I to 6 ins.) on thesehummocks,
and a varied floraof ratherstuntedindividuals.The phanerogamsare typical
"fjaeldmark" species, Saxifraga oppositifolia,S. caespitosa,and Catabrosa
whilethe presenceof Cochleariais signialgida beingthe most characteristic,
on these hummocksin Spitsbergen.
list
of
a
plants
ficant. Wulif(70) gives
The followingcryptogamsoccur:
Bryophytes:
BryumpallescensSchl.
B. pseudo-triquetrum
Cynodontiumvirens
var. arcticum
Dicranella varia Schimp.
DicranumfuscescensTurn.
var. congestumHusnot.
Dicranoweisiacrispula
var. atrata
Encalypta commutataN. & Horsch.
Hypnumuncinatum
Myurellajulacea B. & S.
Pseudoleskeatectorumformas
RhacomitriumlanuginosumBrid.
Tortularuralis
Lichens:
AlectoriaochroleucaNyl.
BiatorinaregelianaKorb.
Cetrariaaculeata,
f. hispida Cromb.
Cladonia furcat4Schrad.
var. spinosa Leight.
C. rangiferina
Lecanora epibryonAch.
L. tartareaAch.
SphaerophorusglobosusWain.
Algae:
Nostoc sphaericumVaucher.
(among moss)
is the mostprominentmoss.
Hypnumuncicnatum
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224 Contributions
to Ecology ofSpitsbergen
andlBecarisland
The invertebratefauna is of the same type as on the "fjaeldinark." The
alliinalsfoundwere:
Collembola: Isotoma multisetis
Xenylla humicola
Diptera:
Diamesa ursus
Lauterborinacaracina
Spiders:
Coryphaeusholmgrenii
Mites:
Bdella groenlandica
Oligochaeta: +
The Skua lives by robbingotherbirds. It attacks se'-birds (Guillemots,
Kittiwakes,etc.) causingthemto disgorgetheirfood,and also sucks the eggs
both of the cliff-birds
and of the Red-throatedDiver (Colymbusstellatus
Pontopp.) and NorthernEider. Thus the hummockcommunitiesdepenidfor
theirexistenceultimatelyon sea animals.
4. Moss
HEATH.
The communityoccursin the centreof Bear Island west of MounitMisery,
coveringlargeareas, and also locallyin otherparts. It seemsto be developed
onidry, stony flats,and agrees generallywith the various Moss Heaths describedforotherarctic countries(9, 25). The famous "Grimtmia
Heath" of
Iceland is ofthe same type(25). A certainamountof shelterfromwindseems
advantageous.
The dominantplant is Rhacomitrium
lanuginosumas in othercountries,
but on Bear Island Hypnumuncinatumis relativelymoreimportant,alid ill
slightlydamperplaces, formsalmost pure societies. Saltx polart'sis the onily
commltnon
phanerogam.A mnore
completelist of plants is given below:
Phanerogams,etc.:
Equiisetumvariegatumiiu.
*Oxyria
digyna
*Ranunculuspyginaeus
Salix polaris
Saxifragacaespitosa
*S. cernua
1.
r.
f.
o.
1.
Lichens:
CetrarianiivalisAch.
Hoffm.
Cladonia rangiformis
(f. amongstones)
Bryophytes:
*AulacomniunipalustreSchwaegr.
Dicranum
fuscescens
HylocomiumsplendensB. & S.
*Hypnumuncinatumand var. pluiiiulosuii
Lophozia hatcheriSteph.
*Mniumaffinevar. integrifoliuii
PolytrichumalpinumL.
PtilidiumciliareHampe.
l.a.
Rhacomitriumlanuginosunid.
*Timmiaaustriaca
Tortularuralis
Species withan asteriskare characteristicof the societyof Hypniuma
uncinatut.
The faunaofthisarea was iiot workedmuch,but it is ratherpoor. Isotonta
was found.
V'ridis
5. ROCKAND BOULDERCOMMUNITIES.
In a great imiany
places in Bear Island, owingto the intenseweatherin7g,
thereare large areas, oftenquite flat,which are covered by heaps of loose
blocks,6 inchesto 10 feetin diamneter.On Mount Miserythe SpiriferLinmestoneweathersintolargeblockswhichformordinaryslopingscrees. Thereare
also erraticboulderswhichoftenhave many cracksand crevices,and may be
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V. S. SUMMERHAYESAND C.
S. ELTON
225
broken by frostinto smaller fragments(27). Screes consistingof smaller
particlesare too unstableto supportany life.
The productionof the block plains and heaps may be due to the fact that
by " solifluction"(2) the finerearthhas flowedaway leaving the large rocks
behind. It is knownthat "solifluction"can occur on verygentleslopes,and
such earthmovementshave been noticedon Bear Island.
The block communitycan be dividedintotwo main divisions,whichgrade
into one anotherin many places: (a) Communityon rock surfaces(Lithophytes),(b) Community
in rockcrevicesand betweenboulders(Chomophytes).
(a) The blockssupporta richlichenand moss floraon the partsprotected
by snow in winter. The lichensare usually crustaceous,but may be foliose
or even fruticosein favourablespots. The followinglist includes the more
importantones:
Gyrophoraerosa Ach.
Lecanora galactinasubsp. dispersaNyl.
L. polytropaSchaer.
Lecidea goniophilaSchaer.
L. pantherina
Placodium cerinumEhrh.
P. elegansDC.
P. rupestreBranth. & Rostr.
P. rupestrevar. calvum AL.Sm.
PolyblasteaintercedensLoennr.
RhizocarponcalcareumTh.Fr.
R. geographicum
Massal.
Thelidiumpyrenophorum
The lichencommunitiescertainlyvary withthe chemicalnature of the rock.
Mosses are by no means so commondirectlyon rocksas are the lichens.The
only commonone is Grimmiaapocarpa Hedw. var. alpicola Hook and Tayl.,
whichis veryabundantlocally. Othersare Dicranoweisiacrispulavar. atrata,
W. and M., and D. schistiLindb. (see 5). Animalsare absent.
Dicranumstairkei
(b) The crevicesbetweenthe bouldersand depressionson them contain
a muchmorevariedvegetation. A successionto Moss Heath or "fjaeldmark"
takes place withthe accumulationofhumusin the crevicesand on the surfaces.
viz.: (1) That at the
Two divisionsof the crevicefloracan be distinguished,
mouthsof crevicesand on humus collectedin depressions,and (2) the communityinsidethe crevices.
(1) The florais mainlycryptogamic.There are only a fewphanerogams,
such as Oxyria,Saxifragaoppositifolia,
etc., as usuallythe humusis not deep
enough. Most of the vegetationconsistsof bryophytes,althoughin places,
e.g. on Mount Misery,there is a considerableadmixture of lichens. The
followingplants occur:
Bryophytes:
BrachytheciumsalebrosumB. & S.
Dicranumbonjeani de Not.
D. starkei
HarpanthusscutatusSpruce
Hylocomiumsplendens
Hypnumuncinatum
Jour. of Ecology XI
Lichens:
CetrariahiascensTh.Fr.
C. islandica var. tenuifoliaWain.
Cladonia bellidifoliaSchaer.
C. foliaceaWilld.
C. furcata
C. gracilisWilld.
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15
226 Contributions
to Fcology ofSpitsbergenand Bear Island
Bryophytes(cont.):
HypnumvernicosumLindb.
Lophozia hatcheri
L. quinquiedentataCogn.
L. ventricosaDum.
Polytrichumalpinum
Ptilidiumciliare
Rhacomitriumlanuginosum
Tortula ruralis
Webera cruda Schwaegr.
Lichens (cont.):
Cladonia pyxidata Hoffm.
C. sylvatica
Lecanora tartarea
Parmelia omphalodesAch.
Peltigeracanina
Sphaerophorusglobosus
Stereocaulonpasehale Fr.
The large numberof lichens shows the dry nature of the substratum,but
thereis a definiteadmixtureof wet-lovingtypesamongthe mosses.
(2) Betweenthe rocksthereare oftenconsiderablecavitiesin whichit is
possiblefora man to crouch. The innerpartsoftheseholesare verydark,and
indeed one reachesfinallya regionin whichthe lightis insufficient
forplant
life. The sides of the blocksare usuallywet withwatertricklingfromabove,
and a hygrophilous
typeof vegetationis able to exist. In thesecrevicesliverworts predominate,the more importantbeing Blepharostomatrichophyllum
Dum. and LophozialongidensMacoun. The mosses Weberacrudaand Swartzia
montanaLindb. are also important.
Mosses:
Blindia acuta B. & S.
Dicranummolle Wils.
HypnumsarmentosumWahl.
H. stramineumDicks.
H. uncinatum
Swartzia montana
Timmiaaustriaca
Webera cruda
W. nutans Hedw.
Algae:
CosmariumpseudoholmiiBorge.
StaurastrumgrandeBulnh.
Liverworts:
Blepharostomatrichophyllum
Cephalozia bicuspidataDum.
C. leucanthaSpruce
C. serriflora
Lindb.
Cephaloziellabyssacea Warnst.
Diplophyllumalbicans Dum.
Lophozia alpestrisEvans
L. longidens
L. porphyroleucaSchiffn.
L. quinquedentata
L. ventricosa
Scapania curtaDum.
S. irriguaDum.
and Diplophyllumoccurin caves and cleftsin Iceland (25).
Blepharostoma
The Snow Buntingnestsin the crevices,but findsits food in many other
places. It eats sawfliesto a great extent. One cock when shot was foundto
be carryinghalf-a-dozenin its mouth (Amauronematus
and other species).
Amauronematus
occurredin the stomachof one birdand Pontania in another
with leaves of Sedum rhodiola. Swenander(62) foundseeds of Cochleariain
the stomachof one bird.
The loweranimals werenot workedout.
6. MOSS-MAT.
This is similarto Cleve's (9) moss-mat,but thereare morephanerogams.
It occursin hollowswherethe snowmeltsslowly,or wheresnow-watertrickles
downcontinuallyfromabove. Mossescoverthe soil,but theyare not the very
damp-lovingspecies. The Hypnumuncinatumsocietydescribedunder "Moss
Heath" is closelyallied to this,but drier. The areas may dryup completely
late on in the season, and thisdistinguishes
moss-matfromwettundra. Salix
polaris is commonin many places, whileRanunculussulphureusSol. is often
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V. S. SUMMERHAYESAND C.
S.
227
ELTON
encountered. Other species not common in drier places are Saxifraga rivu-
laris L. and Polygonumviviparum.Thereare also:
Phanerogams,etc.:
Draba alpina
Equisetum variegatum
Oxyriadigyna
Saxifragacaespitosa
S. cernua
S. nivalis
S. oppositifolia
Bryophytes:
Cynodontiumvirens
Hypnumuncinatum
Timmia austriaca
a.
Lichens:
Cladonia rangiferina
Lecidea vernalisAch.
- StereocaulonalpinumLaur.
The Kittiwakeswerewatchedpickingmoss fortheirnestsfrommoss-mat,
and areas up to 1 ft. in diameterare torn up in that way. Swenander(62)
states that the Kittiwakes use moss throughoutthe season for renovating
theirnests,and thismustbe ofimportancewhenone considersthevast numbe-r
ofbirdsconcerned.The less commonGlaucous Gullsalso use thismoss.
Similarvegetationoccurson the flatplateau at the top of Mount Misery,
plicatumB. and S., Amblywhereon a damp mossyarea were Brachythecium
stegiumserpensB. and S., and the lichensSolorina croceaAch., and Psoroma
hypnorumS. F. Gray. A few Ptarmigan (Lagopus mutusSund.) occur here
and also one or two foxes. The Ptarmiganfeeds on all kinds of plants,and,
to a certainextent,on insects (39).
7. WET TUNDRA.
This consists of regions which are wet throughout the spring and summer.
The largest stretches are in Ymers Valley between a number of ponds along
the river. Mosses usually predominate while flowering plants are of little
importance. Three divisions can be recognised:
(a) Pond Marginal Community-with still, fairly well-aerated water.
(b) StreamMarginalCommunity-withrunningwell-aeratedwater.
(c) Moss Bog-with partiallystagnantwater.
(a) This type of wet tundra occurs around nearly all the ponds, except
wherethereis a rockybank. The vegetationformsa prominentmossybank
around the pond. Grasses and other phanerogamsoccur in places, but not
frequently. The mosses were not worked very thoroughly,but probably
listshowsthetypeofvegetation:
includemostofthebog species. The following
AulacomniumturgidumSchwaegr.
Dicranoweisiacrispula
Lindb.
Hypnumbrevifolium
var. gracilisBerggr.
Hedw.
Hypnum cordifolium
H. vernicosum
H. badium Hartm.
H. sarmentosum
Berggren(5) recordsthe last two species.
Under stones,on the marginof Ella Lake by an inflowingstream,the
followinganimals occurred:
Collembola:Achorutesviaticus
Isotoma multisetis
I. viridis
SminthuridesmalmgreniTullb.
Xenylla humicola
Mites:
Cyta brevirostris
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15-2
toEcology of SpitsbergenantdBear Island
228 Contributions
(b) Thisis presentonlylocally,as owingto the erosionthestreamsides
Vegetationis practically
are usuallyan unstablemass of rockfragments.
in theselatterplaces,but Isotomaviridisis oftenfoundthere.In
inhibited
reportsthe
placesthereis a narrowbandofmossby thestreams.Berggren
following
nearMountMiseryand elsewhere:
BryumobtusifoliumLindb.
B. turbinatum
var. latifoliumSchleich.
HypnumochraceumWils.
H. polygamumSchp.
Paludella squarrosaEhrh.
PhilonotisfontanaBrid.
Swartzia montana
Webera albicans Wahlb.
var. glacialis Limpr.
polareOfversand
On stonespartlycoveredby the wateroccurHypnrum
L.
H. filicinum
(c) Thisis presentin flatareas-in valleysand consistsmainlyofmosses,
at WalrusBay. The chiefphanerogams
a well-developed
exampleoccurring
L. and Ranuncutus
S. hirculus
sulphureus.The first
are Saxifragarivularis,
in WalrusBay bog,whiletheothersare locallyabundant
namedis frequent
mossesoccurin bogson BearIsland:
in YmersValley. The following
CinclidiumstygiumSw.
Hypnumbrevifolium
H. fluitansL.
Lindb.
H. intermedium
H. polygamum
Meesia triquetraAngstr.
Hypnumsarmentosum
and var. fontinaloidesBerggr.
H. stellatumSchreb.
H. vernicosum
H. turgescensSchp.
SplachnumvasculosumL.
animalsoccurin WalrusValleybog:
Thefollowing
Collembola:Achorutesviaticus
Agreniabidenticulata
Diptera:
Camptocladiuseltoni
C. oxonianusEdw.
Metriocnemusursinus
ofoldwalrusboneslyinghere.Theenormous
numbers
Therearelargenumbers
ofwalrusformerly
occurring
(e.g. a thousandwerekilledin sevenhoursin
1608(12), whilelargeherdsstillvisitedtheislandin 1825(46)) musthave
the valleys. Underwalrusskullswerea good
in manuring
beenimportant
andExechiafrigida).
manyflies(Metriocnemus
was not workedout by us. SeveralTardigrades
occur
The micro-fauna
and Macrobiotes
ofthegeneraEchiniscus
(55).
(b) Freshwater Communities.
1.
STILL WATER
(Lakes and Ponds).
The pondsare all ofan arctictype,and are notunfrozen
formorethan
twoto threemonths.It was too earlyin the seasonto do completework.
Almostall thepondsin thesouthoftheislandare rockbasins,but mostof
mossvegetation.
themhavea decidedmarginal
and
Ella
Lake
17th
19th). Thisis a largelake,whosegreatest
(a)
(June
brownloamymud,butthe
depthis 120feet.The bottomconsistsofgreyish
shoresarestony.Thewateris alkalineat thesurface.
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V. S. SUMMERHIAYESAND C. S. ELTON
229
Some of our collections(algae and crustacea) were destroyedby accident.
boryannum1
Lag., Pediastrurn
Lagerheim(35) recordedHormosporasubtilissirna
Grun. That and Hormosporas
Menegh., and the diatom Synedrafiliformlis
folluld
formedthe bulk ofthe planktonat his visit. The morecommondiatomrls
by Cleve (10) and Lagerheim,were:
CampylodiscushibernicusEhb.
Diatoma tenue Ag. var. elongata Ag.
Navicula rotaena Rabh.
Pinnulariacurta Cl.
AchnanthesmicrocephalaGrun.
CocconeisplacentulaEhb. var. eiiglyptaElhb.
The last two wereabundant on filamentousgreenalgae.
Little is knownof the fauna,but the followingoccur:
(Lilljeborg(37) June 1898, July 1899)
ChydorussphaericusMuller
CyclopsstrenuusFischer
C. vicinusClaus.
Chironomidlarvae
Diptera:
Trichoptera: Apatania aretica Boh. (larvae and 1 adult 9)
Hydrachnidae: SperchonlineatusS.Thor. (,, Y, nymphs)(57)
Lonnb.
Salmo umbla var. salvelinus-insularis
Fish:
Crustacea:
The caddis-flyApatantamakesa case out ofsand and mud. It and the watermite Sperchonare abundant on the stones among algae, etc. The fishis a
deep-waterchar,but nothingis knownofits ecology. The fauna of Ella Lake
formsan exceptionto the usual type in Spitsbergenand Bear Island, owing
to the extentand depth of the lake.
(b) Pond 1. (June 13thand 15th) Cleve (10) Habitat 13 and Lagerheii
(35) "Pond near Russian Haven."
In a rockybasin in the dolomite,100 to 200 yards long. The water was
unfrozen,and was very clear and alkaline. The depth is 7 metres,and the
bottomis black mud (10). The shoreis stony,and almostdevoidofvegetatioln
exceptalgae and a littlemoss. The followingoccurred:
PLANTS:
Naeg.
Chlorophyceac: Coelastrummicrosporum
GongrosiradebaryanaRab.
*Pediastrumboryanumvar. longicorneReinseb.
P. constrictumHassall.
P. integrumNaeg.
P. muticumKuetz.
P. sturmiiReinsch.
Rhizocloniumsp.
*Scenedesmusbijugatus Kuetz.
S. denticulatusLagerh.
S. obliquus Kuetz.
*S. quadricauda Breb.
Cyanophyceae: *OscillatoriatenuisAg.
*Diatomns:
Amphoraovalis Kuetz.
Diatoma elongatumvar. tenue VH.
FragilariamutabilisGrun.
F. construensEhb.
Navicula peregrinavar. polarisCl.
N. rhyncocephalaKuetz.
(Those withasterisksrecordedby Lagerheimand Cleve.)
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and Bear island
230 ContributionstoEcology oj Spitsbergen
ANIMALS:
Rotifera:
Oligochaeta:
Tardigrada:
Diptera:
Crustacea:
Plankton:
Littoral:
PolyarthraplatypteraEhrb.
Nais josinae Vejd.
MacrobiotesmacronyxDuj.
Orthocladiusconformis(adults on surfaceof water)
Larvae.
Daphnia longispinaMiiller
Cyclopsstrenuus
LepidurusarcticusPallas
Chydorussphaericus
Cyclopsgigas Claus.
Pond 2. (June 17th.) A small pond in Ymers Valleyfedby a largestream
flowingfromthe hills. These hills are occupied by sea-birdcolonies,and so
the wateris probablyrichlymanured. It was alkaline. Cleve gives a longlist
ofdiatoms(his Locality 10). Cyclopsgigas occurshere.
Pond 3. A small pond in a rockybasin 100 or 200 feetabove Ella Lake.
The crustaceashowedthat its fauna resemblesPond 1. Therewereno caddisfliesor water-mites.The followingalgae occur:
Chlorophyceae:Pediastrumboryanumvar. longicorne
Rhizocloniumsp.
StaurastrumbieneanumRabenh.
Cyanophyceae: TolypothrixdistortaKuetz. var. penicillataLemm.
GeneralRemarks.
The plankton includes Daphnia longisptna,Cyclops strenuusand the
rotiferPolyarthraplatyptera. Cyclops vicinus comes in Ella Lake. Other
crustacea recorded(37, 52) fromthis regionare littoraland bottom-living
forms,viz.: MacrothrixarcticaSars., MaraenbiotusbruceiRichard, Eucypris
arcticaSars., Candona candida Muller. Previous recordsconfirmin a general
way the communitiesdescribedhere.
Large flocksofKittiwakeswereseen on the waterbathing,probablypartly
in orderto get rid of marineparasites. Glaucous Gulls and NorthernEiders
also occur. The latterare knownto breedby ponds in the northofthe island.
The Long-tailedDuck (Harelda glacialis L.) and CommonScoter (Oedemia
nigranigraL.) were observedin small numbers,and the former,at any rate,
breeds. Red-throatedDivers are common,and nestby the water'sedge. They
evidentlyfeedon the fishin Ella Lake sincetheyeat Salmo alpinus in Greenlland (39). Purple Sandpipersalso feedby the water. CertainCollembolaare
said by Wahlgren(65) to occur on the surface of water, e.g. Sminthurides
Achorutes
viaticus,Xenyllahumicola.
malmgrent,
2. RUNNING WATER (Streams).
The streamsoftenhave a largeamountoffilamentous
greenalgae attached
to the stones. The followingalgae werefoundin various streams:
Kentrosphaerasp.
Pediastrumboryanumvar. longicorne
P. integrum
PhormidiumuncinatumGomont.
UlothrixsubtilissimaRab.
U. tenerrimaKuetz.
U. sp. (allied to U. scutata)
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V. S. SUMMERHAYESAND C.
S. ELTON
231
Cleve recordsmany diatoms(10). The moreimportantones are: Achnanthes
mtcrocephala,
Diatoma tenuevar. elongata,Fragtilariaarcus Kuetz. and Merdion circulareAg. The firstis epiphyticon filamentousalgae. Ranunculus
Rottb. occursin streamson Bear Island (1).
hyperboreus
No animals were foundamong the algae. This is confirmedby observations(55) that none of the streamscontainsTardigrades.
MISCELLANEOUS.
The northernhalf of the island is different
fromthe southernin that the
ArcticTern (Sternaparadisea Briinn)and Grey Phalarope (Phalaropusfulicarius Iredale) breedthere.
The shoresof Bear Island are mostlyprecipitous,and owingto the small
riseand fallofthe tide in the Arctic(not morethan 4 or 5 feet)the inter-tidal
zone is narrowand appears to be almostdevoid of life.
An exampleof " redsnow" collectedturnedout to consistofthe remainsof
marinecrustacea. Lamont (36) says that the "red snow" examinedby him
in Spitsbergenconsistedofthe droppingsof Little Auks. Of courseSphaerella
nivalis Som. does occur,but the cases givenabove show that it is not safe to
judge by colouralone (see 42).
FOOD AND ENEMIES
("NITROGEN CYCLE").
Food is extremelyscarce in the Arctic,both on land and in freshwater,
thoughit is plentifulin the sea. Most of the scavenginganimals live on decaying plants, and are, therefore,practically equivalent to herbivoresin
the food cycle. There are no elaborate "chains" of species dependingon
animals which eat the dung or decaying bodies of other species. Such a
"short-circuiting"
of the nitrogencycle (whichexistsin othercountries,e.g.
badgerseatingbeetleswhichpreyon dung feedersin England) appears to be
unimportantin Spitsbergen.Dead animals are very rarelyfound (46), and
when they do occur are devouredby vertebrates(e.g. reindeerby Glaucous
Gulls and dead whales by bears). Whereanimalslike whaleshave the chance
of decaying,theydo so veryslowly. The questionsof nitrogen-fixing
bacteria
and ofthe effectofthunderstorms
are leftopen,sincethereis no directevidence
at any rate are not at
eitherway forthe Arcticregions,but thunderstorms
all frequent.Soils collectedfromGreenlanid
(4) on examinationshowedseveral
species of putrefactiveorganismssuch as Bacillus subttils,B. vulgatusand
and denitrifying
Bactertium
zopfii. Nitrifying
organismswere also discovered.
Mr Sandon reportsbacteriaand protozoafromall the soils collectedby us.
The diagram (Fig. 2) gives the food relationsof the land and freshwater
animals and plants in the southernpart of Bear Island. Ella Lake is not
includedin this scheme,since it is an exceptionalcase. The directevidence,
whereexisting,forthe truthof this diagramwill be foundat various places
in thispaper. Data fromSpitsbergenanimalsofthe same speciesare included,
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and Bear Island
232 Contributions
to Ecologyof Spitsbergen
forthatcountry
and the diagramwithsomealterations
wouldsuffice
itself,
butwouldbe rathermorecomplicated.
It willbe seenthata largepartofthefoodsupplyofanimalsand plants
caseofthis,theresultofconcentration
comesfromthesea. Themoststriking
is
ofthe Skua Hummocks
in a smallarea,is the Bird Cliffs.The manuring
of
in theseplacesshowstheeffect
anotherexample.Theluxuriant
vegetation
inN.W.Greenland
an exceptional
supplyofnitrogen.
Thereareno BirdCliffs
frozensea, but plantsoccuralthoughnever
(51) owingto the permanently
I
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quiLLEMoTS
--'
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ZIA
IM11ERAL
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-
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;
.:
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RMRl AN
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EIbERR
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iA
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fPURPLE
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. . OLIGOCHA ETA
FIG. 2. Diagram of "NitrogenCycle" on Bear Island.
.......
-----
.Probable,but no evidencefromhere.
Transformation.
luxuriantly.Here therecan be no help fromthe sea and the presenceof
bacteriaseemsnecessary.However,thereare also methods
nitrogen-fixing
fromthe land. One of theseis the usual washingdown
of loss of nitrogen
is dueto migratory
ofnitrates.Another
birds.ThePurpleSandpiper
maybe
takenas an example.It feedsonCollembola,
etc.,whichfeedon dead plants,
and theyoungsandpipers
are rearedon the samefood. In thelate summer
bothadultandyoungdepart,andthenitrogenous
material
represented
bythe
youngis lostto theArctic.
In freshwaterwe may notethe almosttotalabsenceof carnivores.In
theEntomostraca
forma " key-industry"
temperate
regions
whosefunction
is
the conversion
of microscopic
plant foodinto a formutilisableby larger
animals(fish,insects,etc.). Here the Entomostraca
have no enemiesin the
wateritself(exceptinElla Lake) thoughbirdseat Lepidurus
andprobably
the
otheranimals.Therolesplayedbythevariousscavenging
speciesareunknown,
butthereis sureto be somedivisionoflabouramongthem.
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ANDCe S. ELTON
V. S. SUMMERHAYES
233
The parasitic cycles are not included in the diagram, since nothingis
known of them. Parasites and carnivoresare essentiallythe same in their
are due to the relative
food relationswithotheranimals,and the differences
sizes of the eater and the eaten. If the animal is below a certainsize it may
The resultofthis
if above that size it is free-living.
dwellin or on its food-host,
is that the species in food-chainsof parasitesget smallerand smaller,whilein
food-chainsofcarnivorestheyget,on the whole,largerand larger(see Fig. 2).
III. SPITSBERGEN.
INTRODUCTION.
Unlike Bear Island, Spitsbergenis quite an extensive country,being
280 miles fromNorth to South, and over 200 miles fromEast to West (see
Map, Fig. 3). West Spitsbergenand PrinceCharlesForelandonlywerevisited
by us. West Spitsbergenis cut into by many deep fjords,the chiefone of
thesebeingIcefjord,wheremostof our observationsweremade.
Spitsbergenis a mountainouscountry,therebeing very littleflatground
or even lowland. Apartfromraisedbeaches,etc.,the mountainsabut directly
on to the sea. As mightbe expected fromthe latitude, glaciation is very
severeand much of the interiorof the countryis coveredby permanentneve
and the glaciersarisingtherefrom,
and is devoid of life. Since many of the
glaciersreach the sea the narrowcoastal stripis cut up into isolated parts,
and dispersalis hindered.
The geology of the countryis very varied, the rocks being of all types
chemically,but as faras can be seen this has littleeffecton the lifein Spitsbergengenerally,althoughsome plants seem to be restrictedto certainsoils,
or at any rate growbetteron them. In NorthGreenlandaccordingto Ostenfeld(51) thevegetationis richeron limestonethan on othersoils. The physical
propertiesofthe rocksseem moreimportantin relationto denudation.
The climateofSpitsbergenis relativelymildforsuch a highlatitude,owing
to the Gulf Stream driftwhich approaches the west coast. The following
table (Table I) is compiledfromvarious recordsfordifferent
parts of Spitsbergen,but is in no sense complete. It includesrecordsfrom1872-1922.
TABLE I.
Mean
temperature Maximum
0C.
January
February
March
April
May
June
July
August
September
October
November
December
-13 9
-15 9
-17-7
- 15 2
- 70
0C.
40
1-6
3-8
50
125
1-2
16-0
- 2-5
- 8-4
8-3
-0 6
4-6
26
- 10 6
- 127
160
90
4-2
5-6
Minimum
0C.
- 32 4
-38-2
-40 0
- 32-6
-19-4
-14-0
- 61
- 2-6
-19-0
- 27 2
- 23 6
- 32-2
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ANDCe S. ELTON
V. S. SUMMERHAYES
233
The parasitic cycles are not included in the diagram, since nothingis
known of them. Parasites and carnivoresare essentiallythe same in their
are due to the relative
food relationswithotheranimals,and the differences
sizes of the eater and the eaten. If the animal is below a certainsize it may
The resultofthis
if above that size it is free-living.
dwellin or on its food-host,
is that the species in food-chainsof parasitesget smallerand smaller,whilein
food-chainsofcarnivorestheyget,on the whole,largerand larger(see Fig. 2).
III. SPITSBERGEN.
INTRODUCTION.
Unlike Bear Island, Spitsbergenis quite an extensive country,being
280 miles fromNorth to South, and over 200 miles fromEast to West (see
Map, Fig. 3). West Spitsbergenand PrinceCharlesForelandonlywerevisited
by us. West Spitsbergenis cut into by many deep fjords,the chiefone of
thesebeingIcefjord,wheremostof our observationsweremade.
Spitsbergenis a mountainouscountry,therebeing very littleflatground
or even lowland. Apartfromraisedbeaches,etc.,the mountainsabut directly
on to the sea. As mightbe expected fromthe latitude, glaciation is very
severeand much of the interiorof the countryis coveredby permanentneve
and the glaciersarisingtherefrom,
and is devoid of life. Since many of the
glaciersreach the sea the narrowcoastal stripis cut up into isolated parts,
and dispersalis hindered.
The geology of the countryis very varied, the rocks being of all types
chemically,but as faras can be seen this has littleeffecton the lifein Spitsbergengenerally,althoughsome plants seem to be restrictedto certainsoils,
or at any rate growbetteron them. In NorthGreenlandaccordingto Ostenfeld(51) thevegetationis richeron limestonethan on othersoils. The physical
propertiesofthe rocksseem moreimportantin relationto denudation.
The climateofSpitsbergenis relativelymildforsuch a highlatitude,owing
to the Gulf Stream driftwhich approaches the west coast. The following
table (Table I) is compiledfromvarious recordsfordifferent
parts of Spitsbergen,but is in no sense complete. It includesrecordsfrom1872-1922.
TABLE I.
Mean
temperature Maximum
0C.
January
February
March
April
May
June
July
August
September
October
November
December
-13 9
-15 9
-17-7
- 15 2
- 70
0C.
40
1-6
3-8
50
125
1-2
16-0
- 2-5
- 8-4
8-3
-0 6
4-6
26
- 10 6
- 127
160
90
4-2
5-6
Minimum
0C.
- 32 4
-38-2
-40 0
- 32-6
-19-4
-14-0
- 61
- 2-6
-19-0
- 27 2
- 23 6
- 32-2
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to Ecologyqf Spitsbergenand Bear Island
234 Contributions
The anomaliespresenthere,such as the low maximumforOctober,are probably due to the fewrecordsavailable. A temperatureof 19.50 C. in the shade
was registeredin the summerof 1922.
Temperature.During the winterthe temperatureis verylow, but at that
and most of
protectsthe plants and invertebrates,
periodthe snow-covering
overa series
frost
is
free
from
month
no
Apparently
the birdshave migrated.
IF
~
71~j~W4 ERM E(
~
WFS
S P I TS B
FIG. 3. Map of N.W. Spitsbergen.Places dealt with,or referredto in thispaper are marked.
Ofyears,but in realityJulyand Augustmay be so in some years. The temperaturein the sun is oftenquite highon clear days, especiallyin the interior
of 27? and 28? CD.havingbeen recorded.Nathorst
of the fjords,temperatures
(45) has pointedout that the heads of the fjordsenjoy a more continental
climatethan the coastal regions.
Precipitation.Recent recordsgive 11l6 inches as the mean annual pre-
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V. S. SUMMERHAYESAND C. S. ELTON
235
cipitationfor GreenHarbour. The west coast, e.g. Prince CharlesForeland,
probably has rathermore, the interiorof the fjordsless. In any case the
is verylow. Most ofthisfallsin theformofsnowin autumn
total precipitation
and spring,therebeingliftlesnow or rain duringthe summer. In north-east
Greenland,where the precipitationis comparable (6.0 inches per annum),
two-thirdsfalls between October and March, and less than one-tenthfalls
as rain (38). The durationand amount of snowwaterin summerare of great
the plant communities.Owingto the prevalence
importancein determining
of thick mists on the coastal areas much condensationtakes place. Many
lichensmustdependlargelyupon these mistsfortheirwatersupply.
Humidityand Cloudiness. The Relative Humidityis very high owingto
the low temperatures,although the Absolute Humidity is probably rather
small. Cloudy days are frequentin the summerespeciallyaround the coast,
but once the temperaturerises the actual small amount of water vapour is
easily absorbed and a very clear atmosphereresults. This is importantin
increasingthe directinsolationwhich the groundreceives. The distribution
of such animals as Springtails(Collembola)is correlatedwith the humidity
of the air.
Wind. Little is knownof the directionand velocityof the windin Spitswinds occur at various times and help to raise the
bergen. South-westerly
temperature.The directionof the wind depends on the orientationof the
mountain ranges. The 'wind causes many stormsthroughoutautumn and
winterwhichtend to breakup the ice, and thusto raise the temperature.This
breakingup is also probablyan importantfactorin dispersal(see Cassiope).
On the plants,especially,windis a verypotentinfluence(66). Kihlman (33)
has pointedout the drying-outeffectof strongwindswhenthe soil is at a low
and probably"fjaeldmark"is determinedlargelyby windeffect.
temperature,
Denudation. These and otherfactorsproduce a most intenseweathering
and erosionby frost,ice and water.
Of these the two firstoverlap as frosteffectdepends on the formationof
ice and its splittingpowers. By "ice" is meant more particularlythe work
of glaciersand of floatingice. Frost acts primarilyas an agent in splitting
offrockfragments,
but it also producesothersecondaryeffects.Even in the
summerthe soil is generallyfrozenbelow a depth of 18 inches. Hogbom (27)
has shownthat thisfrozensubsoilmay act as a glidingplane down whichthe
soil above, brokencontinuouslyby alternatefreezingand thawing("regelation"), graduallyslides. If one also includesscreesit seems that practically
the whole surfacewherethereis the slightestslope, is on the move. This is
verydestructiveto plant life,and resultsin the almosttotal sterilityof many
areas.
The effectof wateris eitherin rivererosion,or in connectionwith snowwater whichis very destructiveduringthe spring. The latter worksin conjunctionwith "regelation" in manyplaces.
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236 Contributtions
to Ecology of Spitsbergenand Bear Island
SPECIAL REGIONS.
We willdeal withthe specialregionswhichwe visitedin Spitsbergen.They
will be foundmarkedwithdots on the generalmap (Fig. 3).
A. PRINCE
CHARLES
FORELAND.
(North Eastern Region.)
The area dealt withis includedin the map (Fig. 4). It consistsessentially
of threeparts,a centralcore of mountains,an undulatinglowland area next
,q*b
VOG,r:L HoP-KSCALE SVCALA
I
\~~~~~.......
-
q_d9
. 0
= 1s,oo
L
1
T
E
34CD
4 Ip
CARM"IC.1AEL
S]
L
6~~
ln~~~~~~~'T
TAYme
FIG. 4. Map ofnorthernportionofPrinceCharlesForeland. Aftersurveyby Dr W. S. Bruce and
Mr J. Mathiesonlpublishedby the Prince ofMonaco. (By kinldpermission.)
to it, and raisedbeaches of muchlater date on the coast. Th-efirsttwo parts
consistof various rocks,quartzites,sandstones,conglomerates,etc., all poor
in
lime
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V. S. SUMMERHAYESAND C.
S. ELTON
237
The characteristic
featureofthe"Foreland"is therawnessand fogginess
Sea. Clear
mistsfromtheGreenland
of its climate.Thisis due to frequent
variationsare
days are muchrarerthanon themainland,but temperature
probablynotso greatas in thefjords.Thisrawclimatehelpsfrostand other
and thewholecountry
showstheresultsofthese.
erosion,
can be distinguished:
subdivisions
Thefollowing
1. BirdCliffs.
Communities.
2. Maritime
(a) BrackishWater.
(b) Inter-tidal
Zone.
(c) RecentShingle.
3. RaisedBeachSystem.
4. RockyLowland.
(a) With"polygonsoils."
(b) Without"polygonsoils."
(c) Moss-mat.
(d) Freshwater.
(e) ErraticBouldersand Rocks.
5. UplandRegion.
(a) Screes.
(b) StabilisedSlopes.
(c) WetRavines.
1. BIRD CLIFFS.
insomewaysbeionging
is dealtwithherebecausealthough
Thiscommunity
its
characters
dependon the sea. These
to the uplandregion,yet general
thesea withintervening
aresituatedsomelittledistance
from
flatcountry.
cliffs
in thearea is thatat VogelHoek. Hereis a
The onlycolonyofsea-birds
oftheusualtype. The area maybe dividedintotwoparts:
steeprock-wall
(a) thecliff
face,and (b) thescreebelow.
Guillemot,
Kitti(a) On theledgesthereare largecoloniesofBriinnich's
wake,FulmarPetrel,and usuallyabovethemtheirenemytheGlaucousGull.
Puffin
arcticanaumanni
In crevicesthe Spitsbergen
Norton)and
(Fratercula
Auk
nest.
the
rock
wall
are
Little
On
amounts
of
the
large
Enteromorpha
sp.
In thisalgawasfounda richfaunaofrotifers,
tardigrades,
nematodes,
protozoa
and mites(Scutovertex
lineatus).Plantscouldbe seenon theledges,probably
whichis presenton thescreesbelow.
Cochlearia
Goosenestson the
(b) The LittleAuk nestsherewhilethe Pink-footed
slopesaround.The ArcticFox and GlaucousGullhave beenseenhere(21)
as theyleftthenest.
attacking
youngGuillemots
matofflowering
Thereis a continuous
plantson thescree. The absence
ofSaxifragaoppositifolia
is interesting.
It has longbeenrecognised
as oneof
ofArcticandAlpineplants,growing
thehardiest
undermostsevereconditions,
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238 Contributions
toEcology of Spitsbergenand Bear Island
but always seemsto be sensitiveto competition,and is oftenabsent in closed
communities.Here occur:
Phanerogams:
AlopecurusalpinusSm.?
Cochleariaofficinalis
Ranunculuspygmaeus
R. sulphureus
Mosses:
Aulacomniumpalustre
A. turgidum
Salix polaris
Saxifragacaespitosa
S. cernua
S. rivularis
Hypnumuncinatum
S. rivularisand R. pygmaeusare veryluxuriant. Comparisonwithlater lists
showsthat the florais poorerthan that of similarhabitatsin Icefjord.
2. MARITIME COMMUNITIES.
(a) BrackishWater. This is representedby Richard Lagoon. It is a large
tidal lagoon whosewateris brackishat low tide. The lagoonis about 3-1miles
long and W milewide,and is separatedfromthe sea by a bar of roughshingle,
whichis coveredby stormtides only. The only entranceis a narrowchannel
throughwhichthe sea rushesin at the flow,and out at the ebb. A line of
soundings(see Fig. 4) taken betweenhighand low tidesgave depthsof from
41 to 61 feet,and the lagoon cannot be morethan 3 feetdeeperat hightide.
The bottomis gravellyorsandy. Here occurmassesofvariousalgae. Dredging
was carriedout withMr Huxley's help. The followingoccur:
Algae:
ChaetomorphamelagoniumKjellm.
Cladophoraarcta Kuetz.
Desmarestiaaculeata Lamour.
DictyosiphonfoeniculaceusGrev.
Fucus serratusL. f. arctica J. G. Agardh
Laminaria sp.
PhloeosporatortilisStromf.
Pylaiella litoralisKjellm.
Rhodomela lycopodioidesAg.
Hydroida: Opercularellalacerta Johnston
Polychaeta: Capitella capitata Fab.
HarmothoeimbricataLinn.
Ophelia limacina Rathke
Polyzoa (on algae and alive whentaken):
AlcyonidiumgelatinosumLinn.
Gemellarialoricata Linn.
Crustacea: GammaracanthusloricatusSab.
Mollusca: Liocyma fiuctuosaGould
The surfacewater tasted fairlyfresh,but the lower layers in which the
animalslive (6 feet)may be moresaline. Nearerthe entrance,whenthe tide
was flowingin, the surfacechloridecontent("salinity") was 15*77gramsper
litre. Large numbersof Amphipodaoccurin the shallowwater on the inner
side of the lagoon, wherethe chloridecontentat low tide (July 10th) was
6-34 grams per litre. Here are found Gammaracanthus
loricatus,Gammarus
locustaL. var. zaddachi Sexton and Pseudali,brotus
li,toralisKr6yer. Arctic
Terns were seen fishingfor these crustacea-some of the Gammaracanthus
reacheda lengthof 52 cm. Numbersof Ringed Seals (Phoca hispida) inhabit
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V. S. SUMMERHAYESAND C.
S. ELTON
239
the lagoon. They live to a great extent on crustacea, and other bottom
animals.
The followingbirdswereseen in small numbers:
Northern
Eider
Red-throated
Diver
ArcticTern
Kittiwake
FulmarPetrel
Mandt'sGuillemot
GlaucousGull
The diversuse the lagoonfortheirsexual displays,but obtaintheirfoodfrom
the sea, pointingto the probableabsence offishin the lagoon.
The florais of the type usually occurringin the sub-littoralzone in the
Faroes (6) and Iceland (31). Several of the species,e.g. Pylaiella and Phloeospora tortiliscan growin brackishwater or wherefreshwater is discharged
fromstreams(6). Chaetomorpha,
Dictyosiphonand Pylaiella also occur in
Iceland in veryhighpools whichonlyrarelyreceivesalt water,and oftenmust
be brackish,especiallyafterrain (31).
The brackishcharacterofthefaunais shownmoreby theabsenceofcertain
marineforms,than by the presenceofany freshwater
ones. Dead sea-urchins
were seen on the bottomin one place, and planktonbroughtin by the tide
(Copepods,etc.) was in a dyingcondition.The animalsincludeseveralbrackish
forms. Gammaruslocustavar. zaddachiis a form(probablyenvironmental)
of the typical G. locusta. This lives in a salinityof 2 to 6 gramsper litrein
Denmark, but also occurs occasionallyin highersalinitiesthere(61). The
LamellibranchLiocymawas slightlyabnormal,whichmay have been due to
its environment.The Gammaracanthus
loricatus,
however,showedno approach
to the type.ofG. lacustris,the form(also probablyenvironmental)occurring
relictin freshwater in NorthernEurope and in Spitsbergen(48). Richard
Lagoon is of especialinterest,sinceit is one of the largestofits typein Spitsbergen,and may ultimatelybe cut offfromthe sea by land elevation. It
representsan intermediatestage in the processleading to the formationof
relictlakes likethosein Europe.
(b) Inter-tidalZone. We have onlya fewnoteson thisregion.
Alongthe unstableshingleshoreboth outsideand inside RichardLagoon
thereseems to be no rootedvegetation. Huge piles of Laminaria and Fucus
are cast up by the tide. Collembolawerefoundin large numbers:Achorutes
viaticus
, Agrenia bidenticulata,Archisotomabeselsi Packard, and Isotoma
viridis. A. beselsiis confinedto sea shores. Purple Sandpipers were seen
feedingon these animals. A flyFucomyiafrigidaFln. occurson the shoreat
Point Carmichael,and may breedin the seaweed cast up.
As theice had not meltedfromthe edge ofmostofthelagoonwhenwe left,
it was not possibleto see if thereare any salt marshesthere. Mr Mathieson
tells us that thereare large areas of ice in this positionwhichdo not meltin
ordinaryyears,but in the summerof 1922, whichwas exceptionallywarm,
these completelydisappeared. The conditionsseem, tborefore,
to be too unfavourablefortheproductionofwell-marked
inter-tidalcommunities,
although
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to Ecology of Spitsbergenand Bear Island
240 Contributions
GlyceriavilfoideaFr., whichusuallyis commonin salt marshesin Spitsbergen,
is recordedfromthis district(53).
(c) RecentShingle. This occursonlyon the outsideof RichardLagoon. It
is almostbarren,exceptfora fewCollembola,whileArcticTernsand Northern
Eiders nestin small numbers.
3. THE RAISED BEACH SYSTEM.
The partstudiedstretchesfromRichardLagoon almostto PointCarmichael
(see Fig. 4). At its southernend it blendsinto the recentshingle,in the north
into older rocks. Snow preventedobservationsinland. The Beach System
is on an averageabout 30 feetabove sea-level,and formssteep cliffsseawards.
The systemconsistsof parallel beaches,'whichmeet the coast line at a small
angle. The seaward beaches are higherso that most of the surfacedrainage
in heightbetween "ridge" and "low" is
is into the lagoon. The difference
nevermorethan 10 feetand usually less. Most of the beach is composedof
pebbles withdiameterslyingbetween2 and 8 inches. In the cliffface,there
is a sand layerfrom2 to 15 feetin thickness.As the prevailingwindsseemto
be fromthe NorthEast, thissand layeris continuallybeingerodedand carried
on to the shinglesurfacewherethe sand graduallyaccumulates. In the north
very little sand has accumulated,especiallyon the landward beaches. The
plants
vegetationis verysparse,and consistsonlyofveryfewstuntedflowering
and
Lecidea
and lichens. Gyrophora
proboscidea,Rhizocarpongeographicum,
are the mostimportantof the latter.
confluens
On the seaward beaches thereis a littlemoresand, and as a resultthere
is a richer,but still scattered,floraof higherplants and mosses. Saxifraga
and mostfrequentspecies. Also:
is the characteristic
oppositifolia
Phanerogams:
Cerastiumalpinum
Draba alpina
D. subcapitata Simmons
Luzula confusaLindeb.
Oxyriadigyna
Bryophytes:
Dicranoweisiacrispula
Lichens:
Cetrariaislandica
f.
o.
o.
1.
r..
Salix polaris
Saxifragacaespitosa
S. oppositifolia
StellariahumifusaRottb.
r.
f.
a.
f.
Dicranumfuscescensvar. congestumn
The Dicranoweisiaoccursamong the tuftsof Purple Saxifrage. Many lichens
occuron the pebbles. In additionto thosementionedabove thereare Buellia
sorotiaTh.Fr. and Parmelia alpicola Th.Fr.
Amongstonesand plants were:
Collembola:Isotoma viridis
Diamesa poultoniEdw. var. flavipilaEdw.
Diptera:
Sciara tridentataRubs.
Bdella decipiensThor.
Mites:
Thor.
Typhochrestusspetsbergensis
Spiders:
A fewinsectlarvae occurin theplants. The spiderspinsa fewthreadsfrom
etc.
flowerto flowerofSaxifragaoppositifolia,
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V. S. SUMMERIIAYES
ANDC. S. ELTON
241
In damp hollowsthe florais transitionalto the Luzula-Cetrariacommunity
mentionedbelow. Thereis about 1 inch of humus,and the pebbles are small.
Luzula confusaand Cetrariaislandicaoccurheresparingly.The followingalso
occur:
Mosses:
Cynodontiumvirens
f.
Dicranoweisiacrispula
f.
Lichens:
Cetrarianivalis
o.
Microglaenasphinctrinoides
Nyl.
Rhacomitriumlanuginosum
o.
Dicranumfuscescensvar. congestum o
Stereocaulonalpinum
I.a.
The micro-fauna
of some Rhacomitrium
was:
Rotifera: HabrotrochainsignisBryce
Macrotrachelaplicata var. hirundinellaMurray
Mniobia russeolaZelinka (witha parasite)
PleuretraalpinumEhrb.
Tardigrada,Nematoda,Rhizopoda: r.
Thisis a typicaldrymoss. The rotifers,
especiallythelargerones,showgreater
powersof revivalthan the otheranimals(8).
As the sand accumulates,if not too rapidly,a characteristiccommunity
appears-the Luzula-Cetrariacommunity.The threemain plants in this are
Luzula confusa,CetrariahiascensTh.Fr. and C. islandica. There is usually
3 to 8 inchesof sand, of whichthe top layer (Q to 1 inch) is peaty. Therealso
occur:
Phanerogams:
AlsinerubellaWahl.
f.
Cerastiumalpinum
f.
Oxyriadigyna
o.
Brvophytes:
Cynodontiumvirens
var. arcticum
a.
Hypnumuncinatum
var. foeneumHagen.
Lichens:
Microglaenasphinctrinoidesf.
Saxifragacaespitosa
S. oppositifolia
Stellariahumifusa
o.
o.
a.
Lophozia quinquedentata
Polytrichumjuniperinum
a.
StereocaulondenudatumFloerke
var. pulvinataTh.Fr.
o.
The mossesoccurin damperparts,and are moredamp-lovingtypesthan the
previousones. Large masses of NostocpiscinaleKuetz. occurin wet patches.
The two-crustaceouslichensmentionedabove appear to be thefirstcolonisers
on bare ground. Theyare followedbyfruticoseformssuch as Cladoniafurcata,
C. pyxidata,and C. cervicornis
Schaer.
No animals were found among the plants in the Luzula-Cetrariaarea.
Many Collembolaoccur on the surfaceof meltedsnow-waterin hollows.
Near thecoastis a deep layerofsand,whereLuzula and Cetrariaare absent.
There is much bare ground,Saxifraga oppositifoliabeing dominant,while
Silene acaulis L. and Draba subeapitataare abundant. The mossesare similar
to those on the almost bare shingle. The micro-faunaof the mosses is also
similar,and the same may be said of the higherinvertebrates.The flies
extremus
Holmg. and Fucomyiafrigidawereseen on Saxifraga
Catmptocladius
16
Journ.of Ecology XI
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and Bear Island
toEcology of Spitsbergyen
242 Contributions
similarto that on almostbare shingle
is therefore
The community
flowers.
is quitedifferent.
thesubstratum
although
increases,
a distinctchangetakesplace in
Whenthe sand accumulation
thebiota. In placesa seriesofsmalldunes3 to 6 inchesin heightis being
is SaxifragaoppositiTheonlysuccessful
plantundertheseconditions
formed.
stems,keepspace withthedeposition
folia,which,by meansofits creeping
inthehollows
Otherplantsgrowing
ofsandandbindsitintosolidhummocks.
The Oxyriais
Silene,and Drabasubcapitata.
are Oxyria,
Saxifragacaespitosa,
ofanthocyanin.
and brilliant
redin colourdueto theproduction
verydwarfed
BothSileneand Drabaformlargecompacttussockshereand there.Luzula
and Cetrariaare killedby the sand deposition.A grass,possiblyCatabrosa
algida,is abundanthere,butwasnotin flower.
is too rapid,theplantslosetheirhold,
Aftera whilethesanddeposition
and "bloWT-outs"
occur,2 feetofsand beingremovedin someplaces.These
at thedenudedcliff-edge.
becomeundercut
Thehummocks
originate
probably
by thesandblast,and finally
toppleover(16). Silene,owingto its compact
oppositibySaxifraga
is veryresistant.In placesre-colonisation
tussockform,
foliaofsuch"blow-outs"was observed.
Therewas hardlyanyanimallifein theseareas.
Ponds. Thereare one or twosmallpondson thebeaches. Red-throated
mainlyofDesmids(species
Diversnestnearthe edges. The algae consisted
(MerismobutsomeCyanophyceae
ofCosmarium,
andEuastrum),
Staurastrum
Diatoms,
and
were
also
found.
Lemm.)
perelegans
pediaglaucaNaeg.
Lyngbya
Flagellatesand Ciliateswerealso recorded.
4. ROCKY LOWLAND.
ofa rolling
Thisareaconsists
thelagoonand beaches
tractoflandbetween
byfrostaction
andthemountains.Thelargerpartoftheareais differentiated
areasofvarying
intopolygonal
natureand size.
soil." Thetypefoundis thatcalledby Hogbom
(a) Areaswith"polygonal
materials.
(27) "PolygonSoil Type1," and is producedfromheterogeneous
It consistsofpolygonareasofwhichthecentreis composedoffinesiltwhile
toHogbom,
theedgesaredefined
bylinesofstonesofvarioussizes.In addition
Thoroddsen
have dealtwiththe causesof their
(63) and also otherwriters
formation.
onthedrainage
Thenatureofthesepolygons
apparently
varies,depending
wherethedrainageis good),and on therelativehetero(theyare notformed
oftheparticles.In someplacestherimsare composedofsmallshale
geneity
andin others
only,in othersofstonesup to 4 to 6 inchesin diameter,
particles
of
of
thepolygons
diameters
arepresent.The
againblocks 3-feetin diameter
varyusuallyfrom4 to 10 feet. The centreis usuallymuchwetterthanthe
rimowingto retention
of water. Excavationshowedthat the ground-ice,
whichexistseverywhere
at a depthofabout18inches,wasso shapedthatthe
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V. S. SUMMERHAYESAND C(. S. ELTON
243
thaw-watercould neverdrainaway, and probablytheinteriorsofthepolygons
never dry out in summer. The centresare almost always bare of plants
(Plate III, fig.1). The followingwere foundin the centresof manytypesof
polygons,but representfairlyuniformhabitats.
Phanerogams:
Alopecurusalpinus?
Cerastiumalpinum
Draba alpina
Luzula confusa
Bryophyte:
Polytrichumalpinum
r.
Lichens:
Cetrariahiascens
C. islandica
i.f.
i.f.
o.
o.
r.
o.
Saxifragacaespitosa
S. oppositifolia
Stellariahumifusa
o.
o.-f.
0.
Algae:
Nostoc piscinale
0
Solorinacrocea
f.
Stereocaulonalpinum o.-f.
Solorina croceais the most characteristicplant, and oftenis the only species
here.
The edges of the polygons,on the otherhand, oftenbear a luxuriantand
varied flora. Colonisationof the edges was seen in all stagesfromthe appearto a
ance of a few tuftsof moss or small plant of Saxifraga opposit4folia,
2 to 6 incheshighrampartofvegetation.
Apart fromSaxifraga and Salix polaris the lowerplants predominateon
these rims. The followingis a list compiledfrommany types of polygons,
but thereis considerablevariation.
Phanerogams:
Alopecurusalpinus
Cerastiumalpinum
CardaminebellidifoliaL.
Draba alpina
Luzula confusa
Oxyriadigyna
Papaver nudicaule
var. radicatum
1.
o.-f.
1.
r.
0.
1.
0.
Salix polaris
Saxifragacaespitosa
S. cernua
S. nivalis
S. oppositifolia
Silene acaulis
Stellariahumifusa
f.-c.d.
o.-f.
r.-o.
0.
f.-d.
r.
1.
Bryophytes:
r.
Aulacomniumpalustre
o.-a.
A. turgidum
CamptotheciumnitensSchimp. f.-c.d.
r.
Cynodontiumvirens
r.
Dicranoweisiacrispula
Dicranumbonjeani
Braithw.o.-a.
var. juniperifolium
i.f.
Hylocomiumsplendens
r.
HypnumexannulatumGuenb.
H. uncinatum
f.
f.-a.
Polytrichumalpinum
Ptilidiumciliare
r.
RhacomitriumcanescensBrid.
r.
R. lanuginosum
f.-c.d.
Lichens:
Alectoriaochroleuca
Mudd.
Buellia disciformis
var. insignisAL. Gray.
Cetrariahiascens
C. nivalis
Cladonia bellidifolia
C. pyxidata
Cladonia rangiferina
Lecanora epibryon
L. tartarea
Peltigeranialacea Fr.
Psoroma hypnorum
Sphaerophorusglobosus
Stereocaulonalpinum
o.
1.
o.-d.
o.
i.f.
o.
f.
o.
i.f.
r.-o.
o.
r.
o.-f.
and
Nearly all the lichensare on the crestsof the ramparts. Rhacomitrium
seldom occur togetheron the polygonedge. The mosses are
Camptothecium
not bog species; but dry ones are not very commoneither. At the creststhe
vegetationseemsto dryup, and hereoccurthe two speciesof Lecanora,often
16-2
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244 Contributions
to Ecologyof SpitsbergenatndBear Islantd
growingover the otherplants. In Lapland a similargrowthof Lecanoratartareaon dried-outmossesoccurs,althoughon a muchlargerscale (33).
Animallifeon theserampartsis eitherscarceor absent.
(b) Areas without"polygonalsoil." These occur eitherwhere the slope
allows drainageor else wherethe homogeneityofthe particlespreventsdifferentiation.The chiefareas-arethoseat the tops ofthe lowerhills,that worked
in mostdetailbeingone about 150 feethighnamedSileneHill fromtheamount
ofSileneacaulis on its slopes. Most ofthe animaldata are takenfromhere. In
places the groundis rockyand coveredwithfrostdebris. Generallythereis
verylittlesoil (e.g. Silene Hill) althoughin hollowsit may collect. The communityis reallya "fjaeldmark" similarto that on Bear Island, but rather
richerin species. The presence of Dryas octopetalaand the abundance of
Salix polaris in places mark a transitiontowardsthe "Dwarf-shrubHeath"
of othercountries.The completelist (includingSilene Hill) is as follows:
Phanerogams:
Alopecurusalpinus
Alsinerubella
Cerastiumalpinum
Draba alpina
D. alpina var. oblongataRBr.
D. subcapitata
Dryas octopetalaL.
Luzula confusa
Oxyriadigyna
Papaver nudicaulevar. radicatum
PedicularishirsutaL.
Polygonumviviparum
Salix polaris
S. reticulata
Saxifragacaespitosa
S. nivalis
S. oppositifolia
Silene acaulis
Bryophytes:
Cynodontiumvirens
Dicranoweisiacrispula
Dicranumfuscescens
var. congestum
Grimmiaapocarpa
Gymnomitrium
corallioidesNees.
Hylocomiumsplendens
Lichens:
Alectoriaochroleuca
CeraniavermicularisSF.Gray
Cetrariahiascens
C. islandica
C. nivalis
Cladonia rangiferina
C. sylvatica
Lecanora epibryon
L. tartarea
Peltigeramalacea
Sphaerophorusglobosus
Stereocaulonalpinum
r.
r.
o.-f.
r.-o.
r.
l.a.
f.
r.-o.
l.f.
i.f.
f.
a.
r.
f.
r.
a.
o.-a.
o.
o.
o.
o.-f.
r.
r.
o.
o.
r.-a.
l.f.
f.
f.
r.
r.
o.-f.
l.a. on debrisareas
Uf.on drysummits
On sides of hills
Only on drysummits
On dryslopes
Increasestowardsthe summits
HypnumbambergeriSchp.
H. uncinatum
0.
Meesia trichoidesSpruce
o.
Polytrichumalpinum
P. strictumBanks
Rhacomitriumlanuginosum f.
Webera cruda
1.
Dry parts
Increasestowardssummits
Drier parts betweenstones
Dry parts
Cetrarianivalisis characteristicof places wherethe rock is near the surface.
Generallyspeaking,the lowerplants occupya subordinateposition.
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V. S. SUMMERHAYESAND C.
S. ELTON
24
On AberdeenMachar thereare more mosses and lichens. In this area a
few ArcticSkuas and Purple Sandpipersnest. Seton Gordon(21) observed
that the formernestedon hummockswhichwerethe onlyplaces at that time
freefromsnow(see " Skua Hummocks" in the accountofBear Island). A few
foxesalso occurhere.
In the SileneHill regionthereare no Skuas. The followingoccuron Silene
Hill:
r.
Collembola:Isotomaviridis
r.
Xenyllahumicola
o.
extremus
Diptera: Camptocladius
Bohem.o.
megastoma
Limnophora
spetsbergensis f.
Typhochrestus
Spiders:
f.
Bdellagroenlandica
Mites:
f.
B. decipiens
f.
Cytabrevirostris
+
bipilisHerm.
Ceratoppia
+
exilisNic.
Oribatula
f.
notatus
Sphaerozetes
Oligochaeta:HenleabruceiSteph.
PurpleSandpiper
Birds:
SnowBunting
r.
Mammals: ArcticFox
On Sileneflowers
Amongplants
rocks
Understonesand weathering
Nesting
Feeding
fromthe fauna of the beaches in beingricher,especiallyin mites.
This differs
The two species of Bdella do not seem to live in the same kind of habitat.
Numbersof young B. decipienswere found,each in a sphericalspiny case,
on the under side of stones,and in some cases were just emerging.All the
oribatidmites(the last threegivenabove) are vegetarian,and Mr Hull states
lays 7 or 8 pale orangeeggs,
food. Typhochrestus
that theyprefercryptogamic
whichare fastenedto the underside of a stonein a cocoon of whitethreads.
There are several recordsof mites,thoughnot species foundby us, "in the
" (34). But clearlymostofthe mitesmust
spetsbergensis
nestof Typhochrestus
be vegetarian,as therewould not otherwisebe enoughfood to supportthe
numberof spiderswhichoccur. The foodrelationshereare muchthe same as
those shownin the diagramforBear Island. Otheranimals were:
viaticus
Achorutes
Collembola:
Diptera: Diamesapoultoni
Holnmg.
consobrinus
Orthocladius
Vegetarian
Thor.
reticulata
Hermannia
Mites:
The two flieswereflyingabout over almostcompletelysnow-coveredcountry
in the firstdays of July.
(c) Moss-mat. Near Vogel Hoek thereis a large flat mossyplain which
may be included in this community,though it shows transitionsto Moss
Heath. The plants here are probablylittle affectedby the sea-birds. There
are practicallyno higherplants.
Bryophytes:
byssacea
Cephaloziella
Brid. f.
Dicranum
groenlandicum
Hedw.
D. scoparium
Boul.
var.spadiceum
Lophoziaalpestris
a.
uncinatum
Hypnum
o.
lanuginosum
Rhacomitrium
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246 Coontributions
to -Ecologyof Spitsbergenand Bear Island
Lichens:
Cetrariahiascens
C. islandica
Cladonia lepidotaNyl.
C. rangiformis
a.
a.
o.
f.
Lecanora tartarea
Microglaenasphinctrinoides
Xanthoriaparietina
Wetmossareasprobablyoccur,butweresnow-covered.
ofsmallpondshereand
(d) Freshwater
Communities.
Therearea number
therealongthe coursesof the streams.Thesewerenotworkedforanimals
orplantssinceit wasnecessary
notto disturb
theRed-throated
Diversnesting
there,whichwereunderobservation.Thisspeciesis limitedin its choiceof
siteby thefactthatit cannotwalkproperly
nesting
on land,and thenestis
hardlyevermorethan 10 feetfromthe water'sedge (21). The eggswere
severaltimesrobbedby Skuas. Thereis nevermorethanonepairofDivers
on eachpond.
ofsnow-water
The streamsat thistimeweremostlyrushing
torrents
and
devoidoflife.
(e) ErraticBouldersand Rocks. Mostof thesesupporta floraeitherin
cracksor on the surfaceof the stone. The crevicefloraconsistsusuallyof
speciesfromthesurrounding
"fjaeldmark."
On the rocksurfacesthe floraconsistsmainlyoflichens.A fewmosses
occur,buttheyarenotimportant.
Themorecommon
lichens,
ofall biological
types,are thefollowing:
Alectorianigricans
1.
Gyrophoraproboscidea f.
Lecidea lapicida Ach.
var. declinansNyl.
Lecanora austeraNyl.
RhizocarpongeminatumKoerb.
R. geographicum
Xanthoriaparietina
f.
f.
lf.
is particularly
The Xanthoria
characteristic
oflargeisolatedboulders.
5. UPLAND REGION.
herereacha heightofabout 1600feet(see Fig. 4), with
The mountains
steepslopes,but theremay be ratherflatplateauxof limitedextent.No
" polygonal
soils"wereseenin theregion.
screesoccurinmanyplaces. Theparticles
areusually
(a) Screes.Ordinary
ratherlargeas therockis a hardquartziteorgneiss.Thescreesarebeingfed
fromabove, and thus plantsare sparseor absent. The main
continually
as inall unstableandseverelocalities,
Luzula
species,
is Saxifraga
oppositifolia;
Rhacomitrium
confusa,
canescens
var.ericoides
B. andS.,andvariouscrustaceous
lichensarealso present.
Theanimalsofa screein themouthofGlenMackenzie(300-400ft)were:
Collembola:Achorutesviaticus
Xenylla humicola
Spiders:
LeptyphantessobriusThor.
Mites:
Bdella decipiens
Sphaerozetesnotatus
Scutovertexlineatus
Cyta brevirostris
Birds:
Snow Bunting. Nesting
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V. S. SUMMERHAYES AND
C. S. ELTON
247
canescensvar. ericoideswere a rotifer,Macrotrachela
In some Rhacomitrium
plicata var. hirundinella,and a few nematodes,tardigradesand protozoa.
sobriusis a fast runnerand occurs almost only upon unstable
Leptyphantes
and otherflat-lowland
moiintainslopes in Spitsbergen,whereTyphochrestus
150
feet,and Mr Huxley found
speciesare absent. It came on one hillhereat
it togetherwithIsotomaviridis,a flyand the Snow Bunting,on a stabilised
slope on Lord Stairs' Heightsat 1350 feet(see below).
(b) StabilisedSlopes. These are usuallyscreesoffineparticles,whichhave
been stabilised by plants. Even now movementtakes place as a result of
"solifluction." A large area of soil, with the layer of vegetationon it, had
slipped down recentlyin one place on Lord Stairs' Heights,and doubtless
theselandslipsare not uncommon.
The slopesare usuallycoveredby a close herbagewhichconsistsof dicoty" (44).
ledonson southaspects,and are identicalwithNathorst's" sluttningar
The luxurianceof the vegetationis doubtless due to the intenseinsolation
received,as the slopes are nearlyperpendicularto the sun's rays. This effect
has been noticed in arctic countriesby many writers. Kihlman(33) gives
figuresshowingthe extraordinaryheating-upwhich the groundexperiences,
soon afterthe snow has melted,and withground-icenear the surface. This is
greatestaccordingto him on the south side of small hummocks;how much
moreso mustit be on steepslopes! Otherfactorsare the good drainageofthe
soil and the earlyremovalof the snow. These slopes,however,are very dry
in late summer,but would never completelydry up owingto the meltingof
the groundice. The followingoccurupon such slopes:
Phanerogams:
Cerastiumalpinum
Draba alpina
D. wahlenbergiiHartm.
Luzula confusa
Papaver nudicaulevar. radicatum
Polygonumviviparum
Bryophytes:
Hypnumuncinatum
Lichens:
Cetrariahiascens
C. nivalis
f.
r.
f.
f.
f.
Saxifragacaespitosa
S. cernua
S. hieraciifoliaW. & K.
S. nivalis
S. oppositifolia
Salix polaris
Silene acaulis
f.
o.
o.
r.-o.
f.
o.-f.
o.-f.
r.
o.
o.
Cladonia pyxidata
r.
plants,many
A strikingpointis thelack ofany dominantamongthe flowering
common.
of thembeing equally
therewere
On the northslope,whichwas thenstillpartiallysnow-covered,
manymoremosses,and thisagreeswithotherwriters'observations.
In someplaces a ratherdampertypeofcommunityoccurs,includingmany
plantsbeingless important.In additionto someof those
mosses,theflowering
givenabove therewere: Alopecurtisalpinus
Cardaminebellidifolia
o.
Lycopodiumselago L.
f.
Oxyriadigyna
Ranunculussulphureus l.a.
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Bear lsland
to Ecologyof Spvitsbergen-and
248 Contributions
Nearlyall ofthesedenotea damperhabitat.Lowerplantsinclude:
Bryophytes:
Aulacomniumturgidum
Dicranoweisiacrispula
DicranumelongatumSchleich.
D. fuscescensvar. congestum
HypnumrevolutumLindb.
Lichens:
Alectoriaochroleuca
a.
Cetrariaislandica
Cladonia furcata
Hypnumuncinatum
Polytrichumalpinum
P. strictum
Rhacomitriumlanuginosum
Cladonia rangiferina
Lecanora tartarea
Sphaerophorusglobosus
The mossflorais alliedto thaton thepolygonrims. The lichensindicatea
somewhat
dryerhabitatthantheravines.
Thefaunaofthestabilisedslopeswas notstudiedcarefully.
(c) WetRavines.Theseare moreofthenatureofdrainagehollowsthan
clefts.The groundis soddenwithwaterwhichtricklesdownamongthick,
oflivermosshags,flowering
foot-high
plantsbeingalmostabsent.A number
withthe mosses. Curiously
wortsoccurintertwined
enoughthereare also
ofthe
quitea numberoflichens,somein thewetparts,someon thesummits
that
the
much
whole
becomes
drier
at a
It
is
hags.
veryprobable
region
are themoreimportant
late periodin theseason. The following
species:
Phanerogams:
Oxyriadigyna
Bryophytes:
Aulacomniumpalustre A
Dicranumelongatum
Hypnumuncinatum
Polytrichumjuniperinum)
Aulacomniumturgidum
Cephalozia bicuspidata
HypnumcallichroumBrid.
H. stramineum
Lichens:
Cetrariahiascens
Peltigeraaphtosa Willd.
P. malacea
r.
d
f.
f.
f.
Lophozia alpestris
L. attenuataDum.
1.. qiiinquodenit-i
() !i o..1it i:. w ( IilyseumB. & S.
Polytrichumalpinum
f.
Ptilidiumciliare
SphenobolusminutusSteph. f.
Cladonia furcatavar. surrectaFlk.
C. pyxidata
C. lepidota
GENERAL CONSIDERATIONS.
The vegetationof the area is characterised
by the greatimportance
of
in manycommunities.
In theArctic,
cryptogams
owingto thefog,thecoastal
areas are muchless favourableforplantlifethan are the clearer,sunnier
ofthefjords.Thedwarf-shrub
interiors
Cassiopetetragona
Don. doesnotseem
to occuron PrinceCharlesForeland,and thisis partlyin agreement
with
inNorthEast Greenland.ThereCassiopeforms
conditions
heathsinlandonly,
whileon thecoastsit occursscattered.It is also possible,thatCassiopehas
notyetbeenableto crossForelandSound,and as thewestcoastis oftenopen
in winter,
due to violentstorms,
the seedsmaynothave beenblownacross
owingto theabsenceofice. See (28, 50).
Arcticanimalsworkalmostthe whole24 hoursowingto theabsenceof
darkness.Feilden(17) states,however,
thatthereis a slightamountofrest
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ANDC. S. ELTON
V. S. SUMMERHAYES
249
among the birdsat midnight.The temperatureis usuallylowerat night,and
thismustaffecttheactivitiesof invertebrates.Johansen(29) states that bees
in Greenlandworkall night,but that thereare fewerat midnight.He says
moths,which is unusual for
and night-flying
also that thereare day-flying
arcticanimals. The absenceof darknessaccountsforthelack ofnightworking
"
animals (except the mothsmentionedabove), and thus the "relay systenm
of day and nightfornms
whichenables more animals to occupy any area in
lowerlatitudes,is absent in the Arctic. This is one cause of the poornessin
species of the fauna. Othercauses in Spitsbergenare geographicalisolation
and severeconditions.
B. CAPE BOHEMAN REGION.
This districtnowhereriseshigherthan 50 feetabove sea level, and can be
divided into two parts. The northernpart consistsof flat boggy country,
wherethe underlyingrockneverappears at the surface. In the southernpart,
the soil is almost always veryshallow,and thereis a seriesof rock outcrops.
The rocksare sandstonesand slates deficientin lime,and dip at a slightangle,
in places formingrock pavements. There are a numberof ponds scattered
whichare nowhere
overthe wholearea. Aroundthe coast thereare oftencliffs,
morethan 30 feethigh.
Dr G. J. van Oordtof Utrechtspent most of the summerin the vicinity.
We are much indebtedto him forinformationabout the birds of the neighbourhood.
The followingcommunitiescan be distinguished:
(a) Land Communities.
1. Inter-tidalZone.
2. MaritimeCommunities.
3. "Fjaeldmark."
(a) Dryas.
(b) Othertypes.
4. Heath.
5. Rock Communities.
(a) Surface.
(b) Crevices.
6. StreamsideCommunities.
7. Wet Tundra.
(a) Moss-SalixBog.
(b) Pond and StreamMarginalCommunities.
(b) FreshwaterCommunities.
1. Ponds.
(a) Permanent.
(b) Temporary.
2. Streams.
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to Ecology of Spitsbergenand Bear Islaud
2,50 Contributions
(a) Land Communities.
1. INTER-TIDAL ZONE.
This may consistof shingle(not common)or of rocks. In the case of the
lifeis entirelylacking.On therocks,especially
shingle,owingto theinstability,
neartheDutch coal mine,thereis a richalgal vegetation. The followingoccur:
a.
Fucus evanescensAg.
ChaetopterisplumosusKuetz.
Phloeosporatortilis
Hydroida: GonothyraealoveniAllman
Mollusca: Littorinarudisvar. groenlandicaUk6.
Crustacea: Numerousamphipoda(Gammarus,etc.)
Purple Sandpiper
Birds:
Feeding
Feeding
GreyPhalarope
Algae:
2. MARITIME COMMUNITIES.
There is little special life on these areas. Mandt's Guillemotand Arctic
Tern nestin small numbersalong the cliffs,and the lattertogetherwithgeese
and the NorthernEider are found on small rockyislands offthe coast (see
lives among shaly rocks
"Edinburgh Island"). The mite Bdella groenlandtca
on the cliffs. In several places the usual "fjaeldmark" plant community
changesin the vicinityof the sea. Stellariahumifusais muchcommonerhere,
and is usually recordedfromsuch places in arcticcountries.Saxifragafiagellaris Willd.also occursin thisregion,but Dryas is absentin manyplaces.
3.
"FJAELDMARK."
fromthat on Bear Island and Prince Charles
(a) This communitydiffers
Foreland in the abundance of Dryas octopetala,in many places this being
dominant. However, it nowhereformsa completeplant covering,the soil
being usually bare betweenthe plant cushions. The groundis very gravelly
withlittlesoil. The communitymay be comparedwith Cleve's (9) "Polster
besides Dryas (Salix
Heide" although there is only one other dwarf-shrub
polaris). The Dryas plantsformclose cushions,as do mostofthe otherplants.
is the nextcommonestplant. The Dryas "fjaeldmark"
Saxifragaoppositifolia
occurs on dry exposed soils, and this distinguishesit fromthe other type
describedbelow.
The speciesare:
Phanerogams:
Cerastiumalpinum
Draba alpina withvar. oblongata
D. wahlenbergii
Dryas octopetala
Luzula confusa
Oxyriadigyna
Bryophytes:
Hylocomiumsplendens
Lichens:
Cetrarianivalis
o.
f.
f.
a.
o.
r.
Pedicularishirsuta
Polygonumviviparum
Salix polaris
Saxifragacaespitosa
S. oppositifolia
Silene acaulis
r.
f.
f.
r.
f.
o.
Lecanora tartarea
a.
f.
o.
In the open partsinvertebrateanimal lifeis almostabsent.
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V. S. SUMMERITAYES
ANDC. S. ELTON
251-
on slightlydamper
(b) Thisis a moremixedtypeof" fjaeldmark" occurring
slopes,wherethereis morereal soil. Dryas octopetalais absent. The species
are those usually growingin damper places than the plants in the last list.
The plant coveringis never continuous. The communitygrades into the
streamsidecommunitybelow and into the Dryas communityabove. The
followingare present:
Phanerogams:
Luzula confusa
Equally
Salix polaris
Equndal
Saxffragaoppositfolia aOxyria
S. caespitosa
o.-f.
r.
S. hieraciifolia
Lichens:
Cetrariaislandica
l.a.
Saxifraganivalis
Cerastiumalpinum
digyna
Silene acaulis
o.
r.
O.
r.
Stereocaulonalpinum 1f.
4. HEATH.
Heath is known frommost arctic countriesand has been describedby
Warming(67), Cleve (9) and others. Generallythe dwarf-shrubs
formingthe
heathare ofmanyspecies,as in Greenland.In the Cape Bohemanregionthere
is only one species, namely Cassiope tetragona.This growsto a heightof a
footor so, and formsa thickcarpetoverthe ground. The heathin theregionis
formslargeheathsin East
developedin shelteredlocalities. Cassiopetetragona
Greenland(15, 24), but is thereassociatedwithEmpetrum,
Cassiopehypnoides
and Vacciniumuliginosum. In East GreenlandCassiopetetragona
onlyoccurs
wherethereis a completesnow-covering
duringthe winterand spring(38),
and its distributionat Cape Boheman can be easily explained on the same
basis. Withthe Cassiope,but in verysmall numbers,are:
Papaver nudicaulevar. radicatum
Saxifragaoppositifolia
Equisetum variegatum
Dicranoweisiacrispula
o.
o.
r.
f.
The Cassiope heath is very dry and poor in animal life. In one area the
mite Sphaerozetesnotatusand a flyLimnophorahyperborea
Bohem. were the
onlyanimalsfoundaftermuchsearch.
The birds of the dry-regioncommunitiesare dealt with together. The
followingbirdsnest here:
Purple Sandpiper
GreyPhalarope
ArcticSkua
NorthernEider
King Eider (SomateriaspectabilisL.)
f.
f.
r.
o.
o.
Pink-footedGeese and Brent Geese feed here and in otherparts of the Cape
Bohemanregion. The Skuas attack and eat young Eiders,and even Purple
Sandpipers(64). The GreyPhalarope feeds both in freshwater and at sea.
The stomachof one containedremainsof flies,etc. One fromAdvent Bay
and another205 largechironomidflylarvae in the
(July,1921) had overfifty,
stomach. One fromthe same place obtainedby MrJourdainin 1922 contained
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to Ecologyof Spitsbergenand Bear Island
252 Contributions
flylarvaeand moss,whiletwo othershad remainsof marinecrustaceainfeedson landand on
locusta.ThePurpleSandpiper
cludingyoungGammarus
oftwobirdsfromCapeBohemanhad in
theshoreat lowtide. Thestomachs
fourand six spiders,anotherthreefliesand someplant
themrespectively
and remainsofseaweed.
threetinymarinegasteropods
remains,
and a fourth
5. ROCKCOMMUNITIES.
Theseare presenton the rockoutcrops.The majorityof the outcrops
witha fewshallowcracksand deslopingrockpavements
consistofgelntly
and
has had a muchgreater
effect,
weathering
pressions.At theescarpments
manyof whichremainalmost
therockhas beensplitintolargefragments,
seriesofparallelcracksis thusformed,
manyofthem
in situ. A complicated
intotwodivisions:(a) Rock
fallnaturally
4 to 6 feetdeep. The communities
and (b) CreviceCommunities.
SurfaceCommunities,
the
flora
of
rocksurfacesis verypoor. Thereis none of the
(a) The
and otherlichensas on Bear Island and Prince
luxurianceof crustaceous
CharlesForeland.Thisis doubtlessboundup withthemuchdrierclimateat
CapeBoheman.
(b) In the smallercracksand in slighthollowson the pavementa little
soil collects,and this can supportstuntedspecimensof the "fjaeldmark"
plants. However,in the deepercracksa morespecialfloradevelops.The
chieffactorhereis thesmallamoulitoflightavailable. Sincethecracksmay
be 6 feetdeep,are usuallyundera footwideand have verticalsides,it is
patentthatwiththelowaltitudeofthesunin theselatitudesverylittlelight
can enter.Thisis shownby the higherplants,whichare muchdrawnout
also is muchdecreased.Plantsof
The production
offlowers
and attenuated.
Oxyriacollectedherewerequitegreen,whereasthoseon the "fjaeldmark"
Mossesare an important
partofthe
outsidewerereddishwithanthocyanin.
whileas on BearIsland,thereareseveralliverworts.
vegetation,
Phanerogams:
Cerastiumalpinum
Oxyria digyna
Ranunculussulphureus
Bryophytes:
Blepharostomatrichophyllum
Hylocomiumsplendensvar. gracilis
Hypnumstellatum
H. uncinatum
Lophozia longidens
r.
o.
o.
f.
Saxifraga cernua
o.
S. oppositifolia
Stellaria longipesGoldie r.
f.
c.d.
Lophozia ventricosa
Polytrichumalpinum
Pseudoleskeacatenulata f.
Scapania curta
c.d.
Tinmia austriaca
f.
Saxifragacernuais theonlyhigherplantwhichis at all normalhere. It has
in somewhat
frommanycountries
similarhabitats.
beenrecorded
in boggy
but wereoftenseenfeeding
SnowBuntingsnestin thecrevices,
places. Stomachsoffivebirdsfromhereeachhad abouta dozenflypupaein
them,whileonehad also remainsofadultflies.
fallenblockswere:
Amongplantsoutside-the
Collembola: Isotoma viridis
Immature
Spider:
sp.
Oligoehaete: Mesenchytraeus
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V. S. SUMMERHAYES AND C. S. ELTON
253
Between the block collectionsand the Cassiope heath is oftenmuch moss in
whichare manyplants of Ranunculussulphureus.
6. STREAMSIDE COMMUNITIES.
Thereare manysmall streams,and some ofthemare probablyrunningall
throughthe season. Some, however,which drain temporaryponds, dry up
in the late summer. The substratumis much damper than any described
previously. In some places the banks may be floodedby large amounts of
snow-water,but this is not general. Many phanerogamsoccur here, often
formingthe main vegetation. Oxyria digyna is the commonestplant, especiallywherethe banks are steeper,and inundationrarer. Anothercharacteristicplant is Ranunculus nivalis L. togetherwith R. sulphureus. Salix
polaris and Luzula confusaare also common. The communitycorresponds
withthat describedby Cleve (9) in arcticSweden. Alongone streamtwozones
werevisible. The one nearestthe streamconsistedofAlopecurusalpinus,that
on slightlyhighergroundof Poa alpina L. and the two speciesof Ranunculus.
Lundager(38) recordsan almost identicalarrangementof zones in NorthEast Greenland. In additionthe followingare found:
Phanerogams:
Alsinerubella
Ranunculus
pygmaeus
cernua
Saxifraga
r.
1.
i.f.
Saxifraga
hieraciifoliar.
r.
S. rivularis
Bryophytes:
Brachythecium
salebrosum
var.arcticum
Berggr.
nitens
Camptothecium
Polytrichum
alpinum
P. juniperinum
7. WET TUNDRA.
This occupiesmost of the northernhalfofthe region:it also occursaround
the ponds and in the valleys betweenthe outcropsin the southernpart. In
nearlyall cases it is formedof mossestogetherwithSalix polaris. Two communitiescan be distinguished.
(a) Moss-Salix Bog (Plate II, fig.1). The wateris stagnant,and in many
places formspools. In the bog are numeroushummocksformedby the Salix,
the spaces between its branches beilngfilledwith interwovenmosses. The
followinghigherplants occur,mostlyon the summitsof the hummocks:
o.
Alopecurus
alpinus
Cardamine
bellidifolia
Drabaalpina
D. wahlenbergii
o.
scheuchzeri
Eriophorum
Hoppe. o.
o.
Pedicularis
hirsuta
1.
Petasitesfrigida
Fr.
Ranunculus
sulphureusf.
c.d.
Salixpolaris
Saxifraga
hieraciifolia r.
The Alopecurusand Eriophorumare scatteredabout betweenthe hummocks,
and forma regiontransitionalto the dampertype of "fjaeldmark."
A numberof lichensalso occur on the hummocks:
Alectoria
ochroleuca
Ceraniavermicularis
Cetraria
nivalis
Cladoniarangiferina
Stereocaulon
alpinum
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254 Contributions
to Ecology of Spitsbergen
and Bear Island
Theseshowthatthe tops of the hummocks
dryup in summer.Thereis a
markedzonationofthe mosseson thehummocks.The following
showsthe
typeofzonation,
starting
fromabove:
1. Camptotheciumnitens
Hypnumuncinatum
2. Brachytheciumsalebrosumvar. arcticum
Cynodontiumvirensand var. arcticum
Polytrichumalpinum
3. Paludella squarrosaand others
4. Hypnumcordifolium,
in lowestpools
H. giganteumSchimp.
H. stramineum
Ranunculushyperboreus
occuramongthemosses,especially
A certainnumberofliverworts
on the
Salix tussocks:
Blepharostomatrichophyllum
CephaloziellabifidaSchiffn.
and var. erosaLophozia kunzeana Evans
Lophozia quinquedentata f.
and f. turgidaLindb. f.
OdontoschismasphagniDum.
withthe "Myrar" in
These bogs agreeverycloselyin moss constitution
Iceland(25), and withthosein East Greenland
is a
(14). The following
generallistofmossesfortheCape Bohemanbogs:
Aulacomniumpalustre
A. turgidum
Brachytheciumsalebrosum
a.
var. arcticum
Bryumobtusifolium
f.
Camptotheciumnitens
Cincidiumstygium
Cynodontiumvirens
and var. arcticum
Dicranumfuscescens
f.
Hypnumbrevifolium
H. cordifolium
H. intermedium
Hypnumsarmentosum
H. scorpioidesL.
H. stellatum
H. stramineum f.
H. revolvensSw.
H. uncinatum
Meesia triquetra
Mniumaffinevar. integrifolium
Orthothecium
chryseum
Paludella squarrosa
Polytrichumalpinum
P. juniperinum
ofthewetterparts.
ThevariousspeciesofHypnumare mostcharacteristic
All thepondsand especially
(b) Pondand StreamMarginalCommunity.
thepermanent
oneshavea thickmossmargin.Thesameis also trueofsome
occursin damperplacesthan
streams.In thelattercasethemosscommunity
thephanerogamic
onealreadydescribed.Ranunculus
nivalisand sulphureus,
L. f. tetrandrum
Lund.occur
Oxyriadigynaand Chrysosplentium
alternifolium
in thesecarpets.The latterwas onlyoncefound,on theedgeofa largepermanentpond.
to thatoftheboggyareas. In additionSphagnum
Themossflorais similar
Warnst.was foundin a fewplaceson the
Wils.var. concinnum
fimbriatum
ofSphagnum
bankoftheabove-mentioned
pond. Theabsenceorrelative
rarity
has beennotedbefore(5, etc.). It is well-known
inmoss-bogs
inSpitsbergen
thatinbogsintheArcticSphagnaaremuchlessimportant
thanin temperate
did we findany
countries.Onlyin one place (AdventBay) on Spitsbergen
and evenhereit is ratherlocal. In thevalleybetweeni
amountofSphagnum,
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JOURNALOF ECOLOGY
VOL. XI, PLATE II
Photo. C. S. Elton
FIG.
I.
etc., and mosses. A temporary
Cape Boheman. Wet Tundra of Salix, Alojr5ecurus,
pooi withxvetmossvegetationis in the middledistance.
Photo. J. Walton
on the right,unsiltedbeach with
Klaas Bi'llen Bay (Bruce City). In foreground
Dryas,etc.; on theleftPond VI I withmossymargin. Behindthisthe CampbellRange.
FIG. 2.
SUMMERHAYESAND ELTON-EcOLOGY
OF SPITSBERGEN
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AND BEAR ISLAND.
V. S. SUMMERHAYESAND C. S. ELTON
255
the two LongyearCity minesmoss-bogsoccurin various places. In addition
to the usual bog mossesgivenabove, the followingspeciesofSphagnumoccur:
Sphagnumacutifoliumvar. subnitensRuss. & Warnst.
S. subsecundumvar. contortumSchimp.
Warnst.
S. squarrosumvar. semisquarrosum
In agreementwiththe rarityof Sphagnumthereis verylittlepeat produced
in Spitsbergen. A small amount was seen at Cape Boheman, but none in
otherbogs. In most cases the mosses grow directlyon the underlyingsoil.
Nathorst(43) mentionsthe bog in ReindeerValleynear Cape Thordsen,where
thereis about 10 inchesofpeat. It is composedofHypnumspp. and theupper
layerscontainremainsof Saltx polaris. It seems as thoughthis was formed
froma bog similarto thosedescribedabove.
The zonation on the pond sides usually resemblesthat on the bog hummocks. The lowestlayer,consistingof speciesof Hypnum(e.g. H. scorpioides
var. fontinaloides),was observed oftencompletelysuband H. sarmentosum
mergedand possiblyis always so.
Thereare oftenmany Collembolaon the surfaceof waterin the bogs and
at the edges of the ponds. Amongtheseis Agreniabidenticulata.In mosses
virens,Hypnumbrevifolium
(includingthe Sphagnumgivenand Cynodonttium
froma pond marginwere:
and H. cordtfolium)
Rotifera: Adinetavaga Davis
Mniobiarusseola
RotifersordidusWestern
also Tardigradesand Nematodes
Thereare a numberof fliesto be seen in the boggyregion:
Orthocladiusconsobrinus
ChironomuslugubrisZett.
PsectrocladiusborealisKieff.
Cricotopusbasalis Stalg.
Metriocnemusursinus
Camptocladiuscurvinervisvar. polarisK.
Phalaropesand Snow Buntingsact as a big checkon the aquatic flies.
Three species of birds nest here-Grey Phalarope (f.), King Eider (r.),
and Long-tailedDuck (r.). The last two feedmainlyat sea, but also on fresh
water.
(b) Freshwater Communities.
1. PONDS.
These are scatteredabout, some in the hollowsbetweenthe rock ridges,
othersin the flatboggyexpanse in the north. The most strikingfeatureat
Cape Boheman is the large numberof temporarypools whichdry up before
the end of the summer. Dr van Oordtkindlyinformsus that Pond 12 dried
up completelybeforethe end of the season. Many otherponds had muchless
this
waterin them,and wouldcertainlyhave driedup also. The biota confirms
idea. The frequencyof thesetemporaryponds is due to the geologicalformation. The groundbeing rocky,hollowsare formedwhichhold meltedsnowgroundwater. Sincethe rockis closeto thesurface,thereis nottheunderlying
ice whichis foundon shinglebeaches as at Klaas Billen Bay, and thus the
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256 Contributtions
toEcology of Spitsbergen
and Bear Island
pondsare notsuppliedwithwaterby thegradualmelting
ofthisice during
thesummer.Thosesmallpondsat Klaas BillenBay whichonewouldexpect
to dryup, do notdo so, and theycontainthesamefaunaas thelargerpermanentpondshere.However,thewaterofsmallpondsis hotterin summer,
andtheanimalsaretherefore
ableto complete
theirseasonalcycleina shorter
time(48). Thismakestheseasonvirtually
shorter
in a largepond,and tends
to equalisetheconditions
in smalland largeones. About20 pondswereexaminedat Cape Boheman,buttypicalexamplesonlyare given.
(a) Permanent
Ponds. Thereare manylargeshallow(1-2 feet)ponds,
usuallywitha definite
bankofmosses,etc. Thesepondscontaina richalgal
rockis siliceousand poorin lime,thealgal flora
flora.Sincethesurrounding
richin Desmidsmainlyofthe generaCosmarium,
is extraordinarily
Stauraand C. construmand Euastrum.The largetuberculate
Cosmarium
biretum
are characteristic.
in permanent
spersum
The following
occurtypically
pools
and werenotfoundin thetemporary
ones:
Chlorophyceae: CosmariumbiretumBreb.
C. conspersumRalfs. var. latum W. & G. S. West
C. impressulumElfv.
C. punctulatumBreb. var. subpunctulatumNordst.
Euastrumspp.
Cyanophyceae: Lyngbyaperelegans
ofspeciesoccurwhichare also foundin thetemporary
A number
pools:
Chlorophyceae: Cosmariumcrenatumvar. bicrenatumNordst.
C. holmiensevar. integrumLund.
C. ochthodesNordst.
C. protumidumNordst.
C. quadratumRalfs.
Staurastrumbieneanum
S. pachyrhynchum
Nordst.
S. subbrebissoniiSchmidlevar. hexagonumSutwinski
PandorinamorumBory.
Cyanophyceae: ChroococcusturgidusNaeg.
OocystissolitariaWittr.
SynechococcusaeruginosusNaeg.
Pond20 is a goodexampleofthistypo. Thebottomis sandy. Thefollowing
thelasttwo:
oftenmingle,
subdivisions
especially
Crustacea:
Plankton:
Daphnia pulex De Geer
CyclopscrassicaudisSars.
Amongmoss,etc.: Chydorussphaericus
Cyclopscrassicaudis
On bottommud: Lepidurusarcticus
Macrothrixhirsuticornis
Norman& Brady
Eucyprisarctica
Candona rectangulataAlm.
and feedon diatomsand otheralgae,while
All theseanimalsare vegetarian
formsare also scavengers.
Thereis a richfaunaof rotithe bottom-living
fers,etc. AmongtheprotozoawerelargecoloniesofthegreenciliateOphrywereseenfeeding.Chironomid
dium,uponwhichyoungLeptidurus
flylarvae
abundant.
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V. S. SUMMERIIAYES
ANDC. S. ELTON
257
Ponds 6, 7, 10, 19 are of the same type. The Copepod Maraenbiotusbrucei
occurs sometimes. The natureof the bottomhas an importanteffecton the
fauna. Pond 14 has a stony bottomand very little plant life,and contains
only Daphnia pulex and Chydorussphaericus. Close to it is Pond 15 which
has a mud bottom,on whichare Lepidurus,Eucyprisand Candona. A rather
different
type of permanentpond is that whichhas a streamflowingthrough.
Pond 1 is about 35 feetacrossand a footdeep. One edge has mosses,Ranunculus hyperboreus,
Eriophorum,etc. Flies (Cricotopusbasalis and others)are
abundant near the water, and larvae and pupae in it. There is a strearn
flowingthrough,and therefore
planktoncrustaceaare almostabsent. On the
bottomand edge are Lepidurusarcticus,Chydorussphaericus,Cyclopscrassicaudis,EucyprisglactalisSars., also rhabdocoelflatworms,
rotifers,
etc. Pond 2
is also of thistype.
(b) TemporaryPonds (Plate II, fig. 1). The algal floraof these ponds is
very rich,and consistsmainly of Desmids. Unlike the other type of pond
these are characterisedby the presenceof Staurastrummegalonotum
Nordst.
and S. oligocanthum
Breb. which are absent in the permanentpools. The
followingspecies of algae are also typical of these temporarypools, and are
absent in the permanentones:
Chlorophyceae: CosmariumarctoumNordst.
C. bioculatumBreb.
C. cyclicumLund. var. aroticumNordst.
C. ochthodesvar. amoebumW. & G. S. West
Euastrumspp. (not the same ones as in the permanenit
ponds)
obesumWest
Nephrocytium
Staurastrumpolymorphum
Breb.
Eudorina elegansEhr.
GloeocystisinfusionemW. & G. S. West
Cyanophyceae: Aphanocapsa grevilleiRabenh.
AphanothecemicroscopicaNaeg.
The faunaofthesevaries,and mustdependto someextenton chancedispersal.
They are not characterisedby any peculiarcrustacea,but by the absence of
Lepidurus,Daphnia and Macrothrix,and usually of flylarvae. We should
expect to findthe richestfauna in those which have been formedby the
encroachmentof plants on a formerly
largerpond. Ponds 3 and 4 appear to
be of this type.
Pond 3. This is about 10 feet in diameterand a few inches deep. The
and a grass. Outside
vegetationat the edge consistsof Hypnumcordifolium
is a broadlayerofH. brevifolium
and H. stramtineum.
mixedwithH. cordifolium
is lacking
On the outside of this again is a ratherdrierarea. H. cordtfolium
here,but Cynodontinum
virenstakes its place. The "fjaeldmark" lies beyond
this. Of the crustaceaAcroperusharpaeBaird has not previouslybeen found
in Spitsbergen.
Pond 4. Here the pond has reacheda furtherstage,being5 feetwideaild
an inch or two deep. In this are foundRotifera:MetopidialepadellaEhrb.,
and others; Protozoa: Peridinium;Nematodes; Tardigrades. The following
Journ.of Ecology XI
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17
toEcologyof Spitsbergenand Bear Island
258 Contriibutions
stagesin
pools showingdifferent
is a table (Table II) of all the temporary
up:
dryillg
TABLE II.
Pond No.
Species
Chydorussphaericus
17
4
11
12
13
22
18
x
21
x
3
x
Cyclopscrassicaudis
x
x
x
x
x
x
x
x
x
x
x
Acroperus harpae
Maraenbiotus brucei
Candona rectangulata
Eucypris glacialis
-
-
x
x
-
x
x
x
x
x
in boggyland. Pond 11 resembles
Ponds11,12 and 13 are closetogether
and a grassoutside)
in centreand H. brevifolium
cordifolium
Pond3 (Hypnum
Metopidialepadellaand EuchlanisdeflexaGosse.
and containsthe rotifers
Theothertwopondsarelargerand someofthebottomconsistsofbaremud.
massesofPhormidium.
LarvaeofDipteraareabundantin Pond13in floating
M. lepaPonds18,21 and 22 are ofthePond3 type. In Pond 18 therotifers
della,EuchlanisdilatataEhrb.,andE. orophaGossewerefound.Pond17isin a
sp.
andMonostyla
rockypocketwithwaterabouta footdeepandhasE. dilatata
Diver,
Theusualaquaticbirdsoccur,viz.: GreyPhalarope,Red-throated
Ducksand King Eider. The youngof the latterfeedin
a fewLong-tailed
plants,etc.(39, 64).
freshwater
pondson crustacea,
2.
STREAMS.
are ratherpoorin lifeas faras couldbe ascerspeaking,
These,generally
greenalgae attachedto subtained. In someplacesthereare filamentous
stones.
merged
C. KLAAS BILLEN BAY. (BruceCityRegion.)
Thisdistrictlies at the head of Klass BillenBay. The area consistsof
a veryextensive
raisedbeach,someofwhichis stillmoreor
twoparts,firstly,
whilethegreaterparthas beensiltedoverby
lessin its originalcondition,
and secondly,the CampbellRangewhichrisesabruptlyfromthis
streams,
beachto a heightof2500feet.
can be distinguished:
communities
Thefollowing
(a) Land Communities.
Zone.
1. Inter-tidal
2. RaisedBeach.
(a) Unsilted.
(b) Silted.
3. UplaindRegion.
(b) AquaticCommunities.
1. Brackish.
2. Fresh-water.
(a) Ponds.
(b) Streams.
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V. S. SUMMERHAYESAND C.
S.
259
ELTON
(a) Land Communities.
1. INTER-TIDAL ZONE.
The plant ecologyof this regionhas been describedby Walton (66). He
has shown the transitionfrommarineconditionsthroughsalt marshto the
unsiltedbeach on the one hand, and to the silted"moor" on the other.
The shingleshoreoutsidethe salt marsharea (see Plate III, fig.2) is much
exposedto thegrindingactionoffloatingice brokenofffromthe Nordenskjold
Glacier. Thereis no plant life,but betweenthe tide marksare the following
animals:
ofsmallspecimens
ofsomespecies
Numbers
Nemertinea;
Collembola:Archisotoma
beselsi
Linn.
Mites:
Bdellalittoralis
On the driftline thereis much seaweed, etc., deposited;in one place a dead
seal had been cast up. The followinglive above the driftline on the shingle,
but feedamongthe organicdrift.The spidersfeedon the otheranimals:
armatusvar.arcticus
Collembola:Onychiurus
Spiders:
ErigonearcticaWhite
LK.
E. tirolensis
Probablyotherspecies
Bdellalittoralis
scavenging
Mites:
Another
mitein deadseal
rare,in deadseal
Oligochaeta:Henleasp.
Lumbricillus
aegialites
Steph. va. in seaweed
L. necrophagus
Steph.
va. in deadseal
Purple Sandpipersfeedalong the shore. Thereis much dry driftwoodon the
beaches, but no animals werefoundin or underit. OInthe mud flatsinside
the bouldersdescribedlater the miteBdella littoralis
is verycommon. A moss
sample of Bryumnitidulumwas taken fromthe salt marsh (Walton, p. 116,
Zone II), and examinedforthe animals. This zone is not coveredby ordinary
tides, and consistsof a close growingmoss layer on black mud with much
decayinghumus. In the moss were:
viaticus
Collembola:Achorutes
Folsomiaquadrioculata
a
F. sexoculataTullb.
armatusvar.arcticus
!
Onychiurus
r.
Larvae
Diptera:
a.
crymodes
Steph.
O]igochaeta:Enchytraeus
Nematoda: +
Therewereno rotifers.The wormswereeatinggreenmossleaves.
Countsof theseenchytraidwormsweremade in orderto get some idea of
theirimportancein the biologyof the soil. The groundconsistedof 1 cm. of
moss and 7 cm. of black mud, the wormsoccurringin the moss only. In the
firstplace examined,14 sq. cm. ofmossweretaken. In thisthewormsaveraged
2*3wormspersq. cm. In anotherslightlydrierplace,an averageof 1P75worms
per sq. cm. was obtained. The estimatesare probablytoo low owingto the
smallsize ofthewormsand thedifficulty
ofcountingthemaccurately.Enough
data wereobtainedto showthat these wormsmustplay a not inconsiderable
role in the soil wheretheyoccur,
17-2
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gen and lPear Island
to Ecology of Spitsber
260 Contributions
2. RAISED BEACH.
(a) The unsiltedpart of this has been describedby Walton withrespect
to the plants. Apparentlythis area is a type of open "fjaeldmark" whichis
on its wayto heath. A heathofDryasand Salix polarisseemsto be theordinary
climaxin thisregion,Cassiope heath onlyrarelybeingproduced.
Onychiurus
armatusvar. arcticusoccursundershingleon the lowerpartsof
the beach.
The larger ponds on the unsilted beach are surroundedby a marked
marginalmoss community(see Plate II, fig.2). It is possiblein Pond VII
(== Walton VII) to distinguishzonation on the bank. The zones noted are
as follows,the area outsidebeing a typical sparse "fjaeldmark" community.
Zone 1 is the highest,whilepart of Zone 3 is submerged:
...
Equisetum variegatum
Luzula
Poa
sp.
confusa
...
...
...
...
...
...
Salix polaris
Saxifraga aizoides
...
...
...
...
...
...
Bryophytes:
...
Blepharostoma trichophyllum
Brachythecium salebrosum var. arcticum
Bryum
pendulum
Cinclidium stygium
Hypnum brevifolium
H. giganteum
H. polygamum
H. revolvens
H. stellatum
H. vernicosum
...
...
...
...
...
...
...
x
x
x
-
-
x
f.
a.
-
-
...
-
...
x
x
...
...
...
...
...
...
...
...
...
...
...
...
x
x
x*
x
...
...
...
x
-
-
...
...
...
.
i.f.
-
...
...
..
.....
...
Lophozia bicrenata Dum.
Orthothecium chryseum
Swartzia montana ...
f.
...
...
...
...
...
...
...
x
x
...
...
...
...
B. pseudo-triquetrum
Camptothecium nitens
1
f.
...
...
...
.f.
...
...
Polygonum viviparum
l.f
-
...
3
x
...
...
...
...
...
...
Cardamine pratensis ...
...
Cassiope tetragona
...
Dryas octopetala
2
1
Nos.
Phanerogams, etc.:
-
x
x
-
x
f.
f.
x
-
x
x*
x
-
Lichens:
Cetraria nivalis
Lecanora
tartarea
x
...
-
x
-
Rotifera:
Adineta barbata
A. gracilis ...
A. vaga
...
Habrotrocha constricta
...
Macrotrachela ehrenbergii
...
H. sp.
...
M. habita
...
M. multispinosa
...
...
...
Pond Rotifers-several
spp.
x
-
...
...
...
..
...
x
x
...
.-
...
...
...
Tardigrada
....
...
...
...
Protozoa
...
...
...
...
Oligochaeta:
...
...
...
...
...
...
...
...
sp ...
x
...
...
...
Mesenchytraeus
-
...
Nematoda
Enchytraeus crvmodes
Henlea heleotrophus
x
-
...
x
x
x
x
x
x
x
x
-
...
x
x
r.
x
x
x
* Plants marked thus grow submerged. The Rotifers may not be constant, but those of
Adinetcapr6bably show their normal distribution.
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V. S. SUMMERHAYES
AND
C.
S. ELTON
261
(b) SiltedRegion. This formsmuchthe greaterpart of the area. The substratumvaries fromdrystonyareas on the ridgesto damp boggyflatsalong
the streams. On the seaward edge the beach mergesinto the salt marshmentionedabove, or else is terminatedby a low cliffwitha shinglebeach below.
Silt broughtdown by the streamsin springand early summeras a resultof
snow meltingon the mountainsis beingdepositedoverthe wholeregion. The
large streamjust next to the NordenskjoldGlacierhas graduallywornaway
the raised beach, leaving a bare shingleflat. A large colonyof ArcticTerns
nestson the slightlyhigherparts.
(i) WetTundra. Owingto thefrequentchangesin thecoursesofthestreams,
bare areas of silt are constantlybeing produced. The firstplants to appear
and DupontiafisheriR.Br. In
in such habitats are Eriophorumscheuchzeri
places JuncusbtglumisL. also occurs. Silt is caught by these colonisers,and
as the level risesa wet type of vegetationis produced.
Phanerogams:
Roth.
Eriophorumangustifolium
f. tristeTh.Fr.
E. scheuchzeri
Bryophytes:
Hypnumscorpioides
H. stellatum
Juncusbiglumis
Luzula confusa
Salix polaris
Splachnumvasculosiim
This gradesoffinto the Dryas heath on driersoil.
fromsucha bog nearthe salt marsh
of moss(H. stellatum)
The micro-fauna
was examined.
Adinetabarbata
A. vaga
Habrotrochainsignis
Macrotrachelaaculeata Milne
PleuretrabryceiWeber
a.
Oligochaeta: Enchytraeussp.
Rotifera:
A wormcount was also made here. There was an average of 3-25 wormsper
sq. cm. in an area of 20 sq. cm.
The small pools of these marshesand theirflorahave been describedby
Walton (p. 116). Hypnumscorpioidesoccurs typically,submergedin these
pools. Sampleswereexaminedand showeda richfaunaofRotifera,Nematoda,
Tardigrada,and Protozoa (forRotiferssee Bryce(8) under "L 21 ").
(ii) Heath. This consists of a more or less closed communityof Dryas
octopetala,on the ridgesbetweenthe streams. In places the Dryas plants are
orientatedat rightangles to the prevailingwind(66). The followingspecies
occur:
Phanerogams:
Braya purpurascensR.Br.
Cassiope tetragona(verylocally)
Draba alpina
D. alpina var. oblongata
D. hirtavar. aretica
d.
Dryas octopetala
Bryophytes:
Bryumglobosum
Dixon
and var. ruberrimum
Festuca brevifoliaR.Br.
Pedicularishirsuta
Polygonumviviparum
Salix polaris
Saxifragaaizoides
S. oppositifolia
B. pendulum
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262 Contributtions
to Ecology of Spitsbergen
and Bear Island
A gooddeal ofthearea consists
of"rute-marks"
(see Plate III, fig.1),which
soil (27) and thusdiffer
are polygonalareas in almosthomogeneous
from
on PrinceCharlesForeland.As intheothertype,the
thosealreadydescribed
centreis bareofplants,whilethecracksbetweenadjacentareasareoccupied
islandicaoftenoccurswiththelatter.
byDryasorSalix. Cetraria
a richfloraof
(iii) ErraticBlocks.Thereare a numberof thesebearing.
mossesandlichensoftheusualspecies,e.g.Xanthoria
parietina,
etc.,and also
the alga Prasiola crispaMenegh.This layet of vegetationformsa more
thantheshingle.Hereoccurunderthemosses,
favourable
habitatforaniimals
etc.:
Collembola: Folsomia quadrioculata r.
Spiders:
Erigonearctica
f.
f.
Hilaira glacialisThor.
r.
Bdella sp.
Mites:
f.
Rhagidia gelida
Oligochaeta: Rare
Nematoda: A largespecies.
Sporadic
thuspossess
Erigonewasseento eat themiteRhagidiagelida.Theseboulders
communities
oftheirown.
self-supporting
Thefaunaofbothsiltedand unsiltedbeacheswillbe considered
together.
Mostofthe observations
on the latterweremadearoundBruceCitywhere
thereare a numberof ponds. Thereis verylittleanimallifeexceptwhat
comesfromthewater(flies).The presenceofmanduringthelast fewyears
has drivenawaymanyofthebirds.
Collembola:
Diptera:
Isotoma viridis
ChironomusripariusMg.
C. lugubris
Metriocnemusursinus
M. brevinervisHolmg.
Cricotopusbasalis
C. glacialisEdw. (on flowersofDryas,Saxifragaoppositifolia,
hirculus,
aizoides)
Orthocladiusconsobrinus
0. decoratusHolmg.
0. festivusHolmg.
Psectrocladiusborealis
P. limbatellusH.
CamptocladiuspumilioHolmig.
C. curvinervisvar. polaris
Aedes alpinusL.
Exechia frigida
Sciara praecox
S. pallidiventrisH.
Limnophorahyperborea(on flowersof Dryas, Silene,and Saxifraga
hirculus)
L. megastoma(on Silene flowers)
Scatophaga varipesHolmg. (on dung)
Hymeinoptera:StenomacruspedestrisHolmg. (on Silene flowers,Aug. 2nd)
Atractodes bicolor var. arcticus Holmg. (on flowersof Saxifrayca
hirculus,Aug. 4th)
Ichneutes hyperboreusHolmg. (on flowersof Dryas and Saxifraga
oppositifolia)
(Pristiphorafrigida)?
Spiders:
Typhochrestusspetsbergensis f.
Hilaira glacialis
r.
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JOURNAL OF ECOLOGY
VOL. XI, PLATE III
...~~~~~~~~~~~~~~.
..
..~~~~~~~~~~~~~~d
Piwlo
I
Walton~
FIG. i. Klaas BillenBay. Area ofsiltedraisedbeach showincr"rute-marks."
Betweenthelarge
polygons are plants of Salixfiolaris and mosses.
Pl'wto. A. M. Carr-Saunders
FIG. 2.
Klaas BillenBay. I ntertidalshinglebeach withdriftline. Note absence ofplants; also
thedrifting
ice, whichis mainlyresponsibleforthepaucityoflife.
SUMMERHAYES
AND
ELTON-ECOLOGY
OF SPITSBERGEN
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AND BEAR ISLAND.
V. S. SUMMERHAYESAND) C. S. ELTON
Mites:
Oligochaeta:
Birds:
Mammals:
Bdella decipiens
B. pallipesL.Koch
Cyta brevirostris
Rhagidia gelida
Sphaerozetesnotatus
Henlea brucei
Pink-footedGoose
Purple Sandpiper
Buffon'sSkua (Stercorarius
longicaudusVieill.)
Snow Bunting
Dogs kept herein 1921
263
r. Nesting
f. Nesting
r. Probablynesting
Feeding
as shed antlerswerefound.There
Reindeermust have roamedhereformerly
are veryfewcollembola,mitesand oligochaeteson account of the sparseness
and drynessof the plants. Spiders are fairlyabundant, especiallyne'arthe
ponds,and theymustlive largelyupon the flieswhichbreed there. One was
Most of these fliesseem to be
seen to attack the flyChironomus
hyperboreus.
and Psectrocladius).Cricotopusbasalis and
aquatic (Chironomus,
Orthocladius
glacialis were seen layingeggs in Pond VII on August 1st. The ichneumons
and Atractodes
are probablyparasiticon Diptera (69). It is clear
Stenomacrus
by now that the Purple Sandpipereats almost anythingthat it can get. The
stomachsof two youngbirdshad Lepidurusarcticusfromthe ponds in them
(August 15th),and anotherhymenoptera.The birds are ofteninfectedwith
bird-lice:
Degeeriellaactophilus
D. zonaria
MenoponlutescensBurm.
Philopterusfusiformis
The Sandpipershad mostlyleftBruce Cityby August 1st.
is a braconidwhichprobablyparasitisesthe sawfly
Ichneuteshyperboreus
Pristiphora
frigida(i9). The latterwas not seen here,but it may have been
in the larval stage. Stenomacruspedestrisis recordedon flowersof Dryas,
Saxifragacaespitosaand Cerastiumalpinumin Spitsbergen(19).
It was surprising
to findthat a verydrymoss(Grimmiacommutata
Huebn.)
had a richmicro-fauna-13speciesof rotifers(see Bryce(8) under"L 25").
3. UPLAND REGION.
Severaldistrictsare includedin theabove term,but mostoftheinformation
applies to the slopes of the CampbellRange. The greaterpart of thisrangeis
composed of sandstones and carbonaceous shales; there are also bands of
Gypsum. This Gypsummaybe 200 feetin thickness,but is generallymuchless.
AcrossAdolfBay is the De GeerRange, composedchieflyof Archaeangneiss
and mica-slate. Almostall the surfaceof the CampbellRange is coveredwith
screes,and as a resultthe vegetationis nowhereclosed,beingalways a typeof
"fjaeldmark" (see Plate IV, figs.1, 2). No flowerslopes as in Prince Charles
Foreland were noticed. This may be due to the northernaspect or to the
greaterdrynessof this region. The slopes are also swept by winds fromthe
snow-coveredinterior,and this increasesthe barrenness. Where the snow-
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and Bear Island
264 Contributionsto EcologyofSpitsbergen
waterdrainsalong depressionsa slightlydampertypeofvegetationthanusual
appears.
The followingplants occuron the CampbellRange:
f. triste 1.
tEriophorumangustifolium
1.
Alsinerubella
1.
tJuncusbiglumis
Carex misandraR.Br.
*Papaver nudicaulevar. radicatum f.
*Cerastiumalpinum
Pedicularishirsuita
*C. caespitosumMalmg.
Poa cenisiaAll.
Draba alpina
tPolygonumviviparum
D. alpina var. oblongata
a.
*tSalix polaris
D. rupestrisWg.
1.
tSaxifragaaizoides
D. subcapitata
o.
S. caespitosa
D. wahlenbergii
f.
S. oppositifolia
a.
*Dryas octopetala
Species markedwithan asteriskoccur on Gypsumscreesat a heightof 300-400 feet. Those
markedthus t occur at lowerlevels (up to 500 feet).
Poa cenisiais a commonplant on limestonescreesin the Alps (56). Saxifraga oppositifoliais the only plant occurringat 2000 feet in this area. In
otherparts of SpitsbergenPapaver and otherspecies have been recordedat
great heights. The followingplants were found at 3000 feet (approx.) on
MountTerrier:
Cerastiumcaespitosum
Draba alpina
Grimmiaapocarpa
Papaver nudicaulevar. radicatum
Saxifragaoppositifolia
and probablyothersoccur. It is interestingto note that threeof the above
plantsreach 2000 feetin GrinnellLand (23).
On the De Geer Range conditionsare rathermorefavourable. This faces
and Dryas octosouthso that in shelteredplaces a heath of Cassiopetetragona
petalais formed.Possiblythelack oflimehereis ofimportancein determining
the luxurianceof Cassiope.
The animalsof the upland area werenot workedout in detail. The spider
is confinedto the flatshingle,and its place is taken by LeptyTyphochrestus
phantessobriusat the point wherescree slopes start. The limitingfactoris
clearly not the height, but the nature of the substratum. A few mites
etc.) occurin the screes. Snow Buntingsnest in
(Bdella sp., Cytabrevirostris,
the boulderand screeregions. The exploringpartybroughtback someCollembola fromnear the top of MountTerrier(2800 ft)-Achorutesviaticus,Agrenia
bidenticulata.
(b) Aquatic Communities.
The ponds and streamsmay be seen in the photo given by Walton (66,
Plate VI, fig.1). On the raised shinglebeachesformingterracesof successive
ages, thereis a seriesof ponds,the lowerbeingstilltidal whilethe upperones
are freshwater.The latter contain relictbrackishcrustacea,and must once
have beentidal. At a laterdate theywerecut offfromthe sea as a resultofthe
risingofthe land whichhas been goingon sincethe Glacial Epoch.
Figs. 5 and 6 showin a diagrammaticway the positionofthe Klaas Billen
ponds. Nos. 1-11Tare tidal,whileIV-XI are freshwater.Waltongivesa map
numbersfor the
showingthe details of the tidal area, but he uses different
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JOURNAL OF ECOLOGY
VOL. Xi, PLATE IV
r
s~~~~~~~~~~~~~~~~~~~~~~I
_Vah
Photo.
J.
-AL.
too
FIG i. Klaas Billen Bay. Scree on Campbell Range with plants of Pa
and Saxifraga oJjfiosi/ifoliti.
izudicauilevar. radicatumzi
c.
J-11ht0ot.
3
Alton
FIG. 2. Gips Valley. "Fjaeldmark" on slope. Note the rocky debris everywhere
and scattered vegetation.
SUMMERHAYES AND ELTON-ECOLOGY
OF SPITSBERGEN
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ANDBEAR
ISLAND.
V. S. SUMMERHAYESAND C.
S. ELTON
265
ponds. The diatomsofthesepools showa transitionfrommarineto freshwater
conditions(66, p. 119).
We shall describethe aquatic communitiesin orderof theirnatural succession,startinlg
withnmarine
and endingwithfreshwater.
K L AAS
ADOLPH
X I1LLS_N
E
:BA
EY
CAPE NAPIBDt
C ir A
. . . .
. . . .
.2
.
.T
77,7,7777777
z
.......
WACCR
f
.....
i
O
celY
SoC -EDE SS"St
UEAcMS
(64UrS)
MubD
-
_
AOM
sIlwreb
FLATr
: 43S
DSA
FIG. 5. Diagrammaticsketchof Klaas Billen Ponds and Beach lines. Heightsare accurate.
FIG. 6. This showsthe relativepositionsof the ponds and the relationbetween
successiveelevations.
1. BRACKISH COMMUNITIES.
The Boulders. These really belong to the inter-tidalzone, but they are
included here since the crustacea can be compared with those of the tidal
ponds. They forma line lyingat the outer edge of an alluvial fan of mud
broughtdown by streams,and are completelyuncoveredat low tide. They
are much batteredby floatingice fromthe NordenskjoldGlacier,and act as
a barrierpreventingmuch ice fromenteringthe ponds. The algae are describedby Walton (p. 118).
On the boulders(at low tide):
Hydroida:
Gonothyraealoveni
Opercularellalacerta
On algae
Gammaracanthusloricatus
Gammaruslocusta var. zaddachi
Pseudalibrotuslitoralis
Several speciesof Copepods(Dactylopusia,Harpacticus,etc.)
among hydroidsand algae.
Halacaridae: On hydroids
Birds:
ArcticTern
Poly-zoa:
Crustacea:
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266 Contributions
toEcology of Spitsbezgenand Bear Island
A Tunicate,
Rhizomolgula
globularis
Pallas,livesonthesandymudaroundthe
boulders,
just above low tide mark. A fewPolychaetewormsoccurin the
nmud.The hydroidsonlygrowon places protectedfromfloating
ice. The
Gammaracanthus
are oftheusualmarinetype,and showno transition
to the
freshwater
formG. lacustris.The Gammarus,
however,are to someextent
intermediate
betweenG. locustaand thevarietyzaddachi.It has beenshown
in Denmark(61) that the structure
dependson the salinityin whichthe
animalsare living.Thisis also trueofthosehere,sincethosein Pond I are
moreliketheextreme
varietyzaddachi,whilethosefromthesebouldersand
fromCapeScottare intermediate.
Thefaunaand floraofPonds1-111areseenin TableIII.
TABLE
Birds:
:Fish:
Crustacea:
Schizopoda:
Amphipoda:
Copepoda:
Collembola:
Hirudinea:
Rotifera:
Tardigrada:
Nematoda:
Protozoa:
Algae:
III. Fauna and Flora oftheTidal Ponds.
Arctic Tern
...
-......(x)
Pond No. I
II
x
(x)
...
...
--
Mysis oculata Fab. var. relictaLov.
young
adult
...
...
III
...
...
Gammaruslocusta var. zaddachi
Pseudalibrotus litoralis ...
Eurytemora raboti Richard
...
..
...
...
Chlorophyceae: Enteromorpha sp.
...
...
...
Ulothrix sp.
...
Zygnema sp.
Cyanophyceae: Chroococcus turgidus
...
Gomphosphaeria aponina Kuetz.
Merismopedia glauca
...
Oscillatoria sp ...
...
...
Rivularia sp.
...
subsalsa
Oersted
Spiruilina
x
x
(x)
-
(x)
x
x
x
x
?
?
x
...
x
x
x
(x)
x
x
Tachidiusspitzbergensis
Olofsson...
T. longicornis Olofsson ...
Archisotoma beselsi
...
Achorutes viaticus
...
Pontobdella muricata
...
...
Colurella colurus Ehrb.
Encentrum? raptor Gosse
...
...
...
...
...
...
...
...
...
x
( x)--
...
...
...
...
x
x
-
...
x
x
x
x
x
x
x
...
x
x
...
...
x
x
?
...
...
...
x
...
...
...
...
...
...
x
-
x
x
x
x
x
x
x
x
.
x
x
x
?
x
x
x
x
x normallypresent; - definitely
absent; (x) accidentalor sporadic;
? probablypresentthoughnot found.
Pond I (= Walton,Pool I). A shallowpond,not morethan a footor
twodeepat hightide. Therearenoplantsexceptalgae. Thebottomis mostly
sand and mud coveredwitha thinbrown"felt" of organicmatter,which
containsmanydiatomsand blue-green
algae. The leechwas attachedto the
of an adult male Mysisrelicta(August10th). These,and a
cephalothorax
smallfishaboutan inchlong,wereclearlybroughtin accidentally
fromthe
are
seem
fjord. Tardigrades noticeably
absent. All the regularinhabitants
orvegetarian.
to be scavengers
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V. S. SUMMERHAYES AND C.
S.
267
ELTON
PondII (= Walton,
tothelast,butsmaller.
PoolIII). Similar
PondIII (- Walton,
PoolIV). A smallpondcontaining
largenumbers
andother
ofEurytemora
raboti
which
liveonithebottom,
andfeedondiatoms
algae.
TheSpring(S onWalton's
butforms
MapandonFig.5). Thisis fresh,
part of the watersupply of the tidal ponds, and flowsinto Pond III. There
are no crustacea. A collembolanSminthuridesmalmgreniwas seen on the
surface.An oligochaeteHenlea heleotrophus
is abundantin submergedHypnum
cordifolium.Also rotifers,
tardigrades,nematodesand protozoa.
GENERAL DISCUSSION.
We proposeto give some account of the conditionsin these ponds,and of
the limitingfactorsaffecting
the distribution
of some of the animals.
The sea penetratesas faras Pond III at hightide. Completewateranalyses
were made in two cases by Mr Manley,and chloridedeterminations
in the
othersby Mr Mills. Two series of samples were taken, one at high and the
otherat low tide.
in gis. per litre.
TABLE IV. Chloridecontent
Low tide Hightide
Locality
Aug.10th Aug.7th
Sea by BruceCity,Aug.1st ...
18-26
17-57
,,
Aug. 3rd ...
Boulders
...
...
...
...
17-11
PondI ...
...
...
...
101
16-12
9
99
II...
...
...
...
III. Mainpond
...
III. Wherestreamenters
0 57
0-22
15-23
9-10
0-55
It will be seen that thereis a gradientof chloridecontentfromthe sea to
Pond III, both at highand low tide,thoughit is much moremarkedat high
tide. The ponds are practicallyfreshat low tide, the time duringwhichthey
remainso increasingas we pass fromPond I to Pond III. This, combined
withthe influenceof the freshwaterflowingin, causes the tides to have less
effectin the innerthan in the outerponds. If Pond III wereraised onlya foot
or two it would then be a relictfreshwaterpond like those higherup on the
beaches. Table V gives the analyses,whichshowtwo oppositeextremecases.
It will be seen that the total amount of salts in Pond III water at low tide
is about 1/40thof that in Pond I water at high tide, and the osmoticpressures, etc., would differaccordingly. Also calcium carbonate is presentin
relativelygreaterproportionthan the other salts in the fresherwater of
Pond III. Thus comparingthe two samples,thereis about 1/9thoftheamount
ofcalciumin one that thereis in the other. But the amountsofotherelements
comparedin the same way are far smaller,e.g. magnesium1/50th,sodium
1/50th,chlorine1/70th. This shows that the effectof tidal influenceis not
merelythat of dilutingor concentratinga given solution,but is much more
complicated.The proportions
ofsaltspresentmaybe altered,and suchchanges
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268 Contribationsto EcologyofSpitsbergenand Bear Island
have great effectson aquatic animals. Thereforethe chloridecontentis not
in itselfthe mainfactorin the chemistryofthe water,but is a roughindicator
ofothermoreimportantfactorswhichare harderto determinedirectly.These
are the amount of mixingof freshand sea water (the chloridecontentis
about halfthe total solids),the osmoticpressure,the proportionsof different
salts, etc.
TABLE V. Pond I. High tide,innerside,August7th.
Amountsgivenin gms.per litre.
0-754= 1-346CaCO3 or 2-179CaH2 (CO3)2
1-664= 3-764MgSO4.2H20
16-120= 1.513 MgC12
24*750NaCi
1-930
S03
Total solids=31-373
of Total Solids gave 31-00and 31-05.)
(Direct determinations
Ammoniumand Potassiumsalts absent.
CaO
MgO
012
Pond III.
Low Tide, August10th. Amountsgivenin gins.per litre.
0.092 =0-164 CaCO3 or 0-266CaH2 (CO)2
0 049=0-191 MgSO4.2H20
0-225=0-371 NaCl
0 079=0-078 Na2SO4
S03
Total Solids==0804
of Total Solids gave 0-797.)
(Direct determination
Ammoniumand Potassiumsalts absent.
CaO
MgO
C12
rabotiis mostabundant in Pond ITI, and less so
The Copepod Eurytemora
in Pond II, only adults occurringin the latter. Experimentswere made to
salinities,obtainedby mixing
testits powersof survivalin watersof different
freshwaterwith sea water. Pure sea water (chloride18l26 and 17.57) killed
the EurytemorafromPond III in about 20 minutes,while controlsin fresh
water were quite healthy. In a salinityof 13-20they died in about 1 hour
after31 hours. These experiments
40 minutes,but in 8-82theywereunaffected
show that it is limitedin its distributionpartlyby salinity,and the results
agree with its distributionin the ponds. (Gammaruszaddachi fromPond I
would eat large numbersof Eurytemorain captivity. Table III shows that
thesetwo specieshardlymeet. This may be anotherlimitingfactorforEurytemora.) It is clear that it is not only the degreeof salinitywhichaffectsit,
but also the time duringwhichthis salinityacts. These ponds lie very near
high water mark,and are only floodedby the sea foran hour or two. The
animals may be able to resistfora shorttime a high salinitywhich would
kill them if it lasted much longer. This "time-salinity"factormay be one
cause of the dwarfingwhich occurs among many marine animals living in
tidal waters. Theirmetabolismis loweredduringtimesof abnormalsalinity,
and thus less growthis possiblein theirlifetime.
Mysis oculata occurs in arctic and subarcticseas (we found it near the
AnserIslands,at the mouthof Klaas Billen Bay). The formor varietyknown
as Mysis relictaresemblesa youngM. oculata,but diflersin some points,and
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V. S. SUMMERIIAYESAND C.
S. ELTON
269
is sexually mature. It is, in fact,a Mysis oculatawhose growthhas been retarded. M. relictahas been found in freshwaterlakes in the Baltic area,
Ireland,N. Americaand Spitsbergen(48). In SpitsbergenM. relitcawas found
by Olofssonin threeplaces: (1) in Sassen Bay (youngoneswashedintoa lagoon
at high tide and out again at low, and thus belongingreally to the fjord
fauna); (2) in theseKlaas Billen ponds (youngones); (3) relictin a freshwater
lake on Credner'sMoraineat the head of Belsound (youngones,August 12th,
1910).
In Pond I youngMysis relictaoccursveryabundantly. Only one or two
adults wereseen. This and the factthat one had a marineleech on it proves
that theseadults belongto the fjordfauna. These facts,combinedwithOlofsson's resultsfromSassen Bay, showthat Mysisoculataundergoesa retardation
of growthin the innerparts of long fjordsin ratherlow salinity,and that
under certain conditionsit may become relict in freshwater. The process
mightoccur by the cuttingoftof an arm of the sea whichgraduallybecame
fresh. This may have occurredin the Baltic and in Belsound. Arcticconditionsare especiallyfavourablefor such processes,on account of the land
elevationassociated witha retreatingice-sheet. The Mysis relictafoundnow
in Europe, etc., were almost certainlyleft behind afterthe Glacial Epoch.
A studyof the ecologyof M. reltctaat Klaas Billen Bay showedsome of the
factorsaffecting
its chancesof becomingrelict. Why has it not become relict
in thesefreshwater
ponds whileEurytemora
has? Now the tidal ponds freeze
solid in winter,and Mysis and Gammarus,which have no adaptation for
survivingthis,would all be killed. Eurytemora,
however,has eggs whichcan
stand freezing. ThereforeMysis and Gammarusmust migrateinto Pond I
every year, only to be killed when wintersets in. The lake on Credner's
Moraine,whereMysis relictaoccurs,is over 9 metresdeep, and waters over
2 metresdeep are not frozensolid (48). How are we to explain the rarityof
adult Mysis in Pond I since they mighthave been expected to inigratein
fromthe fjord? The young ones are veryabundant; Dr Tattersalldescribes
themas being"all quite juvenile,nonelargerthan 9 mm." (July27th). Those
obtainedhereon August5th,1910,by Olofssonwerefrom8-6to 9.3 mm.long.
Now ArcticTernswereoftenseen fishingin Pond I and sometimesin Pond II.
They were also seen to hover over Pond III, but neverto dive into it. Apparentlytheywerenotfeedingupon Mysisrelicta,theyoungofwhichare small,
transparentand very agile, but on Gammarus,whichhides in the mud and
occasionallymakes shortjourneys,duringwhichit is quite conspicuous. The
in Pond I, and rarelyin Pond II, whichcorrespondsto
latteroccurcommnonly
the habits of the Terns. Examinationof some Terns' stomachsfromPond I
showedthat theyhad been eatingAmphipods,no traceof Mysis beingfound.
Ternswereseen feedingon Gammaridsat Cape Boheman (64), and at Richard
Lagoon. Oli the otherhand,remainsofadult Mysis werefoundin the stomach
of a Tern whichhad,been fishingby the Boulders. Thus the depredationsof
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to Ecologyof Spitsbergenand Bear Island
270 Contributions
Ternsmay preventmanyadult Mysis fromlivingin Pond I. Those seen were
muchmoreconspicuousthan the young.
To sum up the probable mannerof relictionof Mysis oculata: the first
retardationof developmenttakes place at the heads of long fjordsor other
places wherethe wateris ofa lowersalinitythan that of the open sea, though
not necessarilyin estuarine waters. The young of this fjord form (or of
M. oculata?), if they get into estuarinewaters, may survive the summer
(partlyon account of theirsmall size and consequentimmunityfromTerns),
but not the winterunlessthe wateris deep enoughto preventthe pond from
freezingsolid. If it is deep enough(over 2 metres)the Mysis willbreedin the
tidal waters,and willthenbe safefromthe attack ofTerns. It may ultimately
becomerelictin freshwater.
.Anotherproblemis theabsenceofMysisand Gammarusfromthetwo inner
ponds. A fewMysis werefoundin Pond III late in the season (August14th),
but beforethat none occurred. They are not limitedby salinitysince those
in Pond I livemostof the timein freshwater,and someMysisand Gammarus
werekeptforover 12 hoursin freshwater(chloride0027 gm.perlitre)without
being affected. Mysis and Gammarushave to recolonisethese ponds every
springon account of the winterfreezing,and the extentto whichthey can
penetratedependspartlyon theirpowersofdispersal. Mysis relictawas found
to be positivelyrheotactic,but owing to its small size and weak powers of
against currentsit onlyoccursfora certaindistanceup the stream
swimming
connectingPonds I and II, and seems unable to pass up the more rapid
stretches.Since the Mysis live on the bottom,wheretheyfeedon algae, etc.,
theirchance of beingwashed into Ponds II and III by tides is small,though
it does occasionallyhappen.
Gammaruszaddachiprobablyfindsmostofits foodin Pond I, wherematter
is broughtin by the tide; Ternswouldact as a checkon any pioneersentering
loricatusoccurs in the relict lake on Credner's
Pond II. Gammaracanthus
formG. lacustris,but was
Moraineand in Scandinavianlakes in its freshwater
not foundin thesetidal ponds at all. It is ratherremarkablethat no chironomidfliesinhabitthe tidal ponds,sincetheyare one ofthefewgroupsofinsects
in brackishconditions.They occurcommonlyin estuarineareas
whichflourish
in temperateregions.
2.
FRESHWATER COMMUNITIES.
(a) Ponds. Figs. 5 and 6 show the relative positionsand ages of the
freshwaterponds on the raised beaches. They are all on shingle,the water
being held in primarilyby the ground-ice,also by the marginalvegetation.
This ground-iceprovidesby its meltinga continuoussupply of water which
is supplementedby drainage fromhigherground. The result of this is the
also from
absenceoftemporaryponds ofthe Cape Bohemantype. They differ
raboti.
the latterin havingthe relictCopepod Eurytemora
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V. S. SUMMERHAYESAND C. S. ELTON
271
TABLE VI. List of Crustaceaoccurring
in theFreshwater
Ponds.
VI
Pond No. IV
VII VIII
x
x
x
x
Lepidurusarcticus
x
x
x
Daphnia pulex
x
x
x
Macrothrixhirsuticornis x
Chydorussphaericus
x
x
x
x
x
x
x
x
Eurytemoraraboti
Cyclopscrassicaudis
x
x
x
x
x
x
Maraenbiotusbrucei
Candona rectangulata
x
x
x
x
x
x
x
'x
Eucyprisglacialis
0 =recorded by Olofsson(our collectionlost).
X
x
x
0
x
x
0
0
0
0
Table VI showsthe crustaceaof theseponds. Olofssongives the crustacea
of some of the ponds, and these agree with our records. He also gives much
information
about the rotifers.
Pond IV (= Walton,V). One ofseveralsmallpools in boggyland, and the
newestof the series.
Pond V. Now driedup, but originallyformingan intermediatelink.
Pond VI (= Walton, VI and Olofsson,X). A large pond 300 yards long
and 1-22 feetdeep. Bottom sandy-mud,moss bank, etc., on side. Its fauna
is the same as that of Pond VII, whichis describedin detail below.
Pond VIII (= Olofsson,VIII). Two small ponds lying in boggy land
connectedby a narrowchannel.
Ponds IX, X and XI (=COlofsson,VI and VII, not comparableindividually). These are all at the same height,and appear to be remnantsof a large
lagoon,sincethe skeletonof a big whale was foundlyingnear them. They are
all of the same type-small ponds withsandy-mudbottomand plant covered
margins. These are the highestponds in the series (about 40 feetabove sea
level), and are probablyslightlyhigherthan Pond VIII, thoughthis is not
certain. The lattermay be ofthe same age.
Pond VII (= Walton, VII and ? Olofsson,IX) (Plate II, fig. 2). 140
yards long and 6 inches to I footdeep. It lies about 26 feetabove sea level.
The bottomis mostlyshingle,witha fewpatchesof mud,as a resultof which
Lepidurus is rather scarcer than in Pond VI. The followingcommunities
exist,but they mingleto a great extent:
Plankton:
Crustacea:
Daphnia pulex
Eurytemoraraboti
Polyarthraplatypteraand otherspecies
A large colonial flagellateUroglena sp. in masses. On
this occursa verylarge epizoic Vorticella
In plants and in littoralregion:
Crustacea: Chydorussphaericus
Cyclopscrassicaudis
Maraenbiotusbrucei
Rotifera:
MonostylacornutaMuller
M. lunarisEhrb.
Lepadella patella
Mytilina ventraLisvar. brevispina Ehrb. and others
includingmany Bdelloids describedunder Zone 3 of
the Pond MarginalCommunity
Rotifera:
Protozoa:
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and Bear Island
to Ecologyof Spitsbergent
272 (iontribtttions
Cricotopusbasalis
C. glacialis(layingeggson Augustlst) and probablymost
listedin Raised Beach
of the chironomids
Oligochaeta: See Zone 3 of Pond MarginalCommunity
Protozoa,Nematoda, Tardigrada
On bottom:
Crustacea: Lepidurus,Macrothrix,Eucyprisand Candona
Diptera:
otherbirds.
occurand occasionally
ArcticTernsand PurpleSandpipers
of the waterduringthe end of Julyand
Fig. 7 givesthe temperatures
of August. Thesewereusuallytakenat 11a.m., but on several
beginning
daystheyweretakenat othertimes(July25that 1.30p.m.,July28that
ID
P
1
L
12
24
As
?uL.Y
26
a7
a
21
So
31
1
AU
A
cu
3
4
5
6
7
e
9Y
It
/13
ST
July26th). Readingsweretakendailyat 11 a.m.
FIG. 7. TemperatureofPond VII (no record-for
in the shade, at a depth of 9 inches. Curvesshow maximum,actual, minimum.
5 p.m.,Aug.12that 2.30p.m.). Theselatterrecordsshowthatthemaximum
is reachedin themiddleoftheday. In thisregionit is usually
temperature
colderat night.It willbe seenthattheaveragedailyrangeis about3.50 C.,
4.30 C. Thesetempera14?C. and the minimum
themaximum
temperature
lowlandpondofthesamesize.
turesaremuchlowerthanthoseofa temperate
and
Mountains,
to watersin thewillowzoneofScandinavian
Theycorrespond
in theRhaeticAlps(22). Thislowtemperature
to thoseat about2000metres
becomesa seriousfactorforanimalslikecrustacea,whichhave to growup
to Mr R. F. Stobart,
and breedduringthe shortarcticseason. According
in themiddleon June14th. Most
thispondand PondVI werejustunfrozen
watersfreezeby the middleof September.This allowsabout
Spitsbergen
theotherninemonths
whileduring
in theyearfordevelopment,
threemonths
The
crustacea
solid.
frozen
etc.)in the
are
Eurytemora,
(Daphnia,
theponds
in
a
week
were.about
and
VIII
ahead, development,
X)
smallerponds(IV,
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V. S. SUMMERHAYESAND C. S. ELTON
273
in temof those in the large ones. This is probablydue partlyto differences
perature. The fliescan most likelywinterin the water,at any rate in some
stages. The eggs of Cricotopusglacialis a day or two old werefrozensolid to
- 3*6?C. forover half-an-hour,
and subsequentlydevelopedinto larvae. The
also
was unharmedafterbeingfrozento - 2-5?C.
crymodes
wormEnchytraeus
and probablymanyanimals
for10 minutes.Theseare onlyroughexperiments,
ifthe processis more
to resistfreezing,
can preparethemselvesphysiologically
gradual.
The wateris veryclear,quite freshand alkaline. The continuousdaylight
by algae to continueall the time.
ofthe arcticsummerenables photosynthesis
This must cause the oxygen contentof the water to be high and the C02
contentlow, whichis probablyone reason forthe constantalkalinityof the
water. All watersexamined(on Bear Island and at Klaas Billen) had a PH of
8-2-8-5. This is the same as that of manyEnglishlowlandpondsin summer.
There is probablyless lightat nightunder water,since the sun is lower
then; but that it may be quite strongon a summernightis shown by the
behaviourofDaphnia pulexin Pond VI. Thisspeciesis verystronglypositively
phototropic,and could be seen on sunnydays in a crowdon the side of the
pond towardsthe sun. Duringthe 24 hoursthiscrowdmade a completecircuit
of the pond, followingthe sun.
animalsare vegetarianor scavengers.Lepidurus
Almostall the freshwater
has been seen to eat Daphnia pulex in captivity,and Eucypristo attack old
individualsof Lepidurus(30), but theseoccurrencesare probablyexceptional.
The Eucyprisand Candona here were foundto feed mostlyon diatoms,togetherwithotheralgae. Lepidurus,exceptwhenmoving,is hardto see, being
but duringAugustsmall partiesof Arctic
colouredto matchits surroundings,
Ternsused to hoverover Ponds VI and VII divingforLepidurus. They then
returnedin the directionof the nestingcolony-obviously carryingLepidurus
back to the young. Since this mightgo on forseveral hours,it will be seen
that Ternsact as a considerablecheckon Lepidurus,but at the same timemay
providea means of dispersalby droppingtheircatch occasionallyinto other
ponds on the way back to the nest. Purple Sandpipers' gizzards contained
remainsof Lepidurus. Probably Phalaropes and otherwadingbirds eat this
speciestoo.
Eurytemoraraboti,really a brackish water animal, occurs in ponds on
raisedbeaches in manypartsof Spitsbergen.Olofssonhas dealt in detail with
this question,and thereseems no doubt that it is in most places a truerelict
form. It may possiblyhave been distributedsecondarilyto some ofthe Klaas
Billen ponds. Various animals mighthave done this accidentally. Reindeer
roamed over these beaches, whilemen have occupied Bruce Cityin
formerly
the summersince 1919. Arctic Terns dive into the water when fishingfor
Lepidurusand theymay carrythingson theirfeathers.Phalaropesand other
aquatic birds may also help in dispersal. The communitiesof crustacea are
18
Journ.of Ecology XI
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toEcology of Spitsbergenand Bear Island
274 Contributions
so constant throughoutSpitsbergen(except in temporaryponds) -that it
appears that they must have good means of dispersal. There is very little
ponds accordingto age,
difference
in the crustaceaof this seriesof freshwater
exceptthat in the newest(IV) Daphnia is absent.
Ponds of the shallowtype such as at Klaas Billen and Cape Boheman are
those most commonlyfound in Spitsbergen. There are, however,a certain
numberofdeeper,cold lakes,whosefaunais different.Cyclopsstrenuusoccurs
in these,and also salmon(Salmo alpinus)in someplaces (e.g. in Lake Richard,
in Red Bay, also at Cape Staratschin(11)).
(b) Streams. These occur naturallyon the silted area since they are the
agentsproducingthe silting. They are very poor in life. The followingalgae
werefoundin one stream:
CosmariumbotrytisMenegh.
C. globosumBulnh.
brebissoniiMenegh.
Cylindrocystis
GonatozygonbrebissoniiDe Bary
StaurastrumalternansBreb.
is the most importantof these. Animal life is
The filamentousGonatozygon
absent.
D. GIPS VALLEY.
This area, likeBruce City,is situatedwell insideIcefjord(see GeneralMap,
Fig. 3). It consistsof a broad flatvalley,whose flooris formedof a number
of raised beaches,on eitherside of whichis a highrangeof mountains. The
Gips River runsalongthevalley. In the lowerpart are sandstones,etc.,while
TempleRange and the upperpartsof the mountainsgenerallyconsistof limestonewhichis hard,and formsnearlyverticalprecipices. Onlythe south part
ofthevalleyto the east ofthe riverwas studied. This area has a dryand somewhat continentalclimatewitha verysmall annual precipitation.
We do not propose in this case to give a general classificationof comones.
munities.We shall simplydescribea fewinteresting
1. RAISED BEACHES.
There are a numberof parallel beaches runningtransverselyacross the
valley bottom. The regioncan be dividedinto threezones.
(a) Seaward Beaches. These consistof loose shinglewell above hightide
level. Near the sea the stones,whichare about 1 to 6 inchesin diameter,and
obviouslysea-worn,are quite bare ofplantlife. On the crestsofthesecondand
thirdbeaches there is a quite considerablefloraof crustaceousand foliose
lichens. The commonestof theseis Xanthoriaparietinawhichgrowseven on
driftwood.Otherlichenshereare:
Buellia disciformis
var. trifragmia
Boist.
Ephebe lanata Wain.
Placodiumelegans
PlacynthiumnigrumSF.Gray
Physcia lithoteaNyl.
Rinodina demissaArn.
Thelidiumpyrenophorum
florafromthat oftheraisedbeach at PrinceCharles
Thisis an entirelydifferent
chemicalconstitution
Foreland. The difference
maybe relatedto the different
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ANDC. S. ELTON
V. S. SUMMERIIAYES
27;5
of the substratum.In "lows" betweenthese beaches a small amount of soil
collects(1 inchin places),and heregrowscatteredplantsofSaxifragaopposttifolia, S. caespi,tosaand Draba alpina. This passes into (b) as one proceeds
inland.
in beingformedof moreangu(b) StabilisedBeaches. These beaches differ
lar fragments,and in theirgreat compactnessdue to settlingdown and the
addition of silt. The crestssupportmuch the same type of vegetationas in
the "lows" just described. In additionto the threespeciesmentionedabove,
all of whichare frequent,thereare Draba hirtavar. arctica,and Dryas octopetala. This community(a type of "fjaeldmark") resemblesthat at Klaas
Billen Bay. Lichens are only subordinate. In the "lows" the vegetation
consistsof largetussocksof Dryas amongwhichare a numberof mosses.
Amongthe plantsand understonesthe followinganimalsoccur:
Collembola: Onychiurusarinatusvar. arcticus
Typhochrestusspetsbergensis
Spiders:
Sphaerozetesnotatus
Mites:
Nesting
Birds:
Purple Sandpiper
Ringed Plover (CharadriushiaticulahiaticulaL.)
Possiblynesting
ArcticTern
Mammals: ArcticFox (seen by Mr Tuckerin 1922)
I'ink-footedGeese also nest furtherup the valley. The Purple Sandpiper is
in severalrespects
alwaysfoundwithineasy reachofthesea. Thisfaunadiffers
fromthat on the raised beach at Prince CharlesForeland.
(c) Moss Bogs. Along the banks of the Gips River there are extensive
moss-bogs,alternatingwith bare sandy areas. The moss commullitypasses
intotheDryas-mosscommunityonslightlyhigherground. Onlya fewflowering
salebrosum
plantsoccurin thesebogs. The chiefmosseshereare Brachythecium
mosses
of
the
usual
most
bog
but
chryseum,
var. arcticumand Orthothecium
Boheman.
Camptothecium
occur,and the list resemblesthat given for Cape
nttensdoes not appear to be so commoxias at Cape Boheman. A numberof
streamsrun into the riverthroughthe moss-bog. These have various algae,
them Zygnemasp., Osctllatorialimosa and 0. amoena Gomont. The
anmong
last two are presentas "felts" on the muddybottom.
2. SHELTERED VALLEY NEAR TEMPLE RANGE.
The slopes along Gips Valleysupportthe usual " fjaeldmark" (Plate IV,
fig.2) describedelsewhere(see especiallyTemple Bay). It is poor in species
bothofplantsand animals.
In a valley cut out of the moraineand scree materialjust below Teiliple
Range, a more luxuriantseries of communitiesexists. Probably the south
aspect and protectionfromwinds, especially those fromthe snow-covered
interior,are the masterfactors.
On the slopes of the valley are several definitezones. The highest,oil a
ratherstony slope,consistsof a rich type of "fjaeldmark"; in fact it could
18-2
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276 Contributions
to Ecology of Spitsbergen
and Bear Island
almostbe termeda "herbslope." Salix polarisand Sax?fragaoppositifolia
arethechiefspecies.Thefollowing
also arefound:
Phanerogams:
Cerastiumalpinum
Draba alpina
D. alpina var. oblongata
D. hirtavar. arctica
Festuca rubraL. var. arenariaOsb.
Papaver nudicaulevar. radicatum
Bryophytes:
Hypnumuncinatum
ThuidiumabietinumB. & S.
Lichens:
Ceraniavermicularis
var. tauricaAL.Sm.
Cetrarianivalis
Lecanora epibryon
f.
f.
f.
o.
r.
r.
Pedicularislanata Willd.
f. dasyanthaTrautv.
Polygonumviviparum
Saxifragacaespitosa
S. cernua
S. nivalis
o.
0.
f.
r.
Tortularuralis
Physcia pulverulentaNyl.
Solorinabispora
S. spongiosaAch.
Lecanora tartarea
Belowthisthereis a damperlayerformedof largehummocks
of Dryas
withmanymosses.Thelowestzoneis alongthebottomofthevalley,andhere
a streamrunsthrougha thickmosscarpet. The mossesare the usual bog
species,e.g. Brachytheciqum
salebrosum,
nitens,Orthothecium
Camptothecium
chryseum,
etc. The drainageprevents
ofa Hypnumbogas
theestablishment
at Cape Boheman.
A number
ofboulders
occurhere,andthesesupport
crevice
a characteristic
community
tricho(see Bear Island) withSwartziamontana,
Blepharostoma
and others.Xanthoria
parietinaand Placodiumelegansare common
phyllum
on theactualrocksurfaces.
A richfaunaoccursin thisvalley:
Collembola:
Isotoma viridis
Onychiurusarmatusvar. arcticus
Diptera:
Syrphustarsatusvar. Zett.
AcropteragrontataZett.
Camptocladiusextremus
Hymenoptera: Pristiphorafrigida
Spiders:
Immature
Mites:
Bdella decipiens
Sphaerozetesnotatus
Birds:
Snow Bunting
Nesting
NorthernEider
r.
Nesting
Mammals:
Reindeerhornswerefound
" gave the following
A carefulsearchon the ordinary" fjaeldmark
poor
collection
ofanimals:
Spiders:
Mites:
Leptyphantessobrius
Immatureof anotherspecies
Bdella decipiens
Sphaerozetesnotatus
Ptarmigan
occurin smallnumbers
further
up thevalley. Theeffect
offavourable conditions
is clearlyseenfromtheabovelists.
E.
TEMPLE
BAY
(Sassendale).
Thisregionlieson thesouth-east
sideofTempleBay justnorthofSassendale. It consistsof the southern
slopesand lowerplateauxofthe Colorado
Range. Towardsthevalleytheslopesaregentle,butthereis a steepprecipice
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V. S. SUMMERHAYES AND C.
S.
ELTON
277
seawards. The main part of the regionconsistsof limestonesand dolomites;
thereare also sandstones,and a few thin layers of Gypsum. The regionis
thus predominantlycalcareous. We shall only describecommunitieswhich
are ofespecialinterest,or illustratetypescommonin Spitsbergen.
1. BIRD CLIFFS.
This regioncan be divided into two parts: (a) the cliff,(b) the stabilised
screebelow.
(a) The cliffis almostvertical,and severalhundredfeetin heightin some
places. On ledges nest large numbersof the usual sea-birds,and thereare a
fewgeesenestingon theslopesjust above. Aroundthe nests,and on any ledges
where soil collects,there is a very luxuriantphanerogamicvegetation. At
alpinumand Saxifragacaespttosaare the
the upperedge of the cliffCerastibrum
chiefplants. On GreenMountainbetweenAdventand Coles Bays, wherethere
are similarcliffs,Mr Tucker saw large quantitiesof Alopecurusalpinus and
these are the characteristicplants of such places. SaxiCochleari'aofficinalis;
verytall here. The Alopecurusgrowsextremelywell,
grows
fraga cernuaalso
and formsalmost continuousturfon the ledges. Lichens cover the rocksin
an almost unbrokenlayer. Mosses occur on the clifffaces in such places, an
interestingspecies in this connectionbeing Bryum argenteumL. This is
recordedfrombird cliffsin N.W. Spitsbergen(5), fromIceland (25), and from
East Greenland(14). It is consideredto have beenlintroducedby the sea-birds
themselves.
(b) The manuringeffectis most strikingon the scree below wherethere
is a continuouscarpetof dicotyledons,grassesand mosses. This type of slope
is describedin Nathorst(44). At Temple Bay the followingoccur:
alpinum
Cerastium
alternifolium
Chrysosplenium
var.tetrandrum
officinalis
Cochlearia
Draba alpina
D. alpinavar.oblongata
D. hirtavar.arctica
D. wahlenbergii
Dupontiafisheri
Fr.
angustata
Glyceria
LuzulianivalisBeurl.
L.
Poa pratensis
humileWilld.
Polemonium
R.Br.
affinis
Ranunculus
R. pygmaeus
R. sulphureus
Salixpolaris
caespitosa
Saxifraga
S. cernua
S. nivalis
S. oppositifolia
Fr.
affinis
Wahlbergella
W. apetalaFr. f.arcticaTh.Fr.
This list is muchlongerthan that givenforPrinceCharlesForeland. This
is correlatedwith the much richerfloraof the interiorsof the large fjords.
Polemoniumis a characteristicplant in such localities,and sometimesforms
continuousmats on the ground(3). Many of the plants grow much larger
sometimesdominatesand forms
herethan elsewhere(44). Cerastiumalpitnum
pure growthsover large areas. Hart (23) recordsseveralof the above species
in GrinnellLand and elsewhere below bird cliffs. Where the snow-water
traversesthese slopes thereare a numberof mosses. They are bog species,
but are muchmoreluxuriant(5). Amongthosefoundin such places,the most
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278 Contributions
to Ecology of Spitsbergenand Bear Island
characteristic
are Splachnum
vasculosum,
Weberaschimperi
0. Mull,W. ludwigiiSpreng.var.subcarnosa
Berggr.,
Hypnumuncinatum
var. orthotheceoides
Lindb.and Ceratodon
purpureus
Brid. Marchantia
polymorpha
is limitedto
thishabitat.Thethirdoftheseis considered
by Berggren
to be a varietydue
to thehighorganiccontentofthesoil.
2.
RAISED BEACHES.
A seriesoffouroftheseoccurson theseawardslopejust northofSassendale. Theyare verynarrow(lessthan20 yardswide),but'stretch
fora considerabledistancealongtheslope. Thelowestis about5 feetabovesea-level,
thehighest100feet. Thereis a steepscreeslopebetweenadjacentbeaches.
Thebeachescanbe dividedintotwopartslongitudinally,
theinnerparthaving
a muchbetterdevelopedplantcommunity
thantheouter.Thisis dueto the
different
lengthsof durationof the snow-covering.
Snow driftswouldbe
formed
in theanglebetweenthebeachandthescreeabove,whereastheouter
partofthebeachwillhavecomparatively
littlesnow.
Onthelowestbeachthereis littlevegetation
exceptSaxifraga
oppositifolia
and a fewmosseson theinnerpart.
The secondbeach,about15 feethigher,
has a moretypicalflora.On the
outerregionis a "fjaeldmark"community
likethaton theunsiltedbeachat
Klaas BillenBay (66) and GipsValley.Dryasoctopetala
is themostabundant
plant. Ontheinnerparta heathofDryasand Cassiopetetragona
is developed.
Thefollowing
also occurhere:
Draba alpina
Luzula confusa
Papaver nudicaulevar. radicatum
Polygonumviviparum
r.
f.
r.
f.
Pedicularislanata
f. dasyantha
a.
Saxifragaoppositifolia r.
Silene acaulis
At the angleof thisbeachis a blocktalus. Amongthe blocksoccurCarex
Boott.(3), Cystopteris
hepburnii
fragilisBernh.,Draba hirtavar. arcticaand
Wahlbergella
affnis.
The thirdterraceresembles
the second,but thereis no Cassiope;this
appearsagainon thefourth.Thesebeachesprovidean exampleofthesuccessionin timewhichoccurson raisedbeachesin Spitsbergen
iftheconditions
are sufficiently'
favourable
(comparewithshinglebeachesat BlakeneyPoint,
Norfolk
(54)). AtKlaas Billenand GipsValleysuchconditions
areonlyrarely
reached.
3. "FJAELDMARK."
The mainpartoftheslopesand summitplateauxis occupiedby a sparse
"fjaeldmark."Dryasand Cassiopeoccurin places,and herethevegetation
maybe called"cushion-heath"
("Polsterheide,"Cleve(9)). The listbelow
givesan idea ofthetypeofcommunity
whichoccupiesmostofthedrier,less
favourable
slopesin thispartofSpitsbergen.
Similarareasoccurat Advent
Bay and GreenHarbour.
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V. S. SUMMERHAYES AND
Phanerogams,etc.:
Cassiope tetragona Hf.
Catabrosa algida
Draba alpina
Dryas octopetala l.a.
Equisetum arvense
var. alpestreWg.
Festuca brevifolia
o.
Luzula confusa
r.
Oxyriadigyna
C. S. ELTON
279
Papaver nudicaulevar. radicatum f.
Pedicularishirsuta
P. lanata f. dasyantha
Poa abbreviataR.Br.
P. alpina
o.
Salix polaris
f.
Saxifragacaespitosa
f.-.a.
S; oppositifolia
o
Wahlbergellaaffinis
Otherrarerspeciesalso occur.
Lichens: Cetrarianivalisand probablyothers.
In many places the hardy Dryas is absent, scatteredrounded cushions of
Saxifraga oppositifoliaand caespitosaonly occurring(see Bear Island). The
Cassiope occursin slighthollowswheresnow collects. In some hollowson the
slopes Salix polaris becomes dominant. With it are foundSaxifraganivalis,
nitensand the
S. cernua,S. hieraciifolia,etc., also the moss Camptothecium
alpinum. Wherethe earthis finerSaxifragacernuais more
lichenStereocaulon
Ach. and Xanthoriaparietina
frequentand the lichensLecidea sanguineo-atra
occuras thincircularcrustson the almost bare soil.
The animalsofthe "fjaeldmark" werenot examinedcarefully.The spider
sobriusoccurs. At one time the neighbourhoodwas celebrated
Leptyphantes
forthe numberof reindeer,whichfed on the pasturesin Sassendale. Their
antlersare foundscatteredabout the slopes. The reindeeris now foundin
presenthere. Tho moss
Wijde Bay undersimilarconditionsto thoseformerly
Grev.and Arn.is apparentlylimitedto reindeerexcrement(5)
Voitiahyperborea
and its distributiondepends on this animal. The Snow Bunting nests in
crevicesbetweenrocks.
4. STREAMSIDE COMMUNITIES.
On the plateau the streamsrun in wide valleys,the whole of whichhave
a damp type of vegetation. A lopecurusalpinus is the dominant plant.
Papaver nudicaulevar. radicatumalso occursin quantity,whichis surprising
sinceit is usuallya plant of verydry habitats. Mosses occurheresubmerged,
Philonotisfontana and var. alpicola Jur. and Hypnumgiganteumbeing the
commonest.
On theslopesis a thinmarginalstripofmossesamongwhichgrowphanerogams. By one streamwere:
Phanerogams:
Cardaminebellidifolia
Cochleariaofficinalis
Draba subcapitata
Bryophytes:
Bryumobtusifolium
B. rutilansBrid.
Cinclidiumstygium
Saxifragaflagellaris
S. hirculus
S. rivularis
l.f.
Hypnumsarmentosum
Mniumaffinevar. integrifolium f.
f.
Orthothecium
chryseum
These mossesare similarto those by streamsin Iceland.
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280 Contributions
to Ecologyof Spitsbergenand Bear Island
F.
LESSER EDINBURGH ISLAND.
Thisis thesecondlargestislandofthegroup(seeFig. 3). Therawclimate
resulting
fromits proximity
to theGreenland
Sea is probablyresponsible
for
ofitstype,is characterised
ofthelifethere.Theisland,likeothers
the.poorness
bythelargenumber
ofsea-birds
whichnestthere,chiefly
theNorthern
Eider.
It cannotbe termeda true"Eider Holm" sincemanhas interfered
withthe
birdsduringthelast fewyears. EiderHolmsoccurnowonlyin remoteunvisitedregions
suchas LiefdeBay and WijdeBay.
On theshinglethereis no plantlifeexcepta fewtinyplantsofCochlearia.
out ofwhich
The mainpartoftheislandconsistsofa flatarea ofmoss-bog,
a fewlowgrassyhummocks
emerge.Thereare also a fewshallowpondsin
themaindepressions.
On thehillocksbesidesthegrass(notidentified,
but Catabrosa
algidaand
Poa arcticaR.Br.occuronthelargerisland)a fewplantsofSaxifraga
caespitosa
werefound.Thisis themainnesting
of
the
Eider
Ducks.
The
place
bogsare
almostentirely
occupiedby mosses,thefollowing
beingthecommonest
Hypnumpolygamum
H. uncinatum
Mniumaffinevar. integrifolium
Splachnumvasculosum
has a well-known
for maritimesituations
H. potygamum
preference
(49),
whilstSplachnum
dependsonthelargeamountofdungdroppedontheisland.
Thefollowing
birdsoccuron theisland:
ArcticTern
ArcticSkua
NorthernEider
BrentGoose
Purple Sandpiper
Dunlin (Erolia alpina alpina L.)
GreyPhalarope
Red-throatedDiver
Snow Bunting
Nestingon shingleand on moss-bog
o.
Nestingon shingleand on hillocks
Feeding. Nestingon the otherisland
o. Feeding
o. Feeding
Nestingamong moss and shingle
Nestingby ponds
Nestingamongstonesby shore
The Northern
Eider,BrentGooseand Diverfeedat sea. The wadersfeedon
land,and in thepondsas well.TwospeciesofDiptera(Metriocnemus
ursinus
and Cricotopus
glacialis)occur,whilethemiteScutovertex
lineatuswas found
undersmallstones.On thesurfaces
ofthepondswereverymanyCollembola.
the birdlifeofislandslikethe Edinburghs
The mainfactoraffecting
is the
absenceoffoxes. The BrentGoose,Eiderand Ternusuallyneston islands
forthisreason.The GlaucousGulland ArcticSkua are theirchiefenemies.
In ArcticAmericatheEiderand Geeseare sometimes
protected
by thefierce
attackswhichtheTernsmakeuponintruders
in defenceoftheirnests(58).
Thisappearsto be partlythe case in Spitsbergen,
a Skua oftenbeingseen
pursuedby a Tern.Skuas eat youngEidersand suck the eggsof various
birds.
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V. S. SUMMERIIAYESAND C.
S. ELTON
281
G. HERMANSEN ISLAND.
This island lies inside Foreland Sound, and enjoys a more favourable
climate than the EdinburghIslands. It is somewhatlarger,but of the same
type,and the same birdsoccur. In the centreis a rockyridgewhichis nearly
75 feetabove sea-levelin places. No detailedanimal notesweremade. There
are fivemain communitiesin the island:
1. MaritimeRegion.
2. " Fjaeldmark."
3. Rock Outcrops.
4. Wet Tundra.
5. Ponds.
REGION.
1. MARITIME
is similarto that in the Edinburgh
and
and
of
cliffs
consists
shingle,
This
Islands. On the AnserIslands the SpitsbergenPuffinand Mandt's Guillemot
and theyprobablyoccurhere. Ternsnestin numbers
nestin cleftsin the cliffs,
on the shingle.
2. "FJAELDMARK."
This is not of such a stunted characteras usually occurs owing to the
dungingeffectof the numerousbirds. It occurs on the slopes betweenthe
bogs and the rocks. The groundis verydry,withmoresoil than usual. Saxiis the chiefplant. So profuselywas thisbloomingthat from
fraga oppositifolia
afar offit resembledsomewhat a floweringheather-moor.The few other
speciesare theusual " fjaeldmark" ones. A fewEider Ducks nestin thisregion.
3. ROCK OUTCROPS.
This area is concentratedin the ridge,but small rockyareas occurat the
tops of the cliffsand elsewhere. Here the richestvegetationoccurs,due to
the largenumberof Eiders whichnest on the ledgesof,or between,the rocks.
The followingplants occur:
Phanerogams:
AlsinebifloraWg.
Cardaminebellidifolia
Cerastiumalpinum
Draba alpina
D. alpina var. oblongata
D. hirtavar. arctica
D. wahlenbergii
Dryas octopetala
Pedicularishirsuta
PotentillaemarginataPersh.
Lichens:
Alectorianigricans
r.
r.
f.
f.
f.
0.
o.
l.d.
r.
o.
Ranunculuspygmaeus
R. sulphureus
Salix polaris
Saxifragacernua
S. caespitosa
S. hieraciifolia
S. nivalis
S. oppositifolia
S. rivularis
r.
f.
f.
o.
f.
r.
o.
f.
r.
Othermossesand lichensalso occur. On small isolated rockoutcropsa mosslichen vegetationoccurs dominatedby Rhacomitrium
lanuginosum,Cetraria
Here
a
few
Eider
Ducks nest.
and
Cladonia rangiferina.
only
nivalis
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282 Contribttionsto Ecology of Slitsbergenand Bear Island
4. WET TUNDRA.
This consistsof moss-bogsand pond marginalcommunities.The ponds
occur as a ringnear the edge of the island. The bogs are chieflymossy,but
there are a few phanerogamslocally, Salix polaris and Saxifraga rivularis
beingthe moreimportant. The mossesare the usual bog species. Significant
in connectionwith the extra nitrogensupply is the presenceof Ceratodon
purpureusand Tetraplodonwormskjoldii.The moss florais similarto that at
Cape Boheman. The Grey Phalarope and Red-throatedDiver nest around
the ponds.
5. PONDS.
The ponds are of the usual type,and are shallow. The animals were not
examined. Large masses of Nostococcurin places,whileHypnumcordifolium,
H. giganteumand H. stramineum
growsubmerged.
H. ADVENT BAY.
Thisarea was visitedon severaloccasions,but onlyforshortperiods. There
are a numberof species,both of plants and animals,whichare not recorded
fromother places, since Advent Bay and Coles Bay show more favourable
conditionsof lifethan do manyotherparts of Spitsbergen.
At AdventBay thefollowingwerefoundon flowersofDryas and Cerastium
in the lateralvalley behindLongyearCity:
Diptera:
Camptocladiuspumilio
Limnophoramegastoma
Psectrocladiusborealis
Rhamphomyiacaudata Zett.
Sciara sp.
Syrphustarsatus
The larvae of Syrphusfeedon aphids,ofwhich one species-Scaeva dryadisis recordedfromAdvent Bay. No Hymenopterawere seen by us here, but
severalspecieshave been recorded. An ichneumonBassus aroticusis parasitic
on the aphid. Pristiphorafrigida
occursand is parasitisedby Ichneuteshyperboreus.Several otherichneumonsoccurhere.
The birds resemblethose of Gips Valley except for the presenceof the
Barnacle Goose (Branta leucopsis),whichnests on cliffsin the valley. A flea
Ceratophyllus
vagabunduswas found abundantly in the down of Barnacle
This
also occursin the nestsofPink-footedGeese. The bird-louse
Goosenests.
TrinotonanserinumFab. was foundon the Barnacle.
IV. SUMMARY AND CONCLUSIONS.
1. The plantand animalcommunitiesofpart ofBear Island and ofvarious
districtsin Spitsbergenare described.
2. A successionofplant communitiescan be traced,startingwith"fjaeldmark'" on unstable ground,such as screes,or on recentlyelevated land, as
raised beaches,and terminating
onistabilisedgroundwitha heath vegetation
of Ccassiopetetragona,
Dryas octopetalaor Salix polaris. In most places this
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282 Contribttionsto Ecology of Slitsbergenand Bear Island
4. WET TUNDRA.
This consistsof moss-bogsand pond marginalcommunities.The ponds
occur as a ringnear the edge of the island. The bogs are chieflymossy,but
there are a few phanerogamslocally, Salix polaris and Saxifraga rivularis
beingthe moreimportant. The mossesare the usual bog species. Significant
in connectionwith the extra nitrogensupply is the presenceof Ceratodon
purpureusand Tetraplodonwormskjoldii.The moss florais similarto that at
Cape Boheman. The Grey Phalarope and Red-throatedDiver nest around
the ponds.
5. PONDS.
The ponds are of the usual type,and are shallow. The animals were not
examined. Large masses of Nostococcurin places,whileHypnumcordifolium,
H. giganteumand H. stramineum
growsubmerged.
H. ADVENT BAY.
Thisarea was visitedon severaloccasions,but onlyforshortperiods. There
are a numberof species,both of plants and animals,whichare not recorded
fromother places, since Advent Bay and Coles Bay show more favourable
conditionsof lifethan do manyotherparts of Spitsbergen.
At AdventBay thefollowingwerefoundon flowersofDryas and Cerastium
in the lateralvalley behindLongyearCity:
Diptera:
Camptocladiuspumilio
Limnophoramegastoma
Psectrocladiusborealis
Rhamphomyiacaudata Zett.
Sciara sp.
Syrphustarsatus
The larvae of Syrphusfeedon aphids,ofwhich one species-Scaeva dryadisis recordedfromAdvent Bay. No Hymenopterawere seen by us here, but
severalspecieshave been recorded. An ichneumonBassus aroticusis parasitic
on the aphid. Pristiphorafrigida
occursand is parasitisedby Ichneuteshyperboreus.Several otherichneumonsoccurhere.
The birds resemblethose of Gips Valley except for the presenceof the
Barnacle Goose (Branta leucopsis),whichnests on cliffsin the valley. A flea
Ceratophyllus
vagabunduswas found abundantly in the down of Barnacle
This
also occursin the nestsofPink-footedGeese. The bird-louse
Goosenests.
TrinotonanserinumFab. was foundon the Barnacle.
IV. SUMMARY AND CONCLUSIONS.
1. The plantand animalcommunitiesofpart ofBear Island and ofvarious
districtsin Spitsbergenare described.
2. A successionofplant communitiescan be traced,startingwith"fjaeldmark'" on unstable ground,such as screes,or on recentlyelevated land, as
raised beaches,and terminating
onistabilisedgroundwitha heath vegetation
of Ccassiopetetragona,
Dryas octopetalaor Salix polaris. In most places this
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V. S. SUMMERLIAYES AND) C.
S. EI;roN
283
climax is apparentlynever reached, owing to contintued
instabilityof the
substratumi,and "fjaeldmark" is retained.
thedrainiageis bad, or wherethereis a conistant
3. Wlihen
supplyofwater,
damipercomnmunities
are established,and may be maintaiined
indefinitely.
4. The comparativeabuindanceof cryptogamls,
especiallythe Bryophyta,
is a characteristic
featureofthevegetation.The relativeincreaseofthecryptogams in the communities
in maritimeregionssuch as Bear Island and Prince
CharlesForelalndas comparedwithdistrictsat the heads offjordscan be seen
fromthe lists,etc. This is probablydue to the greaterhumidityof the air on
the coasts.
a. The effectof abundantsupplyof nitrogenon the plalntsis pointedout
and special communities
dependingonlthis extrasupplyare described. These
communitiesagree to a great extentwith those describedfromotherarctic
countries.
6. In bothplantsand aniimalsthe numberof speciesis small. Thisis due to
(a) severeconditions,(b) geographicalisolation,(c) theabsenceofday and night
as separatehabitatsso thatfewerspeciesofanimalscan live in the same area.
7. The animal species are not usually conifined
to any particulargeneral
(or "'communal") habitat,plant associationor plant species,but occur in or
on more than one. Individuals may occur in more than one communityat
different
times(e.g. birds,flies). Thus "communities"of animals exist only
in the sensethat each generalhabitatand each plant communityhas a fairly
constantset of animal species associated with it, of which only a few are
exclusive(i.e. confinedto it).
8. As a resultofthisthebestmethodofdescribingtheanimalcommunities
appears to be by basingtheaccountlargelyon1thefoodrelations.The method
of Shelfordand others,based on the "physiologicalresponse" of the animal
to the habitat,althoughindispensable,is inadequate,since it only applies to
a fewspeciesin each community(i.e. the exclusite oines),and these oftenare
notveryimportantin thelifeofthe community.
9. The foodcycleon Bear Island is sketched. The chiefpointsare: (a) the
additioniof food by sea-birdsand probablyby nitrogenfixationby bacteria,
(b) the loss offoodby birdmigrationanidwashingdownofsolublesalts,(c) the
importanceof freshwater
fliesas foodsupplyforland animals,(d) the "short"' of the "nitrogencycle" owingto the activityof scaveengers
circuiting
which
feedon decayinganimal imatter
is not ofimportance,(e) the scarcityof carnivores in the freshwater modifiesthe usual positionof the Entomostracaas
time"key-iniduistry"
of freshwater. These coniclusions
apply on the whole to
Spitsbergenalso.
10. In ainimalsverticaldifferences
in habitat are relativelyunimportanlt
or absenit(except" exposed,"" understonesanidplants,"anid" subterraneail
").
The s-amemnay
be said about the planitssinicethereare no treesor slhrubs,
anid
veryfewdwarf-shrubs.
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284 Contributions
to -Ecologyqf Spitsbergenand Bear Island
11. A series of brackishand freshwaterponds at Klaas Billen Bay is
described.The conditionsand successionofcommunitiesdue to land elevation
are dealt with. In particular,an account of the ecologyof certaincrustacea
(Mysis relicta,Gammaruszaddachi,Eurytemoraraboti)is given. A sketch of
the probable mannerin whichMysis relictabecomesrelictin freshwater is
given. The biota of freshwaterponds containingrelictEurytemorarabotiis
describedand also temperaturerecordsfromone pond.
12. The abundance of enchytraidworms in the soil in Spitsbergenis
investigatedand theirimportanceindicated.
V. BIBLIOGRAPHY.
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Beeren Eilands KirlvXxtflora."Bihang tillK. Svensk. Vet.-Akad.Handl. 26, 3, 1900.
(2) Andersson, J. G. "Solifluction,a componentofsub-aerialdenudation."Journ.Geol.1906.
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Svensk. Vet.-Akad.Handl. 26, 3, 1900.
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Vet.-Akad.Handl. 26, 3, 1900.
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(12) Conway, M. Early Dutchand EnglishVoyagestoSpitsbergen
in the17thCentury.London,
1904.
(13) Conway, M. No Man's Land. Cambridge,1906.
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Handl. 27, 3, 1901.
(15) Dusen, P. "Zur Kenntnisder Geffisspflanzen
Ostgrdnlands."Ibid.
(16) Farrow, E. P. " On the Ecology of the Vegetationof Breckland." Journ.Ecol. 6, 2, 1918.
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(18) Fries, Th. " Om BeerenEilands Fanerogam Vegetation."Offvers.
af K. Svensk. Vet.-Akad.
F6rhandl.1869.
(19) Friese, H. D. JFaunaArctica,2, 3. "Die arktischenHymenopteren."1902.
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Copenhagenand London, 1920.
(21) Gordon, Seton. Amid Snowy Wastes. London, 1922.
(22) Haberbosch, P. "Die SiisswasserEntomostrakenGrdnlands." Zeitschriftfilr
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1 Jahrg. Basle, 1920.
(23) Hart, C. "On the Botany of the BritishPolar Expedition." Journ.Bot. Series 2, 9, 1880.
(24) Hartz, N. "Ostgronlands Vegetationsforhold."Medd. om Gronland,4, 1895.
(25) Hesselbo, A. "The Bryophytesof Iceland." The Botany of Iceland, Copenhagenand
London, 1918.
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ELTON
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of WildLife in Canada. New York, 1921.
(26) Hewitt, C. Gordon. The Conservation
(27) Hdgbom, B. "Einige Illustrationenzur den geologischenWirkungendes Frostes auf
Spitzbergen." Bull. Geol.Inst. Univ. Upsala, 9, 1908-9.
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(29) Johansen, F. "General Remarkson theLife ofInsectsand Arachnidsin N.E. Greenland."
Medd. mnGronland,43, 1911.
(30) Johan$en, F. "FreshwaterLife in N.E. Greenland." Medd. om Gronland,45, 1911.
(31) Jonsson, H. "The MarineAlgal Vegetation." The Botanyof Iceland,Part 1. Copenhagen
and London, 1912.
32) Jourdain, F. C. R. "The Birds of Spitsbergenand Bear Island." Ibis, 4, 1, 1922.
Studienaus RussischLappland. Helsingfors,1890.
(33) Kihlman, A. 0. Pflanzenbiologische
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Wissen.St Petersburg,1902.
(35) Lagerheim, G. " Beitragezur Flora derBaren-Insel. 2. VegetabilischesSiisswasserPlankton aus der Biren-Insel." Bihang till K. Svensk. Vet.-Akad.Handl. 26, 3, 1900.
(36) Lamont, J. Seasons withtheSea-Horses. London, 1861.
Exped.
(37) Lilljeborg, W. "Entomostracen wahrend der schwedischenwissenschaftlichen
der Jahre 1868, 1898, 1899 auf der Biren-Inselgesammelt." Bihang till K. Svenslk.Vet.Akad. Handl. 26, 4, 1900.
theVegetationofGermaniaLand, N.E. Greenland."
(38) Lundager, A. "Some Notesconcerning
Medd. om Gronland,43, 1912.
(39) Manniche, A. L. V. "The TerrestrialMammals and Birds of N.E. Greenland." Medd.
om Grdnland,45, 1912.
(40) Middendorf, A. von. "Die Gewichse Sibiriens." SibirischeReise,4, 1864.
(41) Nansen, F. FarthestNorth,1. London, 1898.
(42) Nansen, F. FarthestNorth,2. Ibid.
(43) Nathorst, A. G. "Beitrage zur Geologie der Bairen-Insel,Spitzbergensund des KiinigKarl-Landes." Bull. Geol.Inst. Univ. Upsala, 10, 1910.
(44) Nathorst, A. G. "Nya bidrag till kiinnedomen om SpetsbergensKdrlviLxter,och
dess ViixtgeographiskaforhOllenden."Bihang till K. Svensk. Vet.-Akad.Handl. 20, 3,
1883.
(45) Nathorst, A. G. "Studien uiberdie Flora Spitzbergens." Engler'sJahrb.4, 1883.
(46) Nordenskjold, A. E. The Voyageof the Vega. London, 1885.
Ectinosomidae,Cantho(47) Olofsson, 0. "Beitrag zur Kenntnisder Harpacticiden-Familien
camptidae...undTachidiidae..... Zool. Beitrdgeaus Uppsala, 6, 1918.
(48) Olofsson, 0. "Studien iiberdie SiisswasserfaunaSpitzbergens." Ibid.
(49) Ostenfeld, C. H. "The Land-Vegetationofthe Far6es." The BotanyoftheFar6es, Part 3.
Copenhagen,1908.
omn
(50) Porsild, M . P . " The Plant LifeofHare Island offthecoast ofWestGreenland."Mlledd.
47, 1911.
Grdnland,
(51) Rasmussen, K. GreenlandbythePolar Sea. London, 1921.
(52) Richard, J. "Sur la fauna des eaux douces exploreesen 1898...." Me'm.Soc. Zool. France,
11, 1898.
(53) Rudmose Brown, R. N. 'The Flora ofPrinceCharlesForeland." Trans. Bot.Soc. Edinbutrgh,
23, 4, 1908.
(54) Salisbury, E. J. "The Soils of BlakeneyPoint: a study of soil reactionand successionin
relationto the plant covering." Ann. Bot. 36, 1922.
(55) Schaudinn, F. Fauna Arctica,2, 1. "Die Tardigraden." 1902.
(56) Schroter, C. Das PfanzenlebenderAlpen. Zurich,1908.
(57) Soar, C. D. "A species of Hydrocarinafoundat Bear Island, June17th,1921." J. Quekett
Micr. Club,14, 1922.
(58) Stefansson, V. My Life withtheEskimo. London, 1913.
(59) Stefansson, V. The FriendlyArctic. London, 1921.
(60) Stephenson, J. Proc. Zool. Soc. London,4. 1922.
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286 Contributions
to Ecology of Spitsbergenand Bear Islawd
(61) Stephenson, K. " On a Collectionof GammarusfromRander's Fjord, Denmark." Comm.
2, 1916.
of Zool. Mus. Copenhagen,
fromDept. of Arthropods
(62) Swenander, G. " BeitrigezurFauna derBdren-Insel.1. Die V6gel." BihangtillK. Svens8k.
Handl. 26, 4, 1900.
Vet.-Akiad.
(63) Thoroddsen, Th. "An Accountof the PhysicalGeographyof Iceland." The Botanyof
Iceland. Copenhagenand London, 1914.
derNederid.
(64) Van Oordt, G. J. "OrnithologicalNotesfromSpitsbergen...1921." Tydschft.
Ornith.Vereeng.1921.
(65) Wahlgren, E. " BeitrdgezurFauna derBdren-Insel.4. Collembola." BihangtillK. Svens8k.
Vet.-Akhad.
Handl. 26, 4, 1900.
(66) Walton, J. "A SpitsbergenSalt Marsh...." Journ.Ecol. 10, 1, 1922.
(67) Warming, E. "Uber Gr6nlandsVegetation." Engler'sJahrb.10, 1888-9.
(68) Warming, E. (EcologyofPlants. Oxford,1909.
(69) Waterston, J. "Results of OxfordUniv. Exped. to Spitsbergen,1921. No. 25. HymenNat. Hist. Series 9, 11, 1922.
optera Parasitica;gchneumonoidea." Ann. and M11ag.
(70) Wulff, Th. "ObservationsBotaniques,faitesau Spitzberg." Miss. Scient.pour la Mesure
d'un Arc de Me'ridienau Spitzbergen 1899-1902. Sect. 10, Botanique. Stockholm.
NOTICE OF P UBLICA TION OF
GENVERAL BEARING
AN INTRODUCTION
TO PLANT ECOLOGY
text-figures.
Tansley,A. G. Practical Plant Ecology. Pp. 228, with fifteeni
London, GeorgeAllen and Unwin, 1923. Price 7s. 6d.
The spread of ecological interestin Britain suffers-and has sufferedfor a long timefromthe absence of any text-book dealing with ecological methods, which might be placed
in the hands of those unfamiliarwith these methods. Such a book to be of real value requires
to combine a certain felicityof expression with a rare discriminationas to subject matter.
It needs to be compact in form,moderate in price and yet to cover adequately the variety
of material and method which is included in the scope of plant ecology. It may safely be
said that fromall of these standpoints Mr Tansley's book is an admirable one. While it is
necessarilyconcerned largely with plant communities,plant ecology is viewed as "a means
of approach to a large part of detailed botanical study" rather than as a name fora special
branch of botany, and in this the author expresses the modern reaction from the purely
laboratory (and academic?) aspects of plant life.
The book is divided into five parts, of which the firstis introductory,while the seconld
definesthe units of vegetation and gives outlines of plant succession and of the main types
of British vegetation. The third part deals with methods of studying and describingvegetation in the field,and this is followed by chapters cn the habitat and the estimation of
habitat conditions. The remaining part discusses the opportunitiesfor ecological work in
schools with sugfgestionsas to profitablelines of attack. Useful appendices are added on
the "life forms" of plants, methods of surveyingand photographingvegetation,soil analysis
(by Dr H. J. Page) and the determinationof hydrogen-ionconcentration.There is a classifiecd
list of books and papers suitable forfurtherreference.
The book well fulfilsthe purpose for which it was written,as a "guide for beginnersill
field study of plant communities," and it is difficultto make any suggestion which would
improve it without greatlyextendingits scope. It will prove verv useful to students and to
those desirous of embarking on field or ecological work. At the same time it is interesting
W. H. P.
and suggestive,even to more experienced ecologists.
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266
Further Contributions, to the -Ecology of Spitsbergen
in another paper, in which will be found a general discussion of the means of
dispersal of the various invertebrates (6): in the summer of 1924, vast swarms
of hover flies (Syrphus ribesii) and spruce aphids (Dilachnus piceae) were
blown from Northern Europe on to the ice-cap of North-East Land-a distance
of over 800 miles. They were alive when they arrived, but perished later in
a blizzard; in any case they could scarcely have survived very long. A similar
swarm of aphids was encountered by Parry off the west of North-East Land
in 1827 (24, p. 201). It is to be supposed that the other insects reached the
archipelago by similar means, or by transport on birds. The result of the
isolated position of Spitsbergen is that many groups of animals are absent,
although these are found in Greenland under similar conditions of climate
and vegetation. Examples are various butterflies, also bumble-bees and tipulid
flies.
Furthermore, several mammals are absent, e.g. the Lemming, Arctic Hare,
and Musk Ox. The absence of Lemmings is reflected in the almost complete
absence of Snowy Owls, and the fact that there are no Ermine (which eat
the Lemmings); nor are there any Arctic Wolves (which eat Hares). Since the
wolves also attack deer, it is probable that their absence is due rather to
dispersal difficulties than merely to absence of food. These questions, in so
far as they affect the mammals, will be treated in a separate paper.
The fauna of Spitsbergen differs therefore from that of similar places in the
same zones of climate and vegetation, in being rather more impoverished.
We have already (Section VIII) traced the gradual increase in complication of
the food-cycle as the climate improves, and have shown how the most highly
developed animal communities in Spitsbergen foreshadow in a general way the
very complicated communities of sub-arctic and temperate regions. At the
present time we know very little about the animals of the sub-arctic zone
(Empetrum, etc.) and it is to be hoped that future work in some place like
South Greenland or Lapland will bridge the gap between our knowledge of
the food-cycles of high arctic countries and those of our own regions.
X. SUMMARY.
1. In 1923 we published in this Journal an account (almost entirely
descriptive) of the animal and plant communities of certain parts of Spitsbergen
and Bear Island. Further investigations in Spitsbergen have enabled us to
co-ordinate these and our later observations into a general scheme, which
makes it possible to explain the general distribution of the plant and animal
communities in terms of four master factors: climate (insolation and airtemperature), soil, manuring by sea-birds, and water-supply.
2. The climatic gradient produced by the meeting of the Polar ice-pack
with the Gulf Stream, brings about a corresponding gradient in the types of
vegetation. This gradient can be divided naturally into four zones, (1) Barren
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V. S. SUMMERHAYESANDC. S. ELTON
267
Zone, (2) Dryas Zone, (3) Cassiope Zone, and (4) the Inner Fjord or ETnpetrum
Zone.
3. This gradient in plant communities has also been traced in the communities of land-animals, and in the intertidal fauna. By combining all these
lines of evidence, we have constructed a provisional map showing the main
life-zones for the whole of the Spitsbergen archipelago.
4. These zones can be traced on a much broader scale right across Greenland and Arctic Canada, and in the mouiitains of northern Scandinavia; and
they demonstrate the very high-arctic character of Spitsbergen communities.
5. The gra(dientoutlined above is interfered with by bird-manuring, which
produces distinctive (andcusually grassy) communities even in the most barren
parts of Spitsbergen. The effects of bird-manuring on animals and plants were
studied intensively in several cases.
6. It is impossible in a short space to summarise any of the other poiilts
dealt with, and the reader may be referred to Sections VIII and IX, and to the
map, since these are to some extent summaries of the rest of this paper.
XI.
REFERENCES.
For a more complete list of works dealing with the flora and fauna of Spitsbergen, the reader
is referred to our previous paper (29).
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H. "Verzeichnis der in K6nig Karls Land wiihrend
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(7) Elton, C. S. "The Oxford University Arctic Expedition, 1924: the Biology in Relation to
the Geography." Geogr.Journ. 66, 111-114, 1925.
(8) Elton, C. S. "The Nature and Origin of Soil-Polygons in Spitsbergen." Quart. Journ. Geol.
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(9) Elton, C. S. Animal Ecology. London, 1927.
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(13) Holmgren, A. E. "Bidrac till k-annedoinenom Becren Eilands och Spetsbergens InsektFauna." Kongl. Svensk-. tet.-Akad.JJandl. 8, 58, 1869.
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268 Futrther Contributions to the Ecology of Spitsbergen
(14) Holttum, R. E. "The Vegetation of West Greeiilanid." This JOURNAL, 10, 1, 1922.
(15) Iversen, T. "Hopen (Hope Island), Svalbard." Resultaterav de Norske Statsunder8t0ttede
Spitsbergenekspeditioner,1, No. 10, 1926.
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et Flora Fenn. Helsingfors, 1890.
(17) Kukenthal, W. Petermanns Mlitteil.36 (1890).
(18) Longstaff, T. G. "Notes from Spitsbergen." Ibis, July 1924, 480-495.
(19) Lundager, A. "Some Notes concerning the Vegetation of Germania Land, N.E. Greenland." Medd. om Gronland,43, 1912.
(20) Marret, L. Icon. Flor. Alpin. Plant. Ser. II, Fasc. 6, P1. 259. Paris, 1913.
(21) Montague, F. A. "Further Notes from Spitsbergen." Ibis, Jan. 1926, 136-151.
(22) Nathorst, A. G. ' Nya bidrag till kiinnedomen om Spetsbergens Kiirlvixter, och dess
Vaxtgeographiska forhAllenden." Kongl. Svensk. Vet.-Akad. Handl. 20, 3, 1883.
(23) Nordenskiold,
A. E. Arctic Voyages. London, 1879.
(24) Parry, W. E. Narrativeof an attemnptto reach the NVorth
Pole. London, 1828. Appendix by
W. J. Hooker.
(25) Raunkiaer, C. ' Livsformerns Statistik somn Grundlaa for biologisk Plantegeografi."
Bot. Tidsskr. 29, 42-83, 1908. (German translation in Beih. Bot. Centralbl. 27, Abt. 2,
171-206, 1910.)
(25a) Resvoll-Holmsen,
H. Exploration du Nord-Ouest du Spitsberg.. par la Mission Isach.
sen, Fasc. XLIV, Pt. 5 (1913).
(26) Sandford, K. S. "Summer in North-East Land, 1924: the Climate and Surface Changes."
Geogr.Journ. 68, 200-225, 1926.
(27) Schr6ter, C. Das Pflanzenlebender Alpen. Ed. 2. Zurich, 1926.
(28) Stephenson, J. "The Oligochaeta of the Oxford University Expedition." Proc. Zool. Soc.
1922, 1109.
(29) Summerhayes,
V. S. and Elton, C. S. "Contributions to the Ecology of Spitsbergen
and Bear Island." This JOURNAL, 11, 214-286, 1923.
(30) Tyrrell, J. W. Across the Subarctics of Canada. Toronto, 1897. Appendix, pp. 260, 265.
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