Skip to main content

Full text of "Arbejder fra den Botaniske have i Kbenhavn"

See other formats 


er 
tits 
4 


iik 


HR: 


i 
tk 


svAr 


SEN 


tilsat sge! i Mg fn % £ 
BAR HER HELET 
ANSETE SEERNE SLET 
' His lslig sli: Ci 
i i HEE 
i Ht 


nd 


il 


H 
i 


HH 
Hi 


STS T- 
GRåtaS 
ST STS Tr 2337 2T 


33153333 


…se72327 


3227 
SIKRET 


tri 
men enene 
ISS TET 


737 





"aernd wo WA! 








LNG HAN 


BUDVA 


yd 
KN 
MÅ 


BAN: 


ro 
AND Cs i 
SÅ ML (Xl 



















 SÆRTRYK 2 2 


af ,,Danmarks Havebrug 
og Gartneri til Aaret 1919" 














Arbejder fra Botanisk Have i København Nr. 93 


mM, 


BES 
bre 


« 


6 


Kg 
DÅ 
== 
£ 





BOFAÅNISKE”HAVER 


AF 
BOTANISK GARTNER AXEL LANGE 





EN Interesse, man i gamle Dage nærede for Planterne som de 

fornemste Lægemidler, har uden Tvivl foraarsaget, at Lægerne 
for ret at kunne siges at forstaa deres Videnskab maatte give sig af 
med Botanik, og det var da ikke underligt, at man søgte at skabe 
Plantesamlinger, hvor Studiet af saavel vildtvoksende som dyrkede 
Urter kunde ske i Ro og Mag, og hvor Planterne var helliget dette 
Formaal alene. Ligeledes er det forstaaeligt, at Institutioner eller 
Privatpersoner efterhaanden som Kommunikationen med fremmede 
Lande forbedredes og Kendskabet til disse øgedes, fik Lyst til at 
prøve Dyrkningen af-disse fjærnere Landes Planter. 

Nogen Efterretning om botaniske Haver har man ikke førend en 
Menneskealder ind i det 16. Aarhundrede. Det har ofte været tillagt 
Universitetet i Padua Æren af at have stiftet den første botaniske 
Have anlagt 1545 — men Eichhorn mener dog i sin ,,Geschichte 
der Literatur" (I, S.304), at Euricius Cordus allerede i 1534 i 
Marburg anlagde en privat botanisk Have. Uden Tvivl er Universite- 
terne dog gaaet foran i denne Bevægelse og først senere er det blevet 
almindeligere, at Fyrster, Kommuner eller private Personer har taget 
Tanken op. Bolognas Universitet fik i 1568 sin botaniske Have og 
Leydens Universitet kom efter i 1577, medens Aarhundredet skulde 
løbe til Ende, før vi her i Danmark kunde glæde os ved Besiddelsen 
af en saadan Anstalt. 











Fig. 123. Plan over Universitetskareen med Botanisk Have, 1757. 
A. Auditorium. B. Botan. Have. C. Konsistorium. 247. Professorbolig. 


KØBENHAVNS 
UNIVERSITETS FØRSTE BOTANISKE HAVE 


(HORTUS MEDICUS) . 


Aaret 1600 skulde der bygges en ny ,,Residentz" i Studiegaarden 

som Bolig for en af Professorerne, og det blev da besluttet, at et 
Jordareal, ,,220 Fødder i Længden og 110 Fødder i Bredden”, hvil- 
ket fra først af havde været bestemt til Opførelse af Enkesæder for 
Præsteenker, skulde udlægges til en botanisk Have. I et ,,Kongeligt 
Skiøde og Gave-Brev" af 2den August 1600 læser vi: ,,Og efterdi for- 
nævnte Plads udi saa lang en Tiid haver lagt øde, og ikke derpaa er 
faaet nogen Bygning; Saa og efterdi Universitetet der udi fornævnte 
vor Kiøbstæd Kiøbenhavn er med en nye Residentz blevet forbedret; 
Thi have vi naadigst undt, skiøt og givet, og nu med dette Vort 
aabne Brev unde, skiøde og give formeldte Plads til Universitetet der 
sammesteds, saa det altid herefter skal ligge og bruges til en Have 
til formeldte Residence, hvorudi kan ympes og plantis synderlige 
Simplicia, og skal den Professor, som i formeldte Residentze bli- 
ver boendes, dermed have Opseende.” 

Haven laa ud mod Skidenstræde — den nuværende Krystalgade 
— omtrent paa den Plads, hvor nu Zoologisk Musæum ligger. Der 
blev ikke bevilget Midler til at holde Haven, hortus botanicus eller 
hortus medicus, vedlige med; ,,thi da man af førstningen formodent- 
lig allene har tænkt deri at indtage vores vilde Planter, har man 
troet, at dertil ingen Udgifter behøvedes og derfore ey heller tillagt 
den Indkomster" (Rottbøll). Der var heller ingen Urtegaardsmand 
eller Gartner ansat derved; den Professor, som fik tildelt Embeds- 
boligen, maatte tillige vedligeholde Haven og af egen Lomme betale 
de paaløbende Udgifter. Det har ikke al Tid været de medicinsk-bo- 


BOTANISKE HAVER 301 


taniske Professorer ved Universitetet, som har faaet Boligen tildelt 
og dermed Forpligtelsen til at have Tilsyn med Haven, hvilket vilde 
have været rimeligt, men de Professorer, som benyttede Haven, har 
sikkert haft en vis Indflydelse paa Driften. 

Den berømte Ole Worm har i sin Tid haft Haven under sig, 
ligesaa har Rasmus Caspar Bartholin haft den i sin Varetægt. 
Sidstnævnte nærede saa megen Interesse for den, at han skænkede 
1200 Kronedalere til et Legat (stiftet 5. Dec. 1696, konfirmeret 8. 
Nov. 1698), ,,hvoraf dend Aarlig Rente skal bruges til hielp at holde 




























F—————— z = = == 
Eltiden OJtrædel 
== 
3 T mk, = 
3 F—= Føns, TI Ek 





væ 




































































Grund fegning 


over 





























Cen obolanickeHatige, med 
hosfogede Gerxedt Has og Drive . 


hlitifse . ele 




















$ 
& 
d 
s 






































Fig. 124, 


en Urte Gaards-Mand for in Horto Botanico, samt at kiøbe frem- 
met frøe og Planter." Det synes, som om Haven delvis har været 
anvendt til Lysthave for den derboende Professor. I al Fald klager 
Johannes de Buchwald i 1723 over, at ,,vores hortus botanicus 
haver til dato haft for lidet rom”, og foreslaar, at Haven bliver ,,;med 
et Planckeværck afdeelt, saa at den Hr. Consistorialis, som opterede 
samme Residence, maatte have dend deel af haven, som er neest bag 
huuset, og dend anden part, som ligger mod skidenstrædet, hørte til 
horti botanici brug.” Joh. de Buchwald udgav i 1720 en paa Latin 
skrevet Bog: ,,Specimen Medico-Practico-Botanicum" med Beskri- 
velse af en Del Urters Anvendelse oversat paa Tysk af hans Søn B al- 
thasar Johannes de Buchwald, 1721, af hvilket Værk det 
fremgaar, at Planterne i den botaniske Have er bleven omplantede i 
Aaret 1719 og anbragte i alfabetisk Orden. 


302 DANMARKS HAVEBRUG OG GARTNERI 


Efter Ildebranden i København 1728 blev Haven noget formind- 
sket i Åreal paa Grund af de omliggende Gaders Regulering. Haven 
trængte saaledes allerede den Gang til at blive flyttet til en rumme- 
ligere Plads, men der skulde dog hengaa over 40 Aar inden nogen 
Forandring i denne Henseende skete. I dette Tidsrum hører man 
ikke meget til Haven, og med de Midler, der stod til Havens Dispo- 
sition, lod sig heller ikke meget udrette; 1655 hører vi dog, at Haven 
havde en Gartner, som fik 30 Rdl. i aarlig Løn. I 1769 begyndte det 
at dages for Haven. Christian VII skænkede nemlig ved Reskript 
af 3. Februar 1769 2500 Daler til Universitetet, ,,hvoraf Renten 
skulde anvendes til Havens ,,Istandsættelse og bestandige Dyrkelse.” 
Og nu var der endda Udsigt til Udvidelse paa Stedet, idet Kongen 
1770 lovede at ville øge den botaniske Have med Communitetets 
Have og ved indtrædende Vakancer tillige med de Haver, som Sogne- 
præsten og 1ste Kapellanen ved Frue Kirke sad inde med. 

Det er sandsynligt, at Christen Friis Rottbøll, som var 
Professor i Medicin ved Universitetet og som nylig ud mod Skiden- 
strædet havde faaet opført et Væksthus af 16 Fods Længde og 8 
Fods Bredde, med Glæde har imødeset den i Udsigt stillede Udvi- 
delse, men iøvrigt har Universitetets Styrelse næppe billiget Planen, 
i al Fald kommer den ikke til Udførelse. Kongen fremkommer nu 
med et nyt betydeligt Tilbud, idet han s. Å. skænker Universitetet 
en Del af Oeders botaniske Have ved Amalienborg (hvorom mere 
senere) med samt Bibliotheket, dertil 5000 Kr. til Istandsættelse og 
aarlig 300 ØRbd. af Partikulærkassen til Vedligeholdelse. Rottbøll er 
imod Flytningen og udtaler sig saaledes om Universitetshaven: 
»jeg kan calculo Mathematico bevise, at der med al Magelighed kan 
staae over 2000 Spec., som er alt det den upsalske Have har," men 
Universitetet modtager dog Gaven og allerede samme Aar sker Over- 
tagelsen. Universitetshaven blev dog i Brug endnu i 8 Aar, hvorefter 
den blev overdraget som en Lysthave til den Professorbolig, som 
den havde været knyttet til. 

Af denne ældste botaniske Have i Danmark eksisterer to Planer. 
Den ene, som er gengivet her i Fig. 123, viser Haven, saaledes som den 
saa ud i 1757. Den anden Plan — Fig. 124 — maa formodes at være 
tegnet mod Slutningen af Havens Eksistens; det synes som den fore- 
liggende Plan er Forslag til Udvidelse af Haven. Figuren her er re- 
produceret efter en Tegning, som findes i Raadhusarkivet, rimeligvis 
en Kopi af ret ny Dato efter en ældre Tegning i det store Kongelige 
Bibliothek. 


KØBENHAVNS 
UNIVERSITETS ANDEN BOTANISKE HAVE 


(OEDERS HAVE) 


ET var en Have i Drift, som Professor Rottbøll kom til at over- 

tage, og selv om han ikke havde haft videre Sind til at komme 
bort fra Universitetshaven, har han dog sikkert snart fundet sig til 
Rette paa det ny Sted. (Haven, der overtoges, var anlagt af G. C. 
Oeder paa Foranledning af Frederik V, men om denne Haves tidligere 
Historie, vil der senere under Afsnittet Kongelige botaniske Haver 
blive givet Meddelelse). I Oeders Tid havde der været Gartner ansat 
ved Haven, hvilket næsten var ganske nødvendigt, dels fordi Haven 
i sig selv var af ret betydeligt Omfang (over 11,000 [|] Al1.), dels 
fordi der fandtes flere Væksthuse. Til Gartner ved Haven i Univer- 
sitetets Eje udnævntes i 1770 J. W. Kæsemacher, der var kom- 
men hertil fra Amsterdam. Hans Løn var 200 Rd. + 60 Rd. til Hus- 
leje og Brændsel. .Skønt kun den mindre Del af den Oederske Have 
(se S. 339) var bleven skænket til Universitetet, fik den botaniske Have 
dog Lov til delvis at benytte den større Del (Kommerciekollegiets 
Del) af Amalienborghaven, navnlig havde Haven god Nytte af en 
stor Bygning, som fandtes paa dette Areal. Om Vinteren opbevare- 
des heri en Del Planter, og den afgav Bolig .for Gartneren. Da nu 
Kommerciekollegiet i 1777 fik Brug for sin Grund, maatte Botanisk 
Have fortrække herfra, men fik dog Brugsretten forlænget til For- 
aaret 1778. Professor Rottbøll lod derfor udarbejde Planer til en 
Bygning, som skulde være baade Drivhus, Gartnerbolig, Bibliothek 
og Auditorium og anmodede om at maatte faa dette Kompleks opført 
påa et Jordstykke mellem Frederiks Hospital og Haven. Dette kom 
dog ikke til Udførelse, thi Kongen besluttede at tilbagekøbe Haven 
af Universitetet for 6000 Ød. og skænke Grund til en ny. botanisk 
Have ved Charlottenborg, og i Løbet af 1778 rømmedes Amalien- 
borghaven, og Inventarium og Planter overflyttedes til Charlotten- 
borghaven. 





Fig. 125. Plan over Botanisk Have ved Charlottenborg. 1787. 


KØBENHAVNS 
UNIVERSITETS TREDIE BOTANISKE HAVE 


(CHARLOTTENBORG-HAVEN) 


ne Kommission, bestaaende af Professor Rottbøll, Gehejme- 
raad Holmskjold samt Konferensraad Aagaard nedsattes 
i 1777 for at gøre Forslag angaaende Indretningen af en botanisk 
Have i Slotshaven ved Charlottenborg, og Kongen approberede under 
22. Juli 1778 de Planer, som Kommissionen havde fremlagt ved- 
rørende denne ny Have. Under Arbejderne med Indretningen skulde 
Holmskjold have Tilsyn med Bygningernes Opførelse, C. F. Hars- 
dorff skulde tilse, at de leverede Tegninger fulgtes, medens Rottbøll 
skulde svare til, at Haven blev anlagt og indrettet paa rette Maade. 
I Aaret 1787 blev en med talrige Bilag forsynet Beretning indgivet til 
Kongen af Holmskjold og Rottbøll. Det fremgaar heraf, at Havens 
Anlæggelse og Indretning havde kostet godt 21,425 ØRdl., medens Aars- 
Budgettet paa dette Tidspunkt var 1363 Rdl. og nu blev forhøjet med 
400 Rdl. (Se Fig. 125, der viser den Plan, som var vedlagt Beretningen). 

Havens Styrelse var fra nu af lagt i Hænderne paa to Direktører, 
en valgt af Universitetet og en valgt af Kongen; de første Direktører 
var Rottbøll og Holmskjold. Senere gaves Direktionen videre Ram- 
mer; i 1798 bestod den af Statsminister C. F. Reventlow og Pro- 
fessorerne Wøldike, Vahl og Viborg. 

Kæsemacher var død i 1780, og Niels S. Bache var bleven 
Gartner i hans Sted, medens Martin Vahl i 1779 var bleven Lektor. 
I Aarene omkring 1787 fandt nogle indre Stridigheder Sted, drejende 
sig navnlig om Lektorens Ret til at benytte Havens Materiale. Det 


BOTANISKE HAVER 305 


var en af Vahls Venner, Kammerraad Tønder Lund, og Havens 
Gartner — denne vel nærmest som Straamand for Professor Rottbøll 
— der i flere Piecer, som udsendtes til Offentligheden, havde Havens 
Anliggender under Debat. Vahl søgte sin Afsked, men fik senere Op- 
rejsning ved at blive udnævnt til Professor. Men inden dette skete, 
rar Bache bleven forflyttet til Frederiksborg som Slotsgartner, og i 
hans Sted var Holbøll bleven udnævnt til botanisk Gartner. 

Frederik Ludvig Holbøll fødtes 14. Oktober 1765 paa Fry- 
denlund, hvor hans Fader var Gartner og Slotsforvalter. Efter udstaaet 
Læretid var han Svend hos sin 
Fader, som han senere fulgte i 
Embedet. I 1793 blev Holbøll ud- 
nævnt til botanisk Gartner, men 
tiltraadte dog først Pladsen 1794, 
da han forud for Tiltrædelsen 
foretog en Studierejse til Holland 
og England. Holbøll kom til at 
virke under fire forskellige Pro- 
fessorer, nemlig Rottbøll (d. 1797), 
Erik Viborg, M. Vahl (d. 1804) og 
Jens Wilken Hornemann. Det 
lange Samarbejde og inderlige 
Venskab mellem de to Mænd, 
Holbøll og Hornemann, og deres 
store Dygtighed indenfor deres 
Fag, har uden Tvivl haft sin store 
Betydning for Havens Udvikling, 
saa at man kan sige, at de har sat 
deres Præg paa den botaniske 
Have for lange Tider. 

Det Areal, som Haven omfat- 
tede ved Anlæggelsen, var c. 3. Tdr. Land, et Areal, som snart viste 
sig at være for lille, men til en Udvidelse paa Stedet var der den 
Gang ingen Udsigt. Et Tilbud, som Gehejmeraad Classen i 1803 
havde gjort Universitetet om at overlade dette 8 Tdr. Land Jord paa 
Østerbro, blev ikke modtaget, formentlig paa Grund af de forholdsvis 
store Udgifter, Nybygningerne vilde kræve. Man maatte saa se at 
skaffe Plads indenfor Havens egne Grænser og ved (1805) at rydde 
en Del overflødige Træer og Buske samt nogle Buxbomhække og ved 
at indskrænke Gangarealet lykkedes det for en Tid at skaffe nogen- 
lunde Plads. I 1803 havde Kongen skænket 3000 Rdl. til Opførelsen 
af et Væksthus, det havde en Længde af 71, Fod, en Bredde af 14 Fod 
og en Højde af 10”/, Fod. Atter i 1811 betænkte han Haven med en 
stor Gave, idet han til Erstatning for et mindre Areal, som skulde af- 
gives til Møntbygningen, skænkede en op til Haven stødende Grund, 

20 





Fig. 126. Frederik Ludvig Holbøll. 


306 DAN MAR KS HAVE BRU GRO GER GARTINSE PI 


hvorved Haven fik en Udvidelse paa 2000 Kv. Alen, endvidere skæn- 
kede Kongen 5000 RØRd!. til dette Areals Indhegning og Bearbejdelse. 
Herfor skulde bl.a. indrettes en ny Dam for Ferskvandsplanter, en 
Kanal med rindende Vand for saadanne Planter, der vokser ved 
Aaløb, endvidere en Plantning af Pile og et Anlæg for Alpeplanter. 

I 1817 skete den Forandring i Havens Forfatning, at den gik helt 
over til Universitetet, baade pekuniært og administrativt, og efter den 
Tid har Enkeltmand ståaet i Spidsen for Haven som Direktør. Hor- 
nemann, som havde siddet i Direktionen, blev som Professor i Bo- 
tanik ved Universitetet Havens 
første Ene-Direktør. 

Nu fremstod c. 1823 atter For- 
slag om at flytte Haven, hvilke 
gik ud paa at forene den botani- 
ske Have med Rosenborg Have, 
men disse Planer gled dog ud i 
Sandet, og Haven maatte nøjes 
med sit indskrænkede Åreal indtil 
1844. Holbøll var død i Aaret 1829 
og bleve ulk OF oRJolsikas 
Nite oa eN fø mee 09 
Aalborg, d.”/, 1842 i København), 
medens Joachim Frederik 
Schouw efter Hornemanns Død 
i 1841 blev Havens Direktør. 

Fra 1833 havde den Bestem- 
melse staaet ved Magt, at der til 
Arbejdskraften ved Botanisk Have 
foruden de 300 ØRdl., hvormed før- 
ste Svend lønnedes, maanedlig 
udbetaltes 125 Rdl. til den botani- 
ske Gartner, som derfor var pligtig at holde Haven i forsvarlig Stand. 
Gartneren havde ogsaa drevet en Del Handel med Planter og Frø fra 
Haven, hvortil der i hans Instruks var hjemlet ham Ret indenfor visse 
Grænser, Der blev nu ved Professor Schouws Initiativ indført den 
Regel, at al Handel af Planter, Frø o.l. fra Haven blev forbudt, lige- 
som den omtalte Maade at lønne Arbejdskraften paa blev afskaffet. 
I Stedet ansattes endnu to Svende, medens Resten af Bevillingen blev 
anvendt til Lønning af fast og løs Arbejdskraft. 

Budgettet, som i 1843 var 6312 Rdl. om Aaret, heri indbefattet 
Gartnerens og Åssistentens Løn samt Bygningsudgifter, forhøjedes 
1844 til 7200 Rdl og 1855—56 til 8200 Rdl. 

Husene undergik i denne Periode en Del Forbedringer. Et gammelt 
kapsk Hus maatte i 1837 nedrives; i dets Sted opførtes Aaret efter for 
3090 Rdl. et nyt Hus, bestaaende af 3 Afdelinger, hvoraf den midterste 





Fig. 127. August Weilbach. 


BOTANISKE HAVER 307 


skulde opvarmes ved varmt Vand, de to andre derimod ved de sæd- 
vanlige Røgkanaler. I 1843 blev er bevilget 9000 Rdl. til Opførelse af 
et Palmehus, 26 Alen langt, 12 Alen højt, 11'/, Alen bredt, og et Orchi- 
déhus paa 12 Vinduer i to Afdelinger, 26 Alen langt, 57/, Alen bredt 
og 4"/, Alen højt. Og endelig blev i 1846 et ældre Hus delvis ombygget. 
Mørch var død i 1842, og i hans Sted udnævntes fra 5. Oktober s.A. 
August Weilbach (f. ”/,, 1813 i København, d. sammesteds ”/, 
1868) til Botanisk Gartner. Weilbach, der havde faaet sin Uddannelse 
i Fredensborg Slotshave og i Botanisk Have, tog 1835 Kunstgartner- 
eksamen, var en Tid Medhjælper i Botanisk Have og rejste derpaa 
i Udlandet, hvorfra han blev kaldt hjem ved Mørchs Død. Han var 
en meget nidkær og duelig Embedsmand, som med Iver tog sig af Ha- 
ven og interesserede sig meget for dens Historie og dens Udvikling. 
I 1844 fik Haven en Udvidelse paa 6 Skæpper Land, idet en Have, 
som havde hørt under Søetaten, blev overladt til Universitetet. Der 
forlangtes først en ret stor Sum i Afstaaelse, hvilket foranledigede 
Havens Direktør til at udtale, at man, hvis dette Krav opretholdtes, 
hellere maatte søge Haven flyttet eller subsidiært anlægge en Filial- 
have udenfor Byen. Kongen resolverede derefter, at Jordstykket 
skulde afstaas, mod at der opførtes et Plankeværk som Grænse og 
foretoges nogle Reparationer paa Bolværket mod Mastegraven samt 
udbetaltes Holmens Overekvipagemester en aarlig Sum af 200 Øbd. 
som Afsavn for Haven. 

Professor Schouw trak sig 1852 tilbage fra sin Stilling som Direk- 
tør, og Professor Liebmann blev hans Efterfølger. Efter hans Død 
i 1856 antoges daværende Bibliothekar Joh. Lange til Direktør for 
Haven. 

Professor Schouw havde i 1847 udgivet en ,,Foreløbig Fortegnelse 
over Havens Planter, hvilket Værk var ledsaget af en Plan over hele 
Anlæget. Da Haven efter den Tid, indtil den i Begyndelsen af 70erne 
skulde flytte bort fra Gammelholm, ikke gennemgik væsentlige For- 
andringer, giver dette Kort (se Fig.128) god Oplysning om Havens 
Udseende i dette Tidsrum. Den laa lunt inde mellem Bygninger, vær- 
net mod skarpe Vinde fra Nord og Nordøst, hvilket sikkert har været 
til Gavn for mange Planters heldige Trivsel, men paa den anden Side 
har andre Planter, som netop lykkes bedst under mere aabne For- 
hold, kun kunnet mistrives dér. Husene laa dels direkte i Forbin- 
delse med Beboelseshusene, dels frit, nogle af dem laa med Facaden 
mod Syd, andre vendte mod Sydvest. De urteagtige Planter var sam- 
lede i regelrette Kvarterer, enaarige, toaarige og fleraarige hver for 
sig, ligeledes var Lægeplanter og indenlandske Planter plantede i re- 
gelmæssige Kvarterer. Den øvrige Del af Haven var udlagt i Plæner 
og Busketter, den største Del af Arboretet laa i Havens sydlige Del. 
Alpeplanter dyrkedes paa Stenhøje paa tre forskellige Steder i Haven 
og Vandplanter i et regelmæssigt Bassin, hvilket dog led af den 


20= 


Kort over den botaniske Have i Kjøbenhavn 1847. 


XAN ÅL ————= hede Jordbælte. (Palmer: Cy- 
SN AN cadeer. Bambuser. osv) … 
ners Bolig. EG: 23. Varmt Huzes. Femtalsplan. 

3æg kolde Huse. Plan: 5. Portnerens Bolig. ter (Dicotyledsner) fra hes 

ter fra det varme. 6. Læsesal. Ovenover Plantes de Jordbælte. 
Pe ererte og Frée=Samlinger. Z4. Koldt Huus. Planter fra 
Jordbæltef Mid 7 IEEE SY TS3 det. varmere ererte Jord 
delhavslan . især fra Meazco og fydafrica. bælte,rsær fra Sydafrica og 

denelNord. (Czeteer. Ålver. osv.). Ny holland. i 
2 Z ”/ FEDER 7 75. Hures til at tiltrække kjel 
Ze sæ FE ERE nere Plaziter, tsær Jra 72 
E africa ogNyholland. 

9 Varmt Huus. Yngre Plan. I eg 
fer fra det hede-Jordbælte. Ze: Cs Bede til forskjel. 

10. Varmt Huus. Orchideer : 


77. Steder, hvor de Planter 
fra det hede Jordbætlte. 2653 FEED SR 


£ Koldt Huzs. Sydafri- : , 
GE SAG SE BR holdes i de kolde Hwse - 
raa L3. Steenplanter. 


= = Er …… 79. Planter til rærmere. Un. 





TRSS 
SÅ 











<a å Ben Træer og Buske. 


== E — QN mn Sækken: 














Græsstykker med 


rige Urter. 





































































































Charlottenborg N 





Fig. 129. Fra den botaniske Have ved Charlottenborg. 














Fig. 130. Parti fra Botanisk Have ved Charlottenborg c. 1830. 


310 DANMARKS HAVEBRUG OG GARTNERI 


Ulempe, at dets Vand var for varmt, da Spildevand fra Møntens 
Dampmaskine ledtes ud deri. 

Forholdene var i det hele meget småa og indsnævrede, og Havens 
Embedsmænd saa stadig med Længsel hen til en mulig Flytning (Se 
Fig. 129 og 130). Weilbach var i denne Retning ikke den mindst ivrige, 
og han fremkom i 1856 i Dagbladet ,,Fædrelandet" med en Serie 
Artikler (under Mærket 1—24), i hvilke han behandlede flere af Ha- 
vens Forhold. Han gør her opmærksom paa den skadelige Indvirkning, 
som Røgen, der med visse Vinde føres ind over Haven, udøver paa 
Planterne; han klager over den slette Jordbund, som for en stor Del 
bestaar af Murbrokker, medens andre Dele af Haven er altfor mul- 
dede paa Grund af den tidligere intensive Dyrkning med Køkken- 
urter, og han beretter, at visse Dele af Haven ved Højvande over- 
svømmes med Saltvand. Dertil kommer, at Grundvandet staar saa 
højt, at det allerede træffes i en Alens Dybde, og om Vinteren løber 
Fyrgravene derfor fulde af Vand. Forholdene var saaledes ikke såa 
idylliske, som man maaske ved et kortere Besøg i Aarets skønne Dage 
vilde faa Indtryk af. Weilbachs Artikler i Forbindelse med de Planer, 
som var fremme om Bebyggelse af Gammelholm, gav en frodig Jord- 
bund for Tanken om Flytning; om disse Planers videre Udvikling be- 
rettes i Stykket om Haven paa Glaciet, den nuværende botaniske Have. 

Efter at Planerne for den ny Have var bragt ud i Livet og ført til 
en lykkelig Afslutning — Weilbach skulde ikke opleve at se dette ske 
— blev Haven efterhaanden tømt for sin Bestand, medens Arealet 
blev overladt til Kunstakademiet og til Bebyggelse, og i April 1877 
afleveredes det sidste Stykke. 

I en lille Stump Have tæt op til Charlottenborg Udstillingsbygnin- 
gen staar endnu enkelte Træer: Celtis australis, Fraxinus angustifolia 
og Pyrus prunifolia tilbage som Minde om Universitetets tredie bo- 


taniske Have. 








Fig. 131. Fra Botan. Haves Drivhuse. Koldhus Nr. i med Fr. V's Buste. 


KØBENHAVNS 
UNIVERSITETS FJERDE BOTANISKE. HAVE 


(DEN NUVÆRENDE HAVE) 


E to Forsøg, der havde været gjorte paa at faa Botanisk Have 

flyttet (1803 og c. 1820, se f.0.) havde ikke ført til noget Re- 
sultat; men nu var Tanken atter med Kraft bleven offentlig fremført 
af Weilbach. Spørgsmaalet havde ogsaa været omtalt i Rigsdagen, og 
Ministeriet henvendte sig derefter til Universitetet for at erfare dettes 
Stilling til en mulig Flytning. 

Foranlediget herved nedsatte Konsistorium d. 4. November 1857 
en Komité, bestaaende af Havens Direktør Joh. Lange, Professor 
Jap. Steenstrup og Professor J. G. Forchhammer, for at fore- 
tage forskellige Forundersøgelser angaaende en eventuel Flytning. 
Denne Komité afgav i Juni 1859 en Erklæring, gaaende ud paa, at 
den maatte anse Flytningen for nyttig, ja næsten nødvendig. At an- 
vende en Del af Rosenborg Slotshave til en ny botanisk Have, hvilket 
Ministeriet havde peget paa, maatte Komiteen, selv om denne Haves 
Beliggenhed var bekvem, fraraade, dels fordi Jordbunden her efter 
Udsagn af en Komité af Gartnere, som i sin Tid havde undersøgt den, 
fandtes uskikket til finere Planters Dyrkning, dels fordi man fandt 
det uheldigt at berøve Byens Beboere en saa yndet Spaseregang, 
hvilket man nødvendigen maatte gøre, hvis man lagde en botanisk 
Have der, da denne i al Fald til visse Tider maatte være afspærret 
for Publikum. > 

Komiteen havde haft sin Opmærksomhed henvendt paa Grunde paa 
Frederiksberg, men den fandt det uheldigt at lægge Haven saa langt 
fjernet fra Universitetet; den burde lægges i en saadan Afstand, at 


312 DANMARKS HAVEBRUG OG GARTNERI 


der ikke spildtes for megen Tid for Universitetslærerne og de Stu- 
derende, der skulde benytte Haven. Efter forskellige Overvejelser var 
Komiteen bleven staaende ved den Plan, at foreslaa Haven lagt paa 
Glaciet umiddelbart udenfor de Fæstningsværker, der var bleven ind- 
rømmede til Observatoriet. Komiteen ønskede at erhverve c. 12 Tdr. 
Land til dette ny Anlæg, og den havde tænkt sig, at Skraaningerne 
af den Del af Volden, som laa mellem Sølvgade og Gothersgade, 
skulde anvendes til Beplantning med Træer af den Art, som man 
plejede at optage i en botanisk Haves Arboret, men med særligt Hen- 
syn til Skønheden og Publikums aldeles fri Benyttelse som Spasere- 
gang. Dette Arboret burde da forenes med den øvrige botaniske 
Have paa Glaciet ved Hjælp af en Fodbro over Stadsgraven, og denne 
egentlige botaniske Have burde til visse Tider være aaben for Publi- 
kums frie Besøg. 

For at forberede den botaniske Haves Flytning fandt Komiteen 
det hensigtsmæssigt, at der anlagdes en Planteskole for at man kunde 
tiltrække en Del Træer og Buske til Anvendelse i den ny Have, og i 
Overensstemmelse hermed lejedes af de militære Myndigheder et Åreal 
paa c. 1 Td. Land tæt udenfor Nørreport (omtrent paa Grønttorvets 
Plads). 

Man havde imidlertid endnu ikke faaet Vished for, at den ny 
botaniske Have vilde komme til at ligge paa det af Komiteen i 1859 
foreslaaede Sted, thi Militærautoriteterne raadede endnu over hele 
Fæstningsterrænet, og det var endnu ikke afgjort, i hvilken Udstræk- 
ning Københavns Befæstning skulde nedlægges, og hvorledes det mu- 
lig frigivne Areal vilde blive benyttet. Forhandlinger med en Kom- 
mission, som det var overdraget at udarbejde Bebyggelsesplaner for 
Terrænet ned til Søerne, førte dog til, at denne Kommission paa sin 
i 1865 udgivne ,,Plan til Bebyggelse af Terrainet mellem Kjøben- 
havns Demarcationslinie og den indre By, efter Fæstningsvoldenes 
Sløjfning” havde indtegnet en botanisk Have paa den af Have-Komi- 
teen angivne Plads, og i Loven om Demarcationslinien af 6te Juli 
1867 & 4 bestemtes det, at et Areal af indtil 21 Tdr. Lands Størrelse 
kunde afstaas til Anlæg af en botanisk Have og et astronomisk Ob- 
servatorium. 

Imidlertid var Botanisk Gartner Aug. Weilbach bleven op- 
taget i Komiteen, og i 1866 forøgedes denne yderligere med Professor 
bot. A. S. Ørsted, Professor Gram og Kvæstor Frydensberg 
med Bygningsinspektør Hansen som Medhjælp. 

Der paafulgte nu en lang Række Forhandlinger mellem Komiteen 
og Konsistorium, Kultusministeriet, Krigsministeriet og Finansmini- 
steriet samt Københavns Kommune om en Række Spørgsmaal, 
drejende sig om Universitetets Ejendomsret til den gamle botaniske 
Haves Grund, Muligheden for et helt eller delvist Mageskifte mellem 
den gamle og den ny Haves Areal, den ny Haves Størrelse, Erhver- 


BOTANISKE HAVER 2313 


velsen af nogle Kommunen tilhørende Arealer ved Farimagsvejen 
o.m.a., og som Følge af disse Forhandlinger trak Havesagen i Lang- 
drag. Komiteen undergik i denne Tid nogle smaa Forandringer, idet 
Gartner Weilbach i 1868 døde og erstattedes med Th. Friedrich- 
sen, medens Kvæstor Frydensberg erstattedes med  Kvæstor 
Gedde, og i 1871 indtraadte Kaptajn, Brygger J. C. Jacobsen i 
Komiteen. Og nu var man kommet saa vidt frem i Arbejdet, at der 


i ne eg ig Commune Hospitalet (fyn 
Aj E id 





Farimagsve i 














Gad'e 







SØRESN 
misk 2 V 


Astronomisk < g 
Observatortum SES 











Voldgade LEN, 








me MELET RR oe NR SR 





Fig. 132, Forslag til Botanisk Have. 1870. 


paa Finansloven for 1871—72 kunde bevilges til Anlæg af en ny bo- 
tanisk Have paa Glaciet et foreløbigt Beløb paa 25,000 Rdl. 

Komiteen havde allerede i 1870 udarbejdet en ,,Beretning om Uni- 
versitetets paatænkte nye Botaniske Have". Denne Beretning var led- 
saget af to Planer, en for Musæet og en for selve Haven med de der- 
paa liggende Bygninger. Beretningen var særlig beregnet til Uddeling 
blandt Rigsdagens Medlemmer for at gøre disse nærmere bekendt med, 
hvilke Kaar Haven forhen havde arbejdet under, og hvorledes man 
i store Træk havde tænkt sig den ny Have. 

Som det vil ses af Planen over Haven (Fig. 132) var det Tanken 
at lægge alle Væksthusene i en fortløbende Række langs Farimags- 


314 DANMARKS HAVEBRUG OG GARTNERI 


- 


vejen. Endvidere fremgik det af Beretningen, at det var Komiteens 
Tanke at omgive hele Haven med en Mur og at planere Arealet med 
Undtagelse af en midt i Haven beliggende Bastion, som skulde an- 
vendes til Stentøj og til en Del af Arboretet. Ogsaa Dele af Vold- 
skrænten, som skulde bruges til Arboret, skulde heller ikke planeres. 
Man agtede ogsaa til Dels at gennemføre en plantegeografisk Ordning 
og endelig vilde man ved Havens Anlæg stræbe at fyldestgøre Skøn- 
hedens Fordringer saavel i dens enkelte Dele som i de mange og for- 
skelligartede Deles Forening til et harmonisk Hele. 

Denne Beretning frembragte i Tidsskrift for Havevæsen en livlig 
Diskussion. Tidsskriftets Redaktør J. A. Dybdahl, Docent F. Did- 
richsen og Kammerjunker Rømer (som skrev under Pseudony- 
met: C€. Rasmussen, Havemand) angreb Komiteens Planer, og Be- 
retningen forsvaredes i særlige Piecer af daværende Cand. mag. Eug. 
Warming og Professor ÅA. S. Ørsted. 

Docent Dybdahls Artikel var meget saglig og indeholdt mange 
berettigede Anker mod Forslaget. Hans Indlæg gik i første Linie ud 
påa at foreslaa, at Haven ikke blev lagt paa Glaciet, men nærved 
Landbohøjskolens Have paa Pladsen mellem Sygehjemmet, Biilows- 
vej og Ladegaardsaaen (5 Td. Land), idet han mente, at man ved at 
vælge Pladsen her vilde kunne bygge Væksthuse og Musæum, men 
nøjes med smaa Frilandsanlæg og navnlig helt undvære Arboretet, 
da man i den nære Landbohøjskoles Have havde et saadant og til- 
lige en smuk Stenhøj saavel som Samlinger af Lægeplanter og agro- 
nomiske Planter. Han kritiserede den foreslaaede Væksthusrække (til 
hvilke Weilbachs Tegninger og Modeller var lagt til Grund) og i ikke 
mindre Grad selve Have-Anlæget, og han sluttede med at udtale: 
» Hvis imidlertid det hele af Komiteen til botanisk Have nu udsete 
Terrain skulde blive indtaget dertil, foreslaae vi, at man sikkrer sig 
en landskabsgartnerisk smuk Benyttelse deraf ved at henvende sig 
til en eller anden i denne Retning anerkjendt dygtig Anlægs- 
gartner.” 

I et Bilag til ,,Beretningenf" læste man under det arktisk-alpine 
Rige: ,,Dette plantegeografiske Rige har en særlig Interesse derved, 
at de fleste af de danske Bilande, Grønland, Island og Færøerne, høre 
hertil, hvorfor det ogsaa i den nye Have vil blive gjengivet paa en 
tiltalende og anskuelig Maade. Dette kan nemlig uden Vanskelighed 
ske ved at give ,,Stenhøienf, den med Sten besatte Høi, paa hvilken 
man netop pleier fortrinsvis at dyrke de arktisk-alpine Planter, en 
til dette Øjemed svarende Tillempning. Til denne Brug vil en om- 
trent midt paa den til Haven bestemte Del af Glaciet liggende Bastion 
fortrinlig egne sig. Den har en Høide af omtrent 25 Fod, og en Del 
af Skraaningen vil blive anvendt til at gjengive de Plantebælter, som 
ligge over Skovgrænsen paa Alperne. Paa de nederste 5 Fod plantes 
Dværgfyrren for at betegne Skovgrænsen, der ligger paa 5,500 Fod, 


BOTANISKE. HAVER 915 


derpaa følger et Bælte (den alpine Region), af lave Buske, blandt 
hvilke Alperoserne og Ellen ... ere herskende, som naar op til 7000 
Fod og indtager de næste 5 Fod af Høiens Skraaning. Mellem 7000 
Fod og 8500 Fod ligger det øverste Plantebælte (den subnivale Re- 
gion) beklædende omtrent 10 Fod af Høiens Skraaning. Det er her, 
at talrige Saxifrager, Ranunkler, Drabaer, Primulaer, Entianer, Si- 
lener o.m. fl. strække sig op til den Bjergets Top bedækkende evige 
Sne.” 

Dette gav ,,Havemand Rasmussen" Anledning til i sm Kritik at 
ytre: ,,Der mangler blot for at fuldstændiggjøre Legetøiet, at den 
evige Sne blev fremstillet med Meelkalk og Kridtstene, at der paa 
Steenhøien blev anbragt en Miniatur-Sennhytte og nogle forlorne 
Geder, og at der ved Foden af Højen imod Statsgraven blev opslaaet 
et Brædt med: her er Middelhavet og dets Flora, samt at lade dette 
blive repræsenteret af nogle Potter med Laurbær-, Orange-, Oliven- 
og Myrtetræer, indfattede af Lavendler og Salvier.” 

Heldigvis blev der intet Hensyn taget til Forslaget om at flytte 
Haven ud paa Frederiksberg, hvilket maaske nok vilde have været 
til Gavn for Landbohøjskolen, men sandsynligvis til liden Baade for 
Universitetet. Angrebene gjorde dog deres Gavn, thi de har maaske 
bevirket, at de to Kapaciteter H. Flindt og Tyge Rothe blev 
tagne med paa Raad. 

Ved Behandlingen i Rigsdagen (Samlingen 1870—71) af Forsla- 
get om at bevilge de føromtalte 25,000 Rd. til foreløbige Arbejder op- 
stod en langvarig Diskussion om den botaniske Have; det blev fra et 
Mindretals Side foreslaaet at udstemme Bevillingen, for at man kunde 
faa Tid til at tage under Overvejelse, om Haven ikke kunde lægges 
et andet Sted, ligesom der udtaltes Ønsker om at faa mere detaille- 
rede Planer. Minister Hall udtalte under Behandlingen i Tinget 
Muligheden af, at der vilde blive udskrevet en offentlig Konkurrence. 

Ministeriet tilskrev kort derefter Komiteen og henstillede til den, 
at den tilkaldte landskabsgartnerisk kyndig Hjælp, og i Overens- 
stemmelse hermed foreslog Komiteen, at Landskabsgartner H. 
Flindt og Slotsgartner Tyge Rothe blev tilkaldte som Raadgivere 
mod passende Honorar, hvilket Ministeriet tiltraadte. 

Landskabsgartner, senere Haveinspektør, Flindt udarbejdede 
Planerne angaaende Havens Anlæg paa Grundlag af de Krav, som 
Komiteen maatte stille med Hensyn til den botaniske Ordning, og 
det lykkedes Flindt uden at forstyrre det hele Anlægs systematiske 
Ordning at forbinde den af Komiteens Flertal antydede Gruppering 
med den af Professor Ørsted med saa megen Varme anbefalede Plan 
til et nordamerikansk Arboret paa en særdeles tilfredsstillende Maa- 
de. Tyge Rothe havde i Forbindelse med Brygger Jacobsen Plan- 
læggelsen af og Tilsynet med Opførelsen af Væksthusene under sig. 

Havens Anlæggelse og Husenes Opførelse saavel som Bygningen 


316 DANMARKS HAVEBRUG OG GARTNERI 


af Tjenesteboligerne stod paa i Åarene 1871—74. Bekostningen ved 
Havens Anlæg og Husenes samt Tjenesteboligernes Opførelse androg 
i alt c. 642,000 Kr., hertil nogle private Tilskud fra Brygger Jacob- 
sen. Første Aar udførtes navnlig Entreprenørarbejderne med Afgrav- 
ning og Planering af Volde og Opfyldning af Dele af Stadsgraven; 
disse Planeringsarbejder var meget betydelige, efter en foretagen Be- 
regning flyttedes ialt 168,000 Kubikmeter Jord. Plantningen af Ar- 
boretet fandt i det væsentlige Sted i 1872—73. 

Den 1. Juni 1874 kunde Komiteen aflevere Haven til dennes Di- 





ae rer me; 1288072 åg ar SE sm KEE ERE SST FN 
ER == sån" zØ tat ETTE FIFE EJER | EIN: (237 
BR 1 På + att | ber i msg; KE: n:: 1 88 + AR s KE part så 


REE) 





Era: (æres z 


Fig. 133. Vue over Botanisk Have c. 1885. 


rektion og dermed sluttede Komiteen sin Virksomhed. Samme Åar 
den 9. Oktober aabnedes Haven for Publikum. 

I Regler for Adgang til Haven, som samtidig udstedtes, bestemtes 
det, at Haven skulde være aaben for Publikum daglig fra Kl. 1 til 
Solnedgang, at der til Personer, som ønskede Adgang udenfor disse 
Tider, kunde udstedes Adgangskort, lydende paa Navn, mod en Be- 
taling af 8 Kroner (Familiekort 16 Kroner), medens Studerende kunde 
faa gratis Adgang til Kvarteret for Lægeplanter fra 8 Morgen til Sol- 
nedgang, at Palmehuset var aabent hver Dag fra 3—6 Em.”) og de 
lavere Huse aabne 3- Gange om Ugen samt at Barnevogne og Hunde 
ikke maatte medtages i Haven. 


=£) Senere er dette ændret til 2—6, senere igen til 1—5. 


BOTANISKE HAVER o17 


København havde faaet sig en ny botanisk Have og var derved 
bleven et værdifuldt Parkanlæg rigere. Den af Flindt udarbejdede 
Plan viste sig ogsaa i sin færdige Skikkelse at være et smukt Anlæg, 
som Aar for Aar blev skønnere og skønnere, saa at man med en vis 
Ret kan sige, at kun faa botaniske Haver i Verden i landskabelig 
Henseende staar over vor Have. (Fig. 133.) 

Haven har Form som en uregelmæssig langstrakt Firkant, c. 
380m lang og 270m bred, mod Øst skærer Observatoriets Grund sig 
som en Oval ind i Havens Areal. Haven, der ved sit Anlæg maalte 
9), hekt. (17%/, Tdå. Land), bestaar af et større, Dalparti, hvori den 


å HE 





F i 
X ø 
LZ me; 


Fig. 134. Stenhøjen i Botanisk Have 1918. 


9,000 [] m store Sø — den omdannede Stadsgrav — befinder sig. 
Vandspejlet er 1'/,m over Havets Overflade, Søens Dybde er omkring 
3,20 m., frisk Vand kan tilføres ad en underjordisk Ledning fra Sorte- 
damssøen. Fra Dalpartiet hæver Terrænet sig til den ene Side (Øst 
og Sydøst) temmelig stejlt opimod Boulevarden og Observatoriebak- 
ken, dennes højeste Punkt er 13m; til de andre Sider stiger Terræ- 
net mindre brat og breder sig snart ud i en stor jævn Flade, som 
indtager den største Del af Haven. Paa dette Plateau er Funktionær- 
boligerne, Musæet og senere Botanisk Laboratorium byggede; Vækst- 
husene ligger i Havens nordre Del, dels paa Plateauet, dels i Dal- 
sænkningen. 

. Ved Beplantningen af Arboretet blev der fulgt en systematisk 
Plan, hvorefter hver enkelt Families Træer blev plantede i Grupper 


318 DANMARKS HAVEBRUG OG GARTNERI 


for sig; men ikke alle Familier indenfor hver Orden blev lagt sam- 
lede, saaledes anbragtes. Æblefamilien langt fjærnet fra Stenfrugtfa- 
milien, de skaalbærende langt fra Pilefamilien; dette blev formentlig 
gjort for at fordele de Familier, som havde mere fremtrædende Blom- 
ster, over hele Haven. Ligeledes blev alle de perennerende urteagtige 
Planter familievis plantede i Grupper eller Rabatter over hele Haven. 
Den nærmere Ordning af Træer og Urtegrupper kan ses af Planen 
(Fig. 135). I Havens vestlige Hjørne blev anlagt en Samling af nord- 
amerikanske Træer, plantede uden Hensyn til deres Plads i Systemet. 

Til særlig Brug for de Studerende anlagdes langs Gothersgade i 
regelmæssige Bede et Studiekvarter — benævnet ,,Medicinsk Kvarter" 
— hvori overvejende plantedes Planter af økonomisk eller medicinsk 
Betydning. Omtrent midt i Haven hæver sig Stenhøjen (Se Fig. 134), 
anlagt paa en Dobbelthøj, omformet af en Bastion af det gamle Fæst- 
ningsværk. De højeste Punkter her er 13—14m, og herfra har man 
et smukt Vue over hele Haven. Ved Søens nordre Ende er anlagt et 
Moseparti og hertil slutter sig Kvarteret for de indenlandske urte- 
agtige Planter: ,,Dansk Kvarter.” 

Væksthusene dækker ialt en Grundflade af 2400m”, de har et 
Rumindhold af 12,000m' og en dækkende Glasflade af 3100m?”; 
højeste Punkt af Palmehuset er 19m; de store Huse og visse Dele af 
de smaa har dobbelt Glaslag. Husene er af stor ornamental Virkning, 
som de ligger i Baggrunden af Haven, og omend de ligger ret højt, 
er de dog ved høje Bygninger i Farimagsgade værnede mod Vinde 
fra Nord og Nordvest. Ved Anlæggelsen af Haven plantedes bag 
Væksthusene langs Farimagsgade en Del Popler, hvormed Hensigten 
var dels at skabe yderligere Læ, dels at fremskaffe en smuk Baggrund 
for Anlæget set fra den søndre Del af Haven og samtidig skjule for 
Blikket de ovennævnte Beboelseshuse. Den øvre Væksthusrække be- 
staar af 5 Afdelinger, nemlig to Ende-Rotunder for Koldhusplanter, 
en stor Midter-Rotunde for tropiske Enkimbladede, ,,Palmehuset" og 
to Saddeltagshuse, som forbinder Ende-Rotunderne med Palmehuset. 
I Palmehuset bæres et højere Midterparti af 18 Støbejærnssøjler; ad 
en Vindeltrappe kommer man op paa et omløbende Galleri, hvorfra 
man fra oven kan betragte Planterne i Husets Midterparti. Hele dette 
Anlæg af store Huse har en Længde af 94m; Bagsiden af Husene 
optages af Værktøjsskure, Folkestue og Arbejdsrum. Foran Vækst- 
husrækken løber en bred cementeret Perron, begrænset af en Balu- 
strade; midt foran Palmehuset fører herfra en bred Stentrappe ned 
i Haven. Denne Stentrappe deler den lavere Væksthusrække i to lige 
store Dele. Under Perronen findes Varmeanlæget, Kulkælder, Maskin- 
rum og Oplagsrum. Foran den lave Væksthusrække, bestaaende af 
7 Afdelinger, alle ensidige, ligger to fritliggende Saddeltagshuse. Midt 
imellem disse ligger et Springvandsbassin, der ligesom Vaserne paa 
Balustraden er skænkede af Brygger Jacobsen. Øst for Væksthus- 
komplekset ligger det 12-kantede Akvarium, 9/,m i Diameter og 5m 


BOTANISKE HAVER 319 


højt. Desuden findes bag Husene en Del Bænke og Bakker, et For- 
søgshus og et Formeringshus. 

Varmeanlæget til Husene er baseret paa Opvarmning ved Damp. 
Der findes 3 store Dampkedler; Dampen herfra gaar over i et fælles 
Hovedrør og fordeles derfra til alle Husene. Til Opvarmning af Hu- 
sene medgaar aarlig mellem 300 og 400 Tons Kul. 

Det var fra først af Tanken, at Musæet skulde bygges samtidig 
med at Væksthusene byggedes, men forskellige Forhold gjorde, at 
Bevillingen til Musæets Opførelse blev udsat. Endelig i Aaret 1875 
opførtes denne Bygning, hertil var bevilget 112,000 Kroner, heri ind- 
befattet Montering. 

Johan Lange, der havde været konstitueret Direktør for Ha- 
ven fra 1855, blev i 1875 efterfulgt af F. Didrichsen, der virkede 
som Professor og Direktør ved Haven til 1885, og siden den Tid har 
Direktørstillingen været knyttet til et Professorat i Botanik ved Uni- 
versitetet. Didrichsen har næppe sat noget Præg paa Haven i snæv- 
rere Forstand, den var saa ny, at der ingen Grund var til at foretage 
Forandringer, og ny Opgaver for Haven fremsatte Didrichsen ikke. 

Haven havde under Forarbejderne for Flytningen skiftet Gartner, 
i Weilbachs Sted var Th Friedrichsen (f. i København ”/, 1827, 
død paa Frederiksberg ””/, 1907) antaget. Han havde siddet i Have- 
Komiteen og han havde, efter at Anlæget var afleveret til Haven, 
foretaget de Plantninger, som stod tilbage at fuldføre og endelig 
havde han ledet den store Flytning af Planterne fra den gamle 
til den ny Have. Han kunde derfor siges og følte sig ogsaa at være 
ét med denne ny Have, og han holdt gennem hele sin lange Virksom- 
hedsperiode med rørende Pietet og Trofasthed fast ved det af Ko- 
miteen for Havens Drift fastsatte og ændrede kun deri, naar han af 
Omstændighederne blev tvungen dertil. 

Men Udviklingen førte ret naturligt med sig, at der i Tidernes 
Løb dog maatte foretages Omlægninger og Smaa-Ændringer ved Ha- 
ven. Ny Tider giver ny Krav, Embedsmænds Skiften vil ofte sætte 
sine Spor; ny Naboer vil kaste lystne Øjne paa Havens Omraade, som 
passende Basis for Udvidelse, og Træernes Vækst vil ganske natur- 
ligt fremtvinge Ændringer: her og der. En af de første vigtigste Æn- 
dringer i Haven var Opførelsen af ,,Planteanatomisk Laboratorium". 
Professor Eug. Warming (f. :/,, 1841 paa Manø) var fulgt efter 
Didrichsen i Embedet som Professor i Botanik og Direktør for Ha- 
ven, han kom hertil fra Sverrig, hvor han havde virket i 3 Aar som 
Professor ved Stockholms Hågskola; han kom ladet med Energi og 
Virkelyst til sin ny Stilling, og just i en Tid, da der var Kræfter i 
Gang for at faa et Botanisk Laboratorium. Allerede i 1880 havde deri 
interesserede Botanikere indgivet et Andragende til Kultusministeriet 
om Oprettelsen af et selvstændigt plantefysiologisk Laboratorium. 
Kultusministeren fremsatte derpaa Forslag om Opførelse af en saa- 
dan Bygning, men Sagen nød ikke Fremme i Rigsdagen. Endelig ved 


—— mt] 
i 


D 











i 
DUM 
3 





Hem 


(t. 
i 


Solvga 


n dqan 


de 
z cc 
AM 





SEP BSD BST OASE EEG 
rd > 


ei ON SR. AEV EK vrd ME per uuelnn £Y ss É 
annen rig NR KR ESS na FEED. SE 
SMRT SN, 20 NT rs AF BASEL I] g >: ' 
ANNA REN g w NADVER DEKAN HELSE = 
g 73 ad ) 






oo 


z ØS 


vi 























100 Alen 


Fig. 135. Plan af Botanisk Have 1907. 
Omlægning af Rosenplænen var projekteret som vist paa Tegningen (29), 
men kom ikke til Udførelse 


so 


























Fa) Set 
SØS Nes & ' oa L 
f SIGE DS . Ca F"2 
rn SENE ed Ses = å 
| >» m 


BP: DEDESEDEDS URE DIT DST SØ 


1. Enaarige Planter. 24. Hamamelidaceæ. 46. Anacardiaceæ og 

2. Planteskole. 25. Kornelfamilien. Rutaceæ. 

3. Forsøgsareal. 26. Hydrangea. 47. Olietræfamilien. 

4. Ubestemte Planter. 27. Ærteblomstrede. 48. Ribsfamilien. 

5. Stenbede. 28. Æblefamilien. 49. Stenhøje. 

6. Sølvbladfamilien. 29. Rosarium. 50. Cypresfamilien. 

7. Pilefamilien. 30. Sommerbede for 51. Alperosefamilien. 

8. Magnetisk Observa- Koldhusplanter. 52. Rosenfamilien. 
torium. 31. Sommerbede for 53. Berberisfamilien. 

9. Skolekvarter. Alpeplanter. 54. Lindefamilien. 

10. Danske Planter. 32. Varmebedsanlæg. 55. Vrietornfamilien, 

11. Dansk Stenhøj. 33. Espaliermure. Benvedfamilien, Sta- 

12. Mosebed. 34. Jordmagasin. phyleaceæ. 

13. Rubus (Brombær). 35. Ælmefamilien. 56. Pibevedfamilien 

14. Granfamilien. 36. Celtidaceæ. (Philadelphus). 

15. Taksfamilien. 37. Skaalfrugtfamilien. 57. Paulownia. 

16. Taxodium. 38. Lønfamilien (ell. 58. Ibenholtfamilien. 

17. Ginkgo. Ahornfamilien). 59. Kristtornfamilien. 

18. Slyngplanter (Løv- 39. Valnødfamilien. 60. Hestekastaniefa- 
gang). 40. Magnoliefamilien. milien. 

19. Birkefamilien. 41. Actinidia. 61. Buksbomfamilien. 

20. Gedebladfamilien. 42. Platanfamilien. 62. Prydgrupper. 

21. Hasselfamilien. 43. Morbærfamilien. 63. Biologiske Grupper. 

22. Mandelfamilien. 44. Gøgeurtgruppe. 64. Kjælnere Buske. 

23. Vedbendfamilien. 45. Bignoniaceæ. 


BOTANISKE HAVER 


321 


Forklaring til Tallene paa Plan af Botanisk Have 1905. 


Lov af ”/, 1888 blev der, samtidig med at der blev sikret Midler til 
andre Instituter under Universitetet, bevilget c. 200,000 Kr. til Opret- 
telse af et Planteanatomisk Laboratorium. Laboratoriet opførtes i 
Aarene 1888—90 i Havens vestre Del paa den Plads, hvor det nord- 
amerikanske Arboret laa, og Haven i snævrere Forstand mistede der- 
ved et ret betydeligt Areal (omkring "”/, hektare). Laboratoriet toges 
i Brug i 1890. Kort efter udkastede Professor Warming Plan til at 
faa omdannet en Del af ,,Medicinsk Kvarter" til et ,,Biologisk Kvar- 
ter", Omtrent Halvdelen af det medicinske Kvarter blev omdannet 
til det ny Formaal; endvidere anlagdes i samme Hensigt en Del Grup- 
per i de tilgrænsende Plæner. I det følgende Ti-Aar foregik ikke større 
Ændringer indenfor Haven: men i Åaret 1902 fandt en Personæn- 
dring Sted, idet Gartner Friedrichsen paa Grund af fremrykket Al- 
der søgte sin Afsked. I hans Sted ansattes fra "/,, 1902 Axel Lange 
(f. paa Frederiksberg «|, 1871) som Botanisk Gartner. 

Polyteknisk Læreanstalt, som i Slutningen af 80erne var bleven 
bygget tæt op til Botanisk Have og var taget i Brug 1890, blev, som 
Aarene gik, snævrere og snævrere for dens stærkt voksende Virk- 
somhed. Der fremkom derfor c. 1903 et Ønske om, at Haven vilde 
afstaa et Areal langs Farimagsgade, saaledes at Læreanstalten kunde 
faa den længe tiltrængte Udvidelse. Til Erstatning derfor skulde Ha- 
ven paa et vist Aaremaal (50 Aar) have en Kommunen tilhørende 
Grund, som stødte op til Haven. Haven mente ikke — særlig af Hen- 
syn til den Betydning, det polytekniske Studium har for hele Landet 
at kunne modsætte sig den fremstillede Plan. I Overensstemmelse 


21 





322 DANMARKS HAVEBRUG OG GARTNERI 


hermed afstodes saa i 1904 og 1905 i alt c. ”/, hektare Jord, derved 
mistede Haven sin Forsøgshave og Dele af Mistbænkanlæget. En ny 
Forsøgshave anlagdes paa det af Kommunen lejede Areal. 

Haven led i landskabelig Henseende ved Ændringen, idet den 
smukke Baggrund, den hidtil i Nordsiden havde haft i en Række 
Popler langs Farimagsgade, blev erstattet med en kold og kedelig 
Bygning. Haven forpligtede sig overfor Kommunen til som et Slags 


Vederlag for det paa meget billige Vilkaar lejede 0,7 hektare store 








Fig. 136. Botanisk Haves Drivhuse 1906. 


Areal at lade en Laage anbringe i Stakittet ud mod Sølvgade og at 
lade Færdsel gennem Haven ad Veje Øst for Søen finde Sted hele 


Dagen. Erhvervelsen af dette kommunale Anlæg — det saakaldte 
Sølvgadeanlæg — gav paa den anden Side Haven Lejlighed til inden- 


for sine Rammer at foretage nogle meget ønskelige Ændringer. Træ- 
grupperne havde efterhaanden bredt sig saa stærkt, at de mange Ste- 
der overskyggede Staudegrupperne. Ny Voksekaar for de perenne- 
rende Planter var derfor særlig ønskelige; Pladsen, hvor de enaarige 
og toaarige Planter hidtil havde staaet, blev nu indtaget til Staude- 
kvarter, de toaarige Planter fordeltes mellem Stauderne, medens de 
enaarige anbragtes i koncentriske Bede, anlagte i Plæner i det lejede 
Jordareal. Den 9. Oktober 1905 aabnedes Sølvgadelaagen for Pub- 
likum. 





BOTANISKE HAVER 5724) 


Efter at Professor Warming med Udgangen af 1911 havde søgt 
sin Afsked, blev Professor C. Raunkiær (f. ”/, 1860 paa Raunkiær- 
gaard, Jylland) Direktør for Haven. 

Af Ændringer indenfor Haven, foretagne i Tidernes Løb, kan for- 
uden de ovenomtalte nævnes nogle ret betydelige Omlægninger af 
Stenhøjene, Anlæg af Stenhøj paa Foden af Observatoriebakken, og 
i 1918 Omlægning af Biologisk Kvarter i regelrette Bede, hvorved 
vandtes saa megen Plads, at ,,Medicinsk Kvarter" eller som det nu 
kaldes ,,Undervisningskvarteret" kunde faa noget af det i 1892 mi- 
stede Areal tilbage. 

Havens nuværende Bestand af Embedsmænd og Funktionærer er 
som følger: Hele Institutionen staar under Havens Direktør, dog sty- 
res det plantefysiologiske Laboratorium uafhængigt heraf af Profes- 
soren i Plantefysiologi (Professor W. Johannsen, f. ”., 1857 i Kø- 
benhavn) med Hjælp af en Assistent. Ved det planteanatomiske La- 
boratorium er ansat en Assistent, og ved disse to Laboratorier findes 
en fælles Portner og Laboratoriebetjent. Musæet ledes af Musæums- 
inspektøren med Hjælp af en Amanuensis, en Konservatrice og en 
Musæumsbetjent; Bibliotheket styres af . Bibliothekaren. Plantebe- 
stemmelsesarbejdet i Haven besørges af en Haveamanuensis. Haven 
ledes af den botaniske Gartner med Hjælp af (Jan. 1919) 2 Under- 
gartnere, 3 Medhjælpere, 2 Fyrbødere, 4 Opsynsmænd, samt c. 20 
mandlige og kvindelige Personer, som delvis er gartnerisk uddan- 
nede; det aarlige normerede Arbejdsdageantal er 7,000. 

Til Bygningernes Vedligeholdelse er der beregnet 9,000 Kr., som 
er opført paa den samlede Sum til Universitetets Bygningers Ved- 
ligeholdelse. Hvert dte eller 6te Aar maa der med særlig Bevilling 
finde en Hovedreparation af Husene Sted, ved hvilken Lejlighed al 
Jærnværk og Træværk bliver efterset; alle løse Vinduesrammer ned- 
tages (Fig. 136 viser Palmehuset under Reparation 1906; Vindues- 
rammerne i venstre Side er nedtagne), efterses med Kitning og Maling. 


Der er i de tidligere Afsnit talt om fire forskellige Haver, som 
har været knyttede til Universitetet, men mellem disse Haver gaar 
der mange Baand, som gør, at man med fuld Ret kan tale om Ha- 


verne som en enkelt Have, thi ved hver Flytning er Planter — baade 
Frilands- og Væksthusplanter — fra den gamle Have bleven over- 


ført til den ny, Havens Embedsmænd har stedse været med at lede 
Flytningen og Forbindelsen med Universitetet har stadig været til 
Stede, selv om den ikke al Tid har været saa fast og direkte som 
nu. Og- medens der i det foregaaende hovedsagentlig er talt om de 
topografiske, administrative, økonomiske og personelle Forhold ved 
hver enkelt af de fire Haver, skal der i det følgende prøves givet en 
Fremstilling af, hvilken Betydning Haven har haft som Plantesam- 


21 


324 DANMARKS HAVEBRUG OG GARTNERI 


lings- og Undervisningsanstalt, hvorledes Plantetallet er tiltaget og 
Plantematerialet hår vekslet gennem Tiderne, samt hvilke Opgaver 
Haven i det hele har søgt at løse. 

I sine første Dage har Haven uden Tvivl næsten udelukkende 
været en Have for de medicinske Studerende, og den har vel tillige 
tjent til Glæde og Nytte for den Professor, som beboede ,,Resident- 
zen", der hørte til Haven. Der dyrkedes særlig Frilandsplanter og, 
i al Fald fra først af, mest danske Planter. Ole W orm skaffede flere 
udenlandske Planter til Haven, saaledes: Rheum rhaponticum, Sco- 

















Fig. 137. Paulownia imperialis i Botanisk Have. 


rozonera hispanica, Solidago canadensis og Mimosa pudica m. fl. I 
den før omtalte Bog af J. Buchwald findes opført 200 Planter. Ved 
hvert Navn i Bogen er en tørret Plante, svarende dertil, indklæbet. 
Det er dog ikke givet, ja næppe sandsynligt, at dette var hele Plante- 
bestanden i Haven, endmindre maa man antage, at alle deri nævnte 
Planter dyrkedes i Haven; det giver næppe noget rigtigt Billede af, 
hvad Haven indeholdt. Sit Herbarie-Materiale har Buchwald sikkert 
dels samlet i Haven, dels i Naturen, f. Eks. Lathræa squamaria, dels 
maaske hos Apothekerne, f. Eks. Viscum album. Af udenlandske 
Planter, som findes nævnt eller opklæbet i Bogen, skal som Eksemp- 
ler nævnes: Adiantum capillus veneris, Antirrhinum majus, Canna- 
bis sativa, Cnicus benedictus, Cucumis sativus, Cyclamen europæum, 
Juniperus sabina, Lilium candidum, Physalis alkekengi og Veratrum 


BOTANISKE HAVER sa) 


album. løvrigt er Værket meget upaalideligt; i de forskellige Eksem- 
plarer, der endnu forefindes, ser man tit ved samme Plantenavn for- 
skellige Plantearter opklæbede. 

Om Haven ved Amalienborg opgives, at Antallet af dyrkede Arter 
i 1761 var 650, medens det 1763 var naaet til 1500 Arter. Efter Flyt- 
ningen til Charlottenborg kom der mere Vækst i Plantesamlingernes 
Årtsantal. En skreven Katalog fra 1784, forfattet af Bache, udviser, 
at der dyrkedes 2855 Arter i Haven. 1801 er Tallet c. 5000, i 1813 op- 
giver Hornemann, som dette Aar udgiver en trykt Fortegnelse, An- 
tallet til 7,500, i en Fortegnelse udgivet 1839 af Mørch findes c. 7,700, 
1848 er det steget til 9,000 og 1857 til 9.500, medens der ved en kal- 
kulatorisk Opgørelse i 1917 fandtes c. 12,000 Arter fordelte paa c- 
2,570 Slægter. 

Forholdet mellem Slægter og Arter er al Tid Genstand for Sving- 
ninger; Tendensen bør, som allerede af Schouw i 1847 hævdet. gaa i 
Retning af at søge at faa saa mange Slægter som muligt repræsente- 
ret, og ved Reduktion af Plantebestanden med den Hensigt at skaffe 
Plads til ny Planter, vil man derfor som Regel særlig lade den gaa 
ud over artrige Siægter. Imidlertid har det sin Interesse af og til at 
samle talrige Arter indenfor enkelte Slægter, og ser man tilbage paa 
Plantefortegnelserne for de svundne Aar, vil man kunne gøre mange 
interessante Iagttagelser og danne sig Billeder af, hvad der har in- 
teresseret de styrende indenfor Haven, eller hvad der har været de 
vekslende Tiders Yndlingsplanter. 1803 andrager Holbøll Havens Di- 
rektion om Tilladelse til at købe 25 Arter Erica for 4 £ og samme Aar 
opgiver han Tallet af Mesembrianthemum til 40 Arter; i 1813 dyrke- 
des 42 Arter Erica. Det er de kapske Planter, som er dominerende! 
og endnu i Mørchs Tid ser vi, at disse Planter har Fortrinnet. I Mørchs 
Fortegnelse er opført: 75 Mesembrianthemum, 35 Acacia, 23 Erica og 
over 50 Pelargonium, deraf dog sikkert mange Havesorter. Til Sam- 
menligning kan anføres, at der samtidig dyrkes af Primula 17, af 
Sazifraga 27, af Agave 4 og af Begonia 10 Arter. Tager man de før- 
nævnte Slægter og undersøger, hvad der i 1918 fandtes i Haven, vil 
man for nogles Vedkommende finde en Nedgang, for andres en be- 
tydelig Opgang; Resultatet er: Mesembrianthemum 20, Erica 13, Pe- 
largonium 25, Primula 64, Saxifraga 100, Agave 50, Begonia 65. Næv- 
nes kan ogsaa, at Botanisk Have indtil 1834 kun besad 3 Arter tro- 
piske Orchideer. Nævnte Aar fik den nogle Arter fra Hamburg og i 
1839—41 en større Samling (60 Arter) fra Hofraad Hambroe, samt 
1711 Arter fra Mexiko ved Liebmann. Nu ejer Haven c. 180 Arter 
i Hus. 

Sjældenheder og store smukke Eksemplarer af visse Arter har 
Haven stadig kunnet glæde sig ved at besidde og som nævnt før, har 
enkelte Individer fulgt med fra den ene Have til den anden. Af Plan- 
ter paa Friland er et Eksemplar af Paulownia imperialis flyttet til 


a chinensis. 


Liviston 


anisk Have. 


2 
>0 


[sml 


i 


t 


38. Fra Palmehuse 


> 
(31 





BOTANISKE HAVER PAT 


den nuværende Have i 1871 (se Fig.137); denne Plante eksisterer 
endnu. Af Væksthusplanter, som er Bindeled mellem den gamle og 
ny Have, kan nævnes den store Midterplante i Palmehuset Phoenix 
spinosa, som er kommet til Haven i 1807, og en ualmindelig kraftig 
Livistona chinensis, €. 75 Aar gammel (se Fig.138), som, da den 
truede med at vokse igennem Glasset, nødvendigvis maatte falde for 
Øksen i 1919. Dens samlede Vægt var 615 kg og den bar, da den fældedes, 
15 kg Frugter og 50 Blade. En Sukkerpalme, Årenga sacchari- 
fera, der ligeledes var overført fra Charlottenborghaven, blomstrede 
i Tiden Januar 1904—Eftersommeren 1906 (se Fig. 139) og døde ef- 
ter afsluttet Blomstring, som det 
er denne Plantes Natur; den var 
dengang c. 75 Aar gammel. 

Eksemplarer af Agave ameri- 
cana har blomstret gentagne Gan- 
ge og et var i sin Tid endog of- 
fentlig udstillet. Dette var i 1836, 
i botanisk Gartner Mørchs Tid. 
Han fortæller i en lille Piece, 
skrevet i denne Anledning, at 
Eksemplaret stammede fra Skaa- 
ne, hvor han selv havde været 
ovre at hente det. Blomsterstilken 
var allerede fuldkomment udvik- 
let og Sidegrenene dannede. Efter 
at Planten med største Omhu var = —— 
indpakket, førtes den med 8 Fig. 139. Fra Bot. Haves Palmehus. 
Heste til Malmø, hvorfra den pr. SUS) DANES AD GLIEE 
Skib fragtedes til København, 
hvor den ankom 15. September, uden at en Gren var knækket eller 
Væksten hæmmet. — 

En Sjældenhed paa Friland er det lille orientalske Træ af Æble- 
familien Eriolobus trilobata, som staar paa Stenhøjen (se Fig. 140). 

Haven har gennem Aarene forøget og vedligeholdt sin Plantebe- 
stand, dels ved Køb hos Handelsgartnere, dels ved Gaver af Planter 
eller Frø fra Botanikere, udsendt til fremmede Lande, og dels ved 
Bytte med beslægtede Institutioner i Udlandet. Havens begrænsede 
Budget har gærne lagt Hindringer i Vejen for at den første af de her 
nævnte Veje kunde benyttes i større Udstrækning, og ligeledes er der 
kun i ringe Grad kommet Planter til Haven ved at der her fra Lan- 
det er sendt Botanikere paa Rejser til fjærne Lande, i Sammenligning 
med, hvad store botaniske Haver i Europa har kunnet. Her maa dog 
nævnes, at Gartner Rathsack, som rejste i Mexiko med Botanikeren 
Liebmann, bragte Haven talrige Planter fra sin Rejse. 

Den vigtigste Kilde til Rekruttering af Havens Plantebestand bliver 





328 DANMARKS HAVEBRUG OG GARTNERI 


dog Udveksling af Frø og Planter med fremmede Haver. Det om Som- 
meren høstede Frø renses om Efteraaret og Vinteren, og i Januar ud- 
sendes en Katalog over, hvad Haven kan undvære. Denne Katalog 
udsendes til udenlandske botaniske Haver samt til en Række Skole- 
haver og Institutioner her i Landet. Efter de Ønskelister, som hjem- 
kommer, udsendes 
Frøet i Marts Maa- 
ned. 

Til Gengæld mod- 
tager Haven fra frem- 
mede Haver Katalog 
over det dér høstede 
Frø, hvoraf vor Have 
udvælger, hvad den 
maatte have Brug for. 
Som Regel forlanges 
hvert Aar 2000 å 3000 
Portioner, Haven selv 
udsender omkring 
11,000 Portioner aar- 
lig (Gennemsnit af de 
sidste 5 Aar). Af en 
Uddelingsliste, der 
gaar tilbage til 1841, 
aftrykkes her Antallet 
af uddelte Portioner 
for Aar med 20 Aars 
Mellemrum; 1841 ud- 
deltes 2373 Portioner, 
1861: 3084; 1881: 5733, 
ag AA z. 1901: 5557; 1918 var 
Fig. 140. Stenhøjen i Botanisk Have Tallet 12,250. Den 

med Eriolobus trilobata. stærke Forøgelse fal- 

der særlig påa de in- 
denlandske Institutioner. Udvekslingen af levende Planter mellem 
fremmede botaniske Haver og vor Institution andrager aarlig nogle 
faa Hundrede Numre. 

Det er sikkert en almindelig Opfattelse, at man altid kan stole 
paa de Navne, som findes ved Planterne i en botanisk Have. Des- 
værre er det imidlertid ikke altid Tilfældet. Paa Grund af forskellige 
Fejlkilder, hvorpaa her ikke nærmere skal gaas ind, kan Planterne 
ikke være helt igennem korrekt etiketterede, og da dette er et Forhold, 
alle botaniske Haver er underkastet, er man stedse i Uvished, om de 
Planter, man modtager fra Udlandet, er rigtig bestemte. 

Om Haverne er mere paalidelige nu end i gamle Dage, er vanske- 








ir 


BOTANISKE HAVER 329 


ligt at afgøre uden grundig Statistik, men saa langt tilbage som 1813 
klages der over fremmede Havers Unøjagtighed. Vor Haves Embeds- 
mænd har stadig søgt at holde Planterne rigtig bestemte, i tidligere 
Tider var det særlig Havens Direktør, som besørgede dette Arbejde. 
Professorerne Schouw, Liebmann og Lange har revideret talrige Plan- 
ter, men med Havens stigende Udvikling blev det nødvendigt at an- 
sætte en særlig Plantebestemmer — Haveamanuensis — hvis Opgave 
er stadig at revidere Havens Samlinger af levende Planter. 

Der føres i Haven nøjagtig Bog over, hvorfra hver enkelt Plante 
er kommet, samt naar den er revideret o.s.v.; dette System, indført 
af Holbøll i 1795, er der med Flid vaaget over siden, og man er der- 
ved sat i Stand til at kunne give enhver Plantes Historie her i 
Haven. 

Den botaniske Have holder stadig sin første oprindelige Bestem- 
melse at være en Studiehave højt i Ære. Derfor er den største Del af 
den om Formiddagen afspærret for Publikum, men tilgængelig for 
Studerende og Lærere med Elever, der udstedes uden Vederlag 
Studiekort og Lærerkort til alle dertil berettigede, som andrager 
derom. 

I ældre Tider afgav Medicinerne det største Kontingent af dem, 
som studerede Botanik i Haven, men efter at denne Videnskab fra 
Aaret 1903 er ophørt med at være et tvungent Fag for nævnte Studie- 
gren, er der kun enkelte Medicinere, som con amore søger Haven 
som Studiested, og dog benyttes Studiekvartererne sikkert i stigende 
Grad. Antallet af farmaceutiske Studerende er taget til, Statens Lærer- 
kursus saavelsom Faglærerne o. m. åa. benytter vor Have. Mate- 
riale til Professorernes Undervisning og Eksaminer skal ogsaa af- 
gives fra Haven; desuden afskæres Materiale til Skolelærere og til 
Manuduktører o.s.v.; ialt afgives til Undervisning udenfor Universi- 
tetet og Farmaceutisk Læreanstalt omkring 1000 Portioner aarligt. 

Foruden at være en Studieanstalt skal den botaniske Have tillige 
tjene til Oplysning og Glæde for Publikum og omend de fleste af de 
ca. 1,412,000 Personer, som aarlig passerer Laagerne, blot skyder Gen- 
vej gennem Haven, er der dog en stor Mængde, der nyder de For- 
dele, den yder som behagelig Spaserepark og Frilandsmusæum. Sty- 
relsen har søgt paa forskellig Maade at gøre Haven tiltrækkende. Det 
ovenfor nævnte af Professor Warming udtænkte biologiske Kvarter 
skal ikke alene tjene som Undervisningsmiddel for de Studerende, 
men tillige hos de ikke faglærte vække Interessen for og aabne Øj- 
nene for Planternes Ligheder og Forskelligheder. Planterne i dette 
Kvarter er ordnede efter saadanne Egenskaber og Karakterer, som 
har Betydning for Planternes Liv i Naturen eller som betinger deres 
Værdi som Kulturplanter. Saaledes er i en Gruppe sammenstillet 
Marehalm, Strandkaal og Echeveria secunda, alle med blaa- 
duggede Blade, i en anden Salvia argentea, Gnaphalium petiolatum 


330 DANMARKS HAVEBRUG OG GARTNERI 


og Lammeøre, alle med tæthaarede Blade. Disse Gruppers Planter 
skal illustrere, hvorledes Naturen giver Planterne Værn mod stærkt 
udtørrende Klima. Kapske Vortemælk og amerikanske Kaktus stilles 
sammen for at vise Stængelsukulenter og for at illustrere, hvorledes 
systematisk vidt adskilte Planter kan have ens Vækstformer. 
Husenes Tilgængelighed for Publikum skal ogsaa bidrage til at 
brede Interessen for Planternes Liv og de forskellige Floraomraader. 
Af og til arrangeres særlige Udstillinger af enkelte Plantegrupper. I 











Fig. 141. Udstilling af Araceer og Commelinaceer i 1918. 


1898 udstilledes saaledes, hvad Haven besad af Stenbræk (Sawxifraga); 
i 1915 udstilledes en Samling af den tropiske Slægt Begonia, 1917 bød 
Haven paa en Samling Hængeplanter og endelig var i 1918 udstillet 
en Samling af Araceer og Commelinaceer med henholdsvis c. 125 og 
c. 25 Repræsentanter (Se Fig. 141). 

Botanikere, der har været knyttet til Haven, ofte ogsaa Havens 
Gartnere, har dér, enten i Forsøgskvarteret eller i Husene, foretaget 
videnskabelige eller praktisk-økonomiske Forsøg. Disse Forsøg gaar 
langt tilbage i Tiden. Det ses saaledes af Have-Tidende 1841, at Bota- 
nisk Gartner Mørch samme Aar havde haft Forsøg med Silkeormes 
Fodring med Blade af forskellige Planter, beslægtede med Morbær, 
hvorved han kom til det Resultat, at den amerikanske Elm var en 


BOTANISKE HAVER Bel 


god Foderplante for disse Dyr. Botanisk Gartner Weilbach plan- 
lagde Forsøg med Anvendelse af tykt Glas i Væksthuse og med Luft- 
bevægelse i Væksthuse, som Middel til at yde Planterne naturligere 
Voksekaar; Problemet har en af vore driftige Handelsgartnere for en 
8—9 Aar siden søgt at løse i sine egne Drivhuse. Botanikerne S am- 
søe Lund og Kiærskou har gjort Forsøg med Kaalsorter og be- 
slægtede Planter, Docent Becker anstillede i 1894—99 Forsøg over 
Perleløgets Oprindelse og Magister O. Rostrup foretog i Tiden 1886 
—89 for Dansk Frøkontrol Forsøg med Frøs Spireevne. I de senere 
Aar har Professor W. Johannsen haft Forsøg med rene Linier i 
Bønner, medens Professor Raunkiær og Dr. Ostenfeld har sys- 
let med Artsforholdene hos forskellige af vore Blomsterplanter og 
med Kimdannelse uden Befrugtning. 

Selv om det er Haven i snævrere Forstand, som her omhandles, 
bør der dog tilføjes et Par Ord om det til Institutionen hørende Mu- 
sæum og Bibliothek. Musæet, ledet af Dr. phil. O. Paulsen, omfatter 
dels Samlinger i Spiritus, dels Samlinger af Vedprøver og Frøsorter 
samt andre tørre Plantedele og som det mest omfattende: de paa 
Papir opklæbede tørrede Planter: Herbariet. Dette sidste er delt i 3 
Hovedafdelinger, nemlig det arktiske Herbarium, som anses for det 
bedste i sin Slags, det danske Herbarium og Generalherbariet, hvilket 
rummer Planter, der ikke kan henføres til nogen af de to andre Af- 
delinger. Herbarierne omfatter ca. 475,000 Ark. Musæet er ikke offent- 
lig tilgængeligt, men forevises paa Forlangende. Bibliotheket — ledet 
af Dr. phil. F. Børgesen — indeholdende særdeles mange værdi- 
fulde Bøger, er offentlig tilgængeligt 3 Gange om Ugen. Det rummer 
omkring 20,000 Bind. 

Den botaniske Have har nu eksisteret i 319 Aar og den har i den 
Tid ført en ret omtumlet Tilværelse, idet den, som anført, har ligget 
paa fire forskellige Steder i Byen. Krav fra forskellig Side kan let 
føre med sig, at Haven atter en Gang helt eller delvis maa skifte 
Plads, men selv om Forholdene allerede nu i visse Henseender er ind- 
skrænkede, og selv om dens Beliggenhed midt i Byens Hjærte kan 
have uheldig Indflydelse paa mange Planters Trivsel, maa man dog 
ønske, at den saa længe som muligt maa kunne bevares uindskrænket 
paa sin nuværende Plads, saa at den kan opnaa det Præg af Ærvær- 
dighed, som klæder Haver saa smukt og som sikkert ikke mindst vil 
klæde denne i sig selv saa landskabeligt skønne Park. 


SORØ ARBORET 


I Slutningen af forrige Aarhundrede blev der mellem Bestyrelsen 
for Sorø Akademis Skove og Botanisk Haves Direktion og med Mini- 
steriets Samtykke truffet Aftale om, at Botanisk Have i Sorø Skove 


352 DANMARKS HAVEBRUG OG GARTNERI 


paa dertil egnede Pladser, som Skovvæsenet kunde afse, skulde ind- 
plante forskellige fremmede Træarter. Det var Tanken derigennem 
at forskønne Sorø Lystskov, men ikke at skabe noget botanisk syste- 
matisk Arboret. I Tiden fra 1903 er der nær Filosofgangen indplantet 
en Del Naaletræer, især Graner og Ædelgraner, samt en stor Sam- 
ling Tjørn, medens der i Feldskov er plantet Acer i større Antal og 
talrige andre Træer i enkelte Eksemplarer, og endelig er der i Hunde- 
kildehave plantet en Samling Birke. 


KLELERSUNIVERSITETETS BOTANISKE 
HAVEPTIE 1864 


ED Universitetet i Kiel, indviet 1665 af Hertug Christian Al- 
brecht af Holsten-Gottorp, oprettedes i 1669 en botanisk Have, 
og Hertugen stillede en Del af Slotshaven til Disposition for dette For- 
maal. Det overdroges Johannes Daniel Major, Professor i theo- 
retisk Medicin, at anlægge og bestyre Haven. Major havde haabet at 
faa hele den fyrstelige Have Nord for Slottet overladt til nævnte 
Brug, men den daværende Slotsgartner Heinrich Vack modar- 
bejdede denne Plan og vilde have Major henvist til et lille kvadratisk 
Stykke Jord, Baumhof kaldet; men dette erklærede Major for uegnet 
"til Formaalet, og Hertugen befalede derpaa, at der skulde afstaas 
en Fjerdedel af Slotshaven, samt at der til Hjælp i Haven skulde 
gives Fjerdedelen af den Arbejdskraft, som ydedes af de Bønder, der 
gjorde Pligtarbejde for Slotshaven. 
Anlæggelsen af Haven stod paa i fire Aar og voldte Major megen 


Besvær — han taler et Sted om sit ,,blutsauren vierjahrigen untertå- 
nigst ehrlich gemeinten ersten Einrichtungsarbeit — bl. åa. ved at 


Slotsgartneren ikke fuldtud vilde levere ham den lovede Arbejds- 
kraft. Just som Major var færdig dermed, skete der nogle Forskyd- 
ninger indenfor Lærerkræfterne ved Univesitetet, hvorved Major blev 
Professor i praktisk Medicin, medens Johannes Nikolaus 
Pechlin blev Professor i theoretisk Medicin og Botanik og som saa- 
dan overtog Ledelsen af Haven. Af Afleveringsdokumenterne fra Ma- 
jor til Pechlin ses det, at der paa nævnte Tidspunkt mest dyrkedes 
medicinske Urter; nogle ikke haardføre Planter forefandtes dog 
ogsaa. 

Haven var (se Fig. 142) anlagt i regelmæssig Stil. Fra Gaden kom 
man ind ad Porten (c paa Planen), der foroven var forsynet med 3 
marmorerede Kugler, under hvilke fandtes Hertugens forgyldte Nav- 
netræk i en Laurbærkrans. Ved Indtrædelsen i Haven ad denne Port 
havde man et Vue over hele Haven. Ad en anden Port (ved e) kom 
man ad en Trappe ned til Ridebanen, som skilte Haven fra Slottet. 

Blandt de Planter, som dyrkedes i eller havde været dyrket i 
Haven indtil 1673, kan nævnes: Acorus aromaticus (Acorus cala- 
mus>), Blattaria fl. violaceo (Verbascum phoeniceum), Campanula 


+) Navnene i Paranthes er de nugældende Navne. 
te] 


334 DANMARKS HAVEBRUG OG GARTNERI 


pyramidalis, Cerefolium hispanicum, (Myrrhis odorata), Flos africa- 
nus (Tagetes), Hyucca gloriosa (Yucca gloriosa), Laurus tinus (Vibu- 
num tinus), Pomum amoris (Lycopersicum esculentum), Sophia 
chirurgorum (Sisymbrium sophia, Barberforstand). Ialt dyrke- 
des omkring 500 Arter og Havesorter. 

Endvidere fandtes opført i Plantelisten A/oe Gottorpiensis [in ef- 
figie] d.v.s. et Billede af en Agave Americana, som i 1668 havde 
blomstret i den hertuge- 
| 6 lige Have ved Gottorp og 
| hvorom Major havde 

i skrevet et lille Værk i 4". 
JENS Pechlrnsynes 

| æ ikke at have vist nogen 
særlig Interesse for Ha- 

ven; der foreligger intet 


PER om den fra de Aar, han 
iii beklædte — Direktoratet. 
NEN IN EEN ERE REE RES Det synes, som om Ha- 
ing kg su E DON ven ikke har eksisteret 


ud over Aaret 1684. 
I 1720erne blev der 
i 
| 


























(24 
(& 








anlagt en ny botanisk 

Have: Hortus medicus, 

nærved SsUniversitetsbyg- 

ningen. Den var en Tid 

lejettudtortbleværeiseN 

overladt til Apotheker 

e Christiani, som med 

Fig. 142, Den ældste botaniske et aarligt Vederlag af 10 

HALERRIGER Rthl. overtog Forpligtel- 

sen til at vedligeholde 

Anlæget som en medicinsk Have og levere Urter og Plantedele til 

Forelæsninger og Eksamener. Denne Akkord skulde overføres paa 
senere Indehavere af Raadsapotheket. 

I 1802 fik Universitetet et medicinsk Hospital, hvilket Professor 
Georg Heinrich Weber oprettede, navnlig ved frivillige Bidrag. 
Ved dette Hospital blev det følgende Aar anlagt en ny botanisk Have 
og derefter ophørte det aarlige Tilskud paa 10 Rthl. til Apotheker- 
haven, som dog vedblev at høre til Apotheket indtil 1855, da Jord- 
stykket gik over til Universitetet mod en Afstaaelsessum af 200 Rthl. 

Den ny botaniske Have, hvis Anlæggelse stod under Professor 
G. H. Webers og Overlæge Fischers Opsigt, blev i 1804 over- 
givet til Webers Søn Friedrich Webers (f. 1781, d. 1823) Le- 
delse, som styrede den i omtrent 20 Åar. 

Haven havde efter datidige Udsagn (1811) ,,rigelig Plads, forskel- 





BOTANISKE HAVER 335 


ligartet Terræn, gennemstrømmende Vand, godt indrettede Vækst- 
huse og en dygtig botanisk Gartner.” Sidstnævnte Mand var Peter 
Høhle, som var i Havens Tjeneste fra 1806 til 1831. I Webers Tid 
udvidedes Haven to Gange, saaledes at den i 1822 maalte ÆREN ds 
Land. Drivhusene bestod samme Aar af 7 Afdelinger, byggede dels 
ved Havens Anlæggelse, dels i 1816 og dels i 1820. Havens Plante- 
bestand i 1822 var ret betydelig, nemlig omkring 6000 Arter. Der dyr- 
kedes bl. a. 31 Astragalus, 11 Begonia, 55 Campanula, 108 Mesem- 
brianthemum, 85 Pelargonium, 20 Saxifraga; af Bromeliaceer dyrke- 
des 4 Arter og af Orchideer i Hus 6 Arter. 

Efter Fr. Webers Død var Ernst Ferdinand Nolte (f. 
1791, d. ”/, 1875) Havens Direktør, og omkring 1855 var C. Dahle 
dens Gartner. Haven var i Sommertiden tilgængelig for Studerende 
alle Hverdage fra 7—12 og fra 1—7 og for Publikum var Haven 
aaben fra Juni til September to Gange om Ugen fra 4—7 Em. 


ROSENBORGS 
KONGELIGESBOTANISKE SHAVE 


1606 købte Christian IV en Del Grunde udenfor Voldene i den 

Hensigt at bygge sig et Lystslot og anlægge sig en Have for dér 
at kunne have et Sted at rekreere sig. Her byggede Kongen i Tiden 
1610—1624 Slottet Rosenborg. I Haven var der til Tider ansat to 
Urtegaardsmænd (Gartnere) og flere Svende. Af en Instrux, udstedt 
24, Marts 1633, fremgaar det, at der dette Åar skulde dyrkes et Bed 
fuldt af Simplicia (Lægeplanter). Dette kan maaske regnes for en 
Forløber for Rosenborgs botaniske Have, hvis Skæbne man iøvrigt 
ikke ved meget om; den synes næsten udelukkende at have været 
knyttet til Grundlæggeren Dr. med. Otto Sperlings Virksomhed dér. 

I Aaret 1638 havde Corfitz Ulfeld, som nærede en betydelig In- 
teresse for Lægen og Botanikeren Otto Sperling, anbefalet denne 
til Kongen, ,,som en der var vel forfaren udi Urtevidenskaben, og 
hvem Hs. Majestæt skulde betro Inspektionen over sin Have.” Chri- 
stian IV fulgte Ulfelds Indstilling og udnævnte 7. Juni s. A. Otto 
Sperling til Læge ved Børnehuset i København og ,,derforuden have 
vi naadigst bestilt hannem at skal være vores Botanicus og have flit- 
tig Indseende med Haven ved vor Kiøbsted Kiøbenhavn, og tilholde 
Urtegaardsmanden, at han sig beflitter at have alle de Simplicia, som 
gror dog i vore Lande, hvorfore Vi hannem have aarligen tilsagt 
200 Rd. in Sp. ....” Det synes efter Ordlyden af Ovenstaaende at 
dømme, at Sperling har haft Overopsyn med hele Haven, men han 
har sikkert i særlig Grad taget sig af den botaniske Del. 

Otto Sperling havde set sig vel om i Verden og havde bl. a. 
været i Venedig, hvor han havde været ansat som Botanicus i Haven 
hos en Raadsherre Contarenus. Til ham skrev nu Sperling og for- 
talte om sin Udnævnelse samt bad, om han ikke vilde sende nogle 
sjældne Frø, endvidere skrev Sperling til en Guilielmo Boel i 
Lissabon og herfra fik han iblandt andet to Slags ,,af en sjælden in- 
disk Væxt, kaldet Herba viva, som er saaledes beskaffen, at naar man 
rører nok saa lidet ved den med en Stok eller Finger, bevæger den 
sig straks, lukker sine Blade med stor Hastighed sammen og sænker 
sine Grene mod Jorden. Da jeg (Sperling) nu engang talte med Stat- 


BOTANISKE HAVER 337 


holderen derom, fik han Lyst til at se samme Urt og beskuede den 
med Forundring. Han berettede det til Kongen, og den fromme 
Herre kom ogsaa og kunde ikke noksom undre sig over denne sæl- 
somme Bevægelse og over, hvorledes Væxten snart efter lukkede sine 
Blade op igjen og Grenene hævede sig fra Jorden.” Senere foretog Sper- 
ling en Rejse til Spanien og besøgte den kongelige Have ved Buen 
Retiro. Han købte der ,,en Del Rødder" af en Hyacinthus indicus, hvis 
Blomster lignede Narcisser og tog dem med sig til Danmark, men 
han kunde ikke faa dem til at blomstre dér. 

Sperling har efter Samtidiges Udtalelser været meget bjærgsom 
og dertil paaholdende med sine Planter, saa at han end ikke over- 
lod sine nære Venner noget uden klækkeligt Vederlag, Han udgav i 
Aaret 1642 sin Hortus Christianæus, hvori opførtes 1519 Navne paa 
Planter, som han dyrkede i Haven. Rottbøll siger herom: ,,Hvilke, 
dersom de end for de mange Afvexlingers Skyld formindskes til 1000 
dog ere for den Tid et meget betydeligt Tal, og kunde ikke uden stor 
Flittighed, Kundskab og en vis Paaholdenhed i saa kort en Tid brin- 
ges sammen, besynderlig da iblant samme findes mange rare uden- 
landske Planter.” Rosenborghavens Samling af Planter har været 
langt over Universitetshavens. Vi ser da ogsaa, at Simon Paulli i 
sin Viridaria varia 1653 optrykker Sperlings Liste, medens han ikke 
nævner Universitetshavens Samling. 

Her et lille Udvalg af Planter, som dyrkedes i Rosenborg Have i 
1642. De i Parenthes anførte Plantenavne er de nu anvendte. 

Alcea vesicaria (Hibiscus trionum?), Alsine baccifera (Cucubalus 
bacciferus), Alnus nigra seu Frangula (Rhamnus frangula), Arbor 
vitæ (Biota orientalis), Aster marinus Dodon. Tripoliu (Aster tripo- 
lium), Capsicum americanum siliqu. oblong. tenuissima og 14 andre 
Capsicum-Former (Capsicum annuum, Spansk Peber), Ceratonia 
Teophrasti s. siliqua (Ceratonia siliqua, Johannesbrødtræ), 
Chamelæa tricoccos (Cneorum tricoccum), Chelidonium majus offi- 
cinar og Ch. maj. alterum fol. et floribus laciniatus (Man har altsaa 
allerede den Gang haft den fligetbladede Form af Svaleurt under 
Kultur), Chelidonium minus (Ficaria verna), Christophoriana (Actæa 
spicata), Colchicum autumnale tessellatum, Fritillarium  dictum 
(Colchicum variegatum), Convolvulus non convolvulus Hispanicus 
tricolor (Convolvulus tricolor), Cucumis asinius (Ecballium agreste, 
Æselsagurk), Halicacabum officin. Alkekengi (Physalis alkekengi, 
Jødekirsebær), Herba mimosa Brasiliana (Mimosa pudica), La- 
chryma Jobi (Coix lacryma), Malus Punica (Punica granatum, Gr a- 
natæble). Af Narcisser opføres 33 Arter og Sorter i Listen, endvidere 
Rapunculus hortensis fl. dilutiore pallido (Phyteuma spicatum) og 
Staphisagria (Delphinium staphysagria). 

Da en Del af disse Planter kun kan dyrkes under Glas, maa der 
nødvendigvis have været Væksthuse, det vides da ogsaa, at der i 1642 

22 


338 DANMARKS HAVEBRUG OG GARTNERI 


fandtes et forfaldent Blomsterhus i Rosenborg Have, og Sperling om- 
taler det (Æstuariet) ogsaa i sin Selvbiografi. 

Det har været noget ganske nyt for Urtegaardsmændene at have 
saa mange forskellige Planter at påsse, og Sperling har rimeligvis 
været vanskelig at tilfredsstille. Han klager gentagne Gange over 
Urtegaardsmanden David Konigs Dovenskab og Uforfarenhed, og 
navnlig er der efter hans Udenlandsrejse 1641 udgaaet mange Plan- 
ter saavel i Haven som i Æstuariet. At faa Konig afskediget lader sig 
imidlertid ikke gøre, da han ved at gifte sig med en Slægtning af 
Vibeke Kruse havde skaffet sig en høj Beskytterinde. I 1644 faldt 
Sperling selv i Unaade hos Kongen og afskedigedes fra sine Embeder. 
I hans Sted ansattes Helvig Dieterich som Bestyrer. Haven sy- 
nes derefter, hvad angaar de botaniske Samlinger, at være gaaet i 
Forfald. 


EFREDERTREV-KONGEFLIGE BOTANISKE HAVE 
NEDSEREDERIKS: HOSPITAL 


Fr V, som havde store Planer med Hensyn til Bebyggelsen 
af Arealet mellem St. Annæ Plads og Toldbodvejen, lod i Aaret 
1752 Grundstenen lægge til Frederiks Hospital. Hertil skulde knyttes 
en botanisk Have, som kom til at ligge Nord for Hospitalet. Haven 
ar gennemskaaret af Amaliegade og faldt saaledes i to Dele, en min- 
dre, vestlig, 11,106 [|] Al. og en større, østlig, 24,539 [] Al. Til Leder 
af denne Have var valgt Bayreren Georg Christian Oeder, som 
i 1752 ved J. H. E. Bernstorff var indkaldt her til Landet. Det 
var fra først af Meningen, at Oeder skulde have været ansat ved 
Universitetet, men da dette ikke lykkedes, udnævnte Kongen ham til 
Medicus paa Hospitalet og overdrog ham at paabegynde det berømte 
Værk ,,Flora Danica" samt lede den ny Have. 

Havens første Gartner var Kristian Hansen, som fra 14. Maj 
1753 var ,,Gartner ved den anlæggende Botaniske Have og ved det 
ny Kgl. Hospital paa Amalienborg" (Bestalling af ”/, 1753). Anlæggel- 
sen af Haven skred kun langsomt frem. Dels var Oeder meget paa 
Rejse, og dels var det Terræn, Haven skulde lægges paa, ikke umid- 
delbart anvendeligt til Dyrkning af Planter; det maatte først fyldes 
op og bearbejdes. Omkring 1760 var den mindre Have bleven no- 
genlunde færdig, den større Have vedblev derimod at henligge uop- 
dyrket, skønt der var rejst Mur udenom den saavelsom udenom den 
mindre. Ogsaa Drivhuse blev der bygget i denne Have, og i 1763 
aabnedes den for Publikum. 

Oeder var et stridbart Gemyt, han kunde heller ikke forliges med 
Kristian Hansen, som derfor i 1762 afskedigedes, men med fuld Gage: 
300 Rd + 100 Rd. til Husleje. I hans Sted ansattes Friederich 
Christian Voigt som Gartner ved Haven (Bestalling ”/, 1762). 

Havens Drift var imidlertid dyr, og i 1770 indkom Partikulær- 
kammeret derfor med en Forestilling til Kongen om at gøre en For- 
andring gaaende ud paa at overlade den vestlige Del af Haven til 
Universitetet og den østlige til Kommerciekollegiet. (Se videre om 
Havens senere Skæbne under Universitetets 2den Have, S. 303.) 

22% 


340 DANMARKS HAVEBRUG OG GARTNERI 


Ogsaa ved Sorgenfri Slot skal der have været en botanisk 
Have, om hvilken dog næppe noget historisk er levnet. I Parken 
voksede i sin Tid Asarum europæum og endnu findes Phyteuma spi- 
catum dér. Begge disse Planter kunde tænkes at være Relikter fra 
den botaniske Have. 


FORSTBOFANITSK: HAVE. VED. CHARLOTTENLUND 
OGFDEN'KONGELIGE-VETERINÆR- OG 
LANDBOHØJSKOLES HAVE 


NDEN der i 1863 paa Landbohøjskolen blev oprettet Kursus for 

Havebrugere og Skovbrugere, foregik Undervisningen i disse Fag 
andetsteds; Havebrugerne undervistes paa deres Læresteder, medens 
Skovbrugerne nød Undervisning paa den polytekniske Læreanstalt og 
blev examinerede af en Forstexaminationskommission, som i 1833 
var bleven dannet med Jonas Collin som Formand. Denne Exa- 
minationskommission indsendte i 1838 et Andragende til Rentekam- 
meret, i hvilket gjordes gældende, at den botaniske Have ,,ikke til- 
fredsstiller de Fordringer som et grundet Studium af Forstbotaniken 
gør til et sligt Anlæg, og at det derfor var nødvendigt, at et eget, i 
Omegnen af Hovedstaden beliggende Anlæg dertil bragtes tilveje,” 
og der indsendtes derefter fra Rentekammeret allerunderdanigst Fo- 
restilling derom til Kongen. 

En Kommission, bestaaende af Professorerne J. F. Schouw og 
Fallesen, Botanisk Gartner Mørch og Plantageinspektør Sch å f- 
fer, blev nedsat for at bringe et passende Sted i Forslag. Stedet, der 
valgtes, var 8 Td. Land i det sydvestlige Hjørne af Charlottenlund 
Skov, og Planen til Anlæget blev udarbejdet af Gartner Mørch. Ved 
Jordens Bearbejdning, Opførelsen af et Hus for Opsynsmanden, Ind- 
hegning af Arealet og Beplantningen af Arboretet havde Kommissio- 
nen Bistand af Skovrider Riedewaldt. Udgifterne ved Havens An- 
læg var beregnede til 1820 Rd. Kommissionen vedblev iøvrigt at be- 
styre Haven og ved Professor Fallesen og Gartner Mørchs Død blev 
disse erstattede med Forstdocent Hansen og Botanisk Gartner W eil- 
bach; iTidens Løb undergik Kommissionen flere andre Forandringer. 

Da Forstundervisningen i 1863 blev henlagt til Landbohøjskolen, 
udtalte Ministeriet, at omend den forstbotaniske Have nu ikke mere 
var nødvendig til Undervisning i Forstbotanik, maatte det dog anses 
for ønskeligt, at dette skønne Anlæg bevaredes og udvikledes. Mini- 
steriet mente videre, at Haven fremtidig kunde tjene som Forsøgsskole 
for Træer og Buske, der vilde kunne trives i vore Skove og tillige 
burde Træer og Buske, som havde Betydning for Havebruget, optages 
i Samlingen. 






den hel ME AGE! E. 








SE ER PE 






n Sne Ba (DM må Ar S 
mm sa al Ak mr JA tå d 
ar AN in Ø RA AR vd FR 
Me an ff: i AL pen 4) ' DE 

Se HV OSSE d å 
(GT æn mm sø an dt ' == 
NE KT fi EN 


t, m ke Vi == am 


km Y É 
Ya SEE 


MÅL LL. ve EL IN 







he 

















=== = == === 























CAR TA SEJ ASAA DDT ONDERE 


Fig. 143. Plan over Forstbotanisk Have 1847. 


BOTANISKE HAVER f 343 


Et i den Anledning nedsat Udvalg — Overførster Eide, Profes- 
sor Å. S. Ørsted og Gartner Bentzien — udarbejdede en Be- 
tænkning, som blev indgivet til Ministeriet det paafølgende Aar. Ud- 
"valget tilraadede bl.a. en Udvidelse og Forøgelse af enkelte større 
Familier, især af Egene og Naaletræerne samt en Bortrydning af 
visse Bevoksninger i Havens Udkanter for at give Plads til andre 
Planter, som bedre kunde tjene Havens Formaal. Med Hensyn til 
Ministeriets Forslag om at anvende Haven delvis som Forsøgsskole 
for Skovtræer, mente Udvalget dette maatte indskrænkes til Iagtta- 
gelser over Træernes Haardførhed; ligeledes mente man for Havebru- 
gets Vedkommende særlig at kunne anbefale Plantning af Træer og 
Buske, til hvis Haardførhed og Dyrkningsmaade man endnu ikke 
havde fuldt Kendskab. Haven tænktes lagt under det tilstødende 
Skovdistrikts Styrelse, men man formente det heldigt, at en Op- 
synsmand, som tillige skulde passe Haven, bibeholdt Bolig i denne. 
Udførelsen af de af Udvalget foreslaaede Arbejder bifaldtes af Mini- 
steriet og det overdroges Bentzien at lede disse Arbejder samt at føre 
det fremtidige gartneriske Tilsyn med Haven. 

Fra Landbohøjskolen fremkom nu imidlertid Forslag om, at Be- 
styrelsen af Forsthaven lagdes ind under Højskolen, og i Skrivelse af 
27. Maj 1869 bifaldt Ministeriet denne Ordning. Efter dette Tids- 
punkt har Forstbotanisk Have været under samme Ledelse som Høj- 
skolens egen Have. 


Ved Veterinærskolens Have paa Christianshavn havde der været 
en lille botanisk Have, hvor der dyrkedes Lægeplanter, men denne 
har aabenbart kun haft ringe Betydning. 

Da nu Veterinær- og Landbohøjskolen i 1858 skulde oprettes un- 
der ny og rummelige Forhold, ønskede man ogsaa at give den en 
Have, som kunde danne en værdig Ramme om Skolen og tjene som 
et fyldigt Undervisningsapparat for Botaniken. Den Komité, der 
skulde udarbejde Planen for Havens Anlæg, bestod af Professorerne 
Fenger, B. S. Jørgensen og Joh. Lange, Konferentsraad 
Drewsen samt Gartnerne Bentzien og J. F. Koch. Det over- 
droges Bentzien at udarbejde en Plan til Haven (se Fig. 144). Forde- 
lingen af Træer og Buske blev foretaget af Professor Lange med 
Bistand af Etatsraad Rothe og Botanisk Gartner Weilbach; Jord- 
behandlingen saavel som Beplantningen af Arealet blev foretaget af 
Gartner Wagenb last. 

Da Undervisningen i Havebrug blev henlagt til Landbohøjskolen 
i 1863, blev der til den 7”/, Td. Land store botaniske Have lagt de 
" 2 Td. Land, som endnu dyrkes som Frugthave og 1'/, Td. Land be- 
stemt hovedsagentlig til Køkkenhavedyrkning. 

Ved Tidernes Ugunst er begge Haver bleven beskaarne og delvis 
omlagte, værst er det i saa Henseende gaaet ud over Forstbotanisk 


"IZ8T a48H suadJfoysføyoqpueT 1940 UWjgq "ppl "SI 


(E 0 05 00 02 & & 0 


2 


— 


0 0 


000% - 1 PIO Yao Os 











" hon 
T 





H 





NH il nm 





FRE 


lyt tit 








eber 





















bEVTE 











KA 


NÅGÅ 


N 


w 


Å 


Av 
SÅ 
AN 



































BOTANISKE HAVER 345 


Have. I 1895 maatte denne afgive "/, Td. Land langs Vestsiden til 
Jærnbanestationens Udvidelse og samme Aar maatte den afstaa et 
smalt Jordstykke langs Nord- og Østsiden for at der kunde skaffes 
Plads til Anlæggelsen af en privat Vej. Landbohøjskolens Have mi- 
stede i 1892 et mindre Jordareal, fordi Højskolens Hovedbygning 
skulde udvides. ; 

De to Haver har siden 1869 staaet under samme Styrelse. Først 
havde Skolens Botaniklærer den botaniske Del under sig, Havebrugs- 
docenten den økonomiske Have under sig, men senere overtog den 


fø tt ; 








Fig. 145. Fra Forstbotanisk Have. Stenhøjen. 


sidste Driften af begge Haverne, dog saaledes, at Botaniklæreren 
havde Indseende med Nyplantninger og Rydninger i Arboretet saa- 
velsom med Ændringer i det urteagtige botaniske System. Haveassi- 
stenten var dels stillet til Raadighed for Havebrugslæreren, dels for 
Botaniklæreren, han havde Bolig i Haven og havde Tilsynet med den 
daglige Drift. I denne Stilling har en Række bekendte Havebrugere, 
nemlig Tyge Rothe, St. Nyeland, D. T. Poulsen, L. Helweg 
og Alfr. Bruun virket. I 1892 ansattes en fast Gartner, hertil ud- 
nævntes Alfr. Bruun, som skulde have den daglige Drift af Haven 
under sig med Ansvar overfor et Haveudvalg bestaaende af Lærere 
ved Højskolen. Da Bruun i 1902 blev Lærer ved Skolen, blev han 
tillige Efor for Haven og samtidig ophævedes Haveudvalget. 

I Forsthaven blev der fra først af — som før nævnt — især plantet 
Ege og Naaletræer, af disse er yderst faa tilbage, og de store Naale- 
træer, som hører til denne Haves Seværdigheder, er for største Delen 
plantede 1864. Men i 80erne og i Begyndelsen af 90erne blev der 


346 DANMARKS HAVEBRUG OG GARTNERI 


atter plantet en overordentilg Mængde Coniferæ, som daværende Do- 
cent Carl Hansen skaffede til Veje fra mange forskellige Ste- 
der. ! 
Der er i Tidernes Løb i Forsthaven ført Bog over, hvad der er 
plantet, saaledes at man som Regel ved Besked om hvert enkelt Træs 
Alder paa Stedet. I 1845 rummede Forsthaven 332 Arter og 32 Varie- 
teter, dette fremgaar af en Fortegnelse, som Botanikeren Kamp h ø- 
vener, der paa det Tidspunkt vikarierede for Professor Schouw 
som Lærer for Skovbrugerne, udarbejdede og som foreligger trykt. 





Fig. 146. Fra Forstbotanisk Have. 





I ,,Fortegnelse over de i Veterinær- og Landbohøjskolens Have og i 
Forsthaven dyrkede Frilandstræer og Buske" er opført 681 Arter og 
Varieteter, og i 1908 regnedes Tallet til c. 1000, hvoraf c. 300 var 
Naaletræer. Forsthaven er, trods sin bekvemme Beliggenhed, ikke 
særlig stærkt besøgt, dette skyldes sikkert udelukkende Ukendskab 
hos Publikum til Haven, men den har en fast Vennekreds, som sæt- 
ter overordentlig Pris paa den og som vil beklage, om den i Frem- 
tiden skulde blive nedlagt. 

Landbohøjskolens Have nyder en stor Popularitet hos den frede- 
riksbergske Befolkning, den er af en egen Hygge og Skønhed; den 
smukke Stenhøj, den lille Sø med den gamle Sørgepil, der ligesom 
danner hele Havens Midtpunkt, og de talrige Foraarsblomster, som 
myldrer frem under Arboretets Træer, virker særlig tiltrækkende paa 
Publikum. 

Havens Arboret rummede i 1871 (ifølge Joh. Langes Liste) 899 Arter 
og Varieteter, medens i 1908 Antallet ansloges til 1020. Af Stenhøjs- 


BOTANISKE HAVER 347 


planter dyrkes nu 830 Arter og Varieteter, af Stauder 550 og af 
Bregner 53, endvidere af botaniske urteagtige Planter i Systembe- 
dene, i Skovbede, Mosebede, Sandbede og Lyngbede 630 Arter (alle 
disse Tal fra 1918). Planter under Glas findes dels i nogle mindre 
Bakker, dels i et ældre ensidigt Hus med to Afdelinger og et nyere 
Saddeltagshus; i det første af disse findes forskellige Varmhus- og 





ea = = = 


Fig. 147. Parti fra Eandbohøjskolens Have. Stenhøjen og Dammen. 


Koldhusplanter, i det sidste dyrkes særlig Florblomster som Cinera- 
ria, Schizanthus og Chrysanthemum. Af Væksthusplanter og Ud- 
plantningsplanter dyrkes 650 Arter og Varieteter, og af Sommervæks- 
ter c. 580 Arter og Sorter. 

Haven beskæftiger — heri indbefattet saavel den botaniske som 
den økonomiske Del — en fast Gartner samt af Medhjælpere og Gart- 
neriarbejdere c. 9 Mand Aaret rundt. 


BOTANISKE HAVER 
VED LÆRDE OG KOMMUNALE SKOLER 


ED SORØ AKADEMI har der tidligt været anlagt en botanisk 

Have, idet Tyskeren Joachim Burser, som virkede ved Aka- 
demiet omkring 1625, har anlagt en botanisk Have. Om dens Anlæg, 
hele Liv eller dens Indhold af Planter ved man imidlertid yderst lidt. 
Lind skriver, at den oprindelig var Klosterhave og efter Bursers Tid 
har været benyttet af Apothekerne der paa Stedet for til Slut at gaa 
over til at blive Akademihave. I 1822 anlagdes atter paa Akademiets 
Grund i den saakaldte Plantagehave en botanisk Have paa en rekt- 
angulær Plads Vest for Gartnerboligen. Her dyrkedes et betydeligt 
Antal Frilandsplanter, og efter at der i 1829—30 var opført et Vækst- 
hus, blev der ogsaa dyrket Planter fra varmere Lande. Ved denne Tid 
var der ansat en Gartner, Hans Ditlev Nissen, ved Akademiets 
Have, han var i 1802 fulgt efter Carl Frederik Døllner, der i 
Aarene 1800—02 havde været Akademiets Gartner. Nissen drev ved 
Siden af sin Gartnergerning ved Haven en Del Handel med Planter, 
og formodentlig har der ligefrem været hjemlet ham Ret hertil for at 
bøde paa den sikkert meget ringe Pengeløn. Af Annoncer i ,,Stats- 
tidende” for den Tid ser man, at han har dyrket en ikke ubetydelig 
Mængde Træer og Buske med Salg for Øje. Efter hans Død 1834 
afkøbte Akademiet hans Bo for 50 Rdl. en Del Potteplanter, som han 
efterlod sig. 

Aaret forud var det blevet overdraget Lektor ved Akademiet 
Jacob Hornemann Bredsdorff at udarbejde en beskrivende 
Katalog over Havens Plantesamling. Det meget omfangsrige Værk, 
som er affattet paa Latin, blev afsluttet i Løbet af 3 Aar og inde- 
holder Navne paa 1422 højere Planter, fordelte paa 665 Slægter. Det 
var fra først af Meningen, at det skulde trykkes, men dette naaedes 
aldrig; Manuskriptet forefindes i Sorø Akademis Bibliothek. Af Vækst- 
husplanter dyrkedes dengang (1837) bl.a. Begonia discolor, Ilicium 
anisatum, Justicia adhatoda og 6 Kaktus-Arter. 

Efter Nissens Død var F. A. Voetmann Gartner indtil 1880; han 
fulgtes af E. C. Thayssen, som atter i 1914 blev efterfulgt af den 
nuværende Gartner F. C. Lønow. 

Omkring Aaret 1857 blev Sorø-Haven omlagt af R. Rothe, som 


BOTANISKE HAVER 349 


anlagde den nuværende Park; ved denne Lejlighed blev den botani- 
ske Have henflyttet til en ret smal Strimmel Jord, beliggende mellem 
Danneskjolds Allé og Priorgade og her blev Planterne anbragte i 
Smaabede. Da Musæet i Slutningen af 90'erne blev opført paa en Del 
af denne Grund, blev den botaniske Have nedlagt. Der findes nu 
ingen botanisk Studiehave ved Akademiet; Materialet til Undervis- 
ningen i Botanik søges udenfor Terrænet. Haven og Omegnen bærer 
dog Spor endnu af, at der har været dyrket fremmede Planter, idet 
man dér finder flere forvildede Planter (f. Eks. Galinsoga parviflora, 
Myrrhis odorata og Oxalis stricta). 


Ved HERLUFSHOLM blev der i Aaret 1890 af Forstanderen ud- 
vist et Stykke Jord af ,,Store Granplantage" paa ca. Sæde Eands 
Størrelse med det Maal for Øje dér at indrette et ,,Pinetum”, d.v.s. 
en Samling af Naaletræer. Anlægget heraf lededes af Adjunkt M. 
Traustedt, som sikkert ogsaa er den, der fra først af har taget 
Initiativet til Pinetets Oprettelse; men i denne Forbindelse maa Plan- 
tør Jacobsen paa Hæsede ogsaa nævnes, som en Mand, der ydede 
sin betydelige Del til, at Pinetet kom i Gang. I Aaret 1891 udkom en 
Beretning om Anlægget og en Liste over de Arter, som var saaede 
eller plantede og i en anden Beretning, udgivet i 1896, findes en ud- 
førligere, delvis beskrivende Liste, omfattende 164 Navne paa Naale- 
træs-Arter, hvoraf dog kun de 102 var repræsenterede, medens Resten 
for Størstedelen ventedes snarligt anskaffede. Desværre er den Plads, 
som i sin Tid blev valgt til Samlingen, meget uheldig; Pinetet er 
nemlig anlagt paa en Bakke med løst, stenfrit Sand, og Betingelserne 
for Træernes gode Vækst er derfor meget daarlige. Baade Traustedt, 
som ledede Pinetet til sm Død, og den nuværende Leder, Adjunkt 
J. Ferdinand, har vist stor Interesse for Pinetets Trivsel, og havde 
der været gode Jordbundsforhold til Stede, er der ingen Tvivl om, at 
der vilde være bragt noget ganske udmærket ud af Samlingen; denne 
rummer, trods de daarlige Kaar, talrige smukke Træer og tæller nu 
ca. 200 Arter og Varieteter. 

Til Skolen hører ogsaa et botanisk System, hvor der dyrkes om- 
kring 550 Arter. Endvidere findes der, ligeledes tjenende den botani- 
ske Undervisning, et lille Parti med Plantesamfund (Klit-, Hede-, 
Mose-Planter) samt et betydeligt Stenhøjsparti, anlagt omkring Sko- 
lens Sceptictanks under Ledelse af Adjunkt Ferdinand. I disse 
Partier dyrkes over 1000 Plantearter. Fra Herlufsholms botaniske 
Have uddeles der gratis Planter til andre Skoler og til Private; den 
aarlige Uddeling andrager 200 å 300 Eksemplarer. 


Endnu kan nævnes, at der muligen allerede i Slutningen af det 
18. Aarhundrede har været en skolebotanisk Have i ODENSE. Peder 
von Westen har 1782 udarbejdet: ,,Plan og Charte til en botanisk 


350 DANMARKS HAVEBRUG OG GARTNERI 


Haves Anlæg ved Odense Gymnasium"; denne Plan findes ikke paa 
vore større Bibliotheker, og om Haven er bleven til Virkelighed, kan 
ikke oplyses. 


Forskellige Seminarier og Skoler her i Landet har i den nyere 
Tid i umiddelbar Tilknytning til Institutionerne udlagt mindre Årealer 
som skolebotaniske Haver. Her at gøre Rede for alle disse Haver, vilde 
dog føre for vidt, der skal blot kortelig gives Oplysninger om den 
vigtigste af Provinsens botaniske Haver, nemlig: 


DEN SKOLEBOTANISKE HAVE I AARHUS 


I 1905 blev der af ,,Jydsk Forening for Naturvidenskab" paa For- 
slag af cand. mag. P. Obel indgivet et Andragende til Aarhus Byraad 
om at indrette en skolebotanisk Have af en saadan Størrelse, at samt- 
lige Skoler kunde forsynes derfra med Materiale til Brug ved Under- 
visningen i Botanik. Det blev overdraget Lærer Poul Larsen at ud- 
arbejde en Plan for Anlæggelsen og Driften af en saadan Have, idet 
der samtidig hertil blev anvist et Stykke Jord paa 1700 Kv. Al. i 
»Jydsk Haveselskabs" Have. Gravning og Dræning af Jordstykket be- 
sørgedes ved Haveselskabet, ligesom dette de følgende Aar lod udføre 
Gravning og Lugning; Stykkets Tilplantning og Tilsaaning udførtes 
af Poul Larsen, og i de følgende Aar paatog samme sig ydermere Ud- 
levering af Planterne til Skolerne. Da Haveselskabets Have senere 
overtoges af Aarhus Kommune udvidedes Skolehavearealet til 4000 [] 
Al. Skolehaven staar dog vedvarende under Lærer P. Larsens 
Ledelse. 

I Haven findes nu ca. 300 Arter af de almindeligste Blomsterplan- 
ter, og det paatænkes at lade plante de danske Buske og Træer. 

Skolernes Benyttelse af Haven foregaar væsentlig paa den Maade, 
at der Sommeren igennem med 2 å 3 Ugers Mellemrum tilstilles Sko- 
lerne en Liste over, hvilke Planter, der er i Blomst eller Frugt, og 
efter denne Liste rekvirerer Botaniklærerne Materiale til Undervisnin- 
gen. Udlevering af bestilt Materiale foregaar hveranden Dag i 2 Timer. 
Antallet af Portioner, d.v.s. saa meget Materiale af hver Art, som 
anses nødvendigt til en enkelt Elev, er i Løbet af den Tid, Haven 
har eksisteret, vokset fra 4000 til 14,000. Haven besøges ogsaa af 
Lærere og Elever under Tilsyn i de Timer, da Skolehavens Leder er 
til Stede. 


PRIVATE BOTANISKE HAVER 


M-æcæ Privatpersoner har i Tidernes Løb næret saa megen Inter- 
esse for Botanik, at de i deres Haver har anlagt Samlinger af 
Planter, hvorfor disse Haver med en vis Ret har kunnet betegnes som 
botaniske. En Grænse, mellem hvad der kan regnes for en botanisk 
Have og hvad der blot maa betegnes som en Plantesamling, synes 
vanskelig at drage. De i det følgende nævnte Haver kan dog maaske 
nævnes blandt de bekendteste af saadanne private, botaniske Sam- 
linger. 

Klavs Urne, Domprovst i Lund, havde ved sit Herresæde Belte- 
bjerg i Skaane, anlagt en ,,besønderlig Urtegaard", som Henrick Smid 
omtaler med Berømmelse i 1546. 

Rottbøll omtaler, at Grev Johann Ludvig af Holstein paa 
»Sit kiere Ledreborg" lod indrette en Botanisk Have, ,,som med saa- 
dan en Hurtighed blev indrettet og forsynet med Planter, at den paa 
2 Aar var stegen mere end til 1200 Planter og pralede af en utallig 
Mængde Aloes, Cacti, Mesembryanthema, Euphorbiæ, Gerania, Hi- 
bisci, Malvæ, Solana etc.” Disse Planter var for Størstedelen ved Pro- 
fessor K. Friis Rottbøll forskrevne fra Holland. Andre Planter, 
som var mindre kælne, var anbragte i Bede paa Friland, ordnede 
systematisk. 


Ved Slottet FRYDENLUND fandtes i Slutningen af det 18. Aar- 
hundrede Samlinger af nordamerikanske Træer og Buske og de Co- 
ninckerne plantede i Haven ved DRONNINGGAARD ligeledes talrige 
sjældne træagtige Planter. 


Ved Slottet AALHOLM paa Lolland blev der i ældre Tider plantet 
et Arboret, som endnu i Slutningen af 1860'erne var ret righoldigt. 
I 1867 har Botanisk Gartner Weilbach i ,,Tidsskrift for Have- 
væsen” givet en Skildring af dette Arboret. Det fremgaar heraf, at 
Arboretet paa Aalholm er plantet i Tiden fra 1796 til 1814, medens 
en anden Del er plantet i Tiden 1830—1835, da Chr. Gentz var 
Gartner ved Slotshaven. En Del af Arboretet kaldtes ,,Systemet"”, 
hvor Træerne har været plantet i Rækker hver Slægt for sig og saa- 
ledes synes at have haft en ren botanisk Ordning. I Weilbachs Be- 


352 DANMARKS HAVEBRUG OG GARTNERI 


skrivelse af Aalholm-Arboretet er optaget ca. 100 Træer og over 50 
Buske; han havde dog kun medtaget de vigtigste Arter eller største 
Eksemplarer; en Del af Arterne kunde ved hans Besøg dér ikke 
identificeres. I ældre Tider har Samlingen været langt righoldigere. 
Det fremgaar af gamle Lister over Indkøb af Planter, at Haven har 
faaet meget righoldige Samlinger fra Booth's berømte Planteskole i 
Flotbeck. Indkøbet begynder i 1829 og kulminerer 1831. Sidstnævnte 
Aar blev bl.a. købt 30 store Plataner, over 150 Quercus coccinea, 
100 Pinus nigra, 29 Pinus tæda o.m.a. Planter i stort Antal, foruden 
adskillige Hundrede Arter og Former i enkelte Stykker. I Følge 
»Dansk Haugetidende", 1. Aargang, 1849, omfattede Samlingen i 
1833 ikke mindre end 1500 forskellige Træer og Buske. 


Noget senere — henimod Midten af det 19. Aarhundrede — er 
Haven i SNOGHØJ, som ejedes af Generalkrigskommissær Riegels, 
en i Plantedyrkning særdeles interesseret Mand, bekendt for sine 
Samlinger af Træer og Buske. I samme Tidsperiode har GJORSLEV 
HAVE paa Sjælland righoldige Samlinger af Frilandsplanter. 


Apotheker GUSTAV LOTZE, der virkede i Odense, nærede en 
dyb Kærlighed til Havebrug og Botanik og samlede i sin Have ikke 
alene talrige sjældne Træer, men anlagde ogsaa store Stenhøje med 
righoldige Samlinger af Alpeplanter. I Væksthusene, der havde en 
samlet Længde af 100m og hvis Palmeafdelinger maalte 10m i Høj- 
den, havde han samlet tropiske Nytteplanter og andre Planter fra de 
varme Lande, navnlig Orchideer. 


Blandt private botaniske Haver bør HÆSEDE PLANTESKOLE 
ikke forbigaas, idet den i al Fald i sin Tid var ganske enestaaende 
1ESInRArE: 

Planteskolen var anlagt i Hæsede Skov, nærmest med det For- 
maal for Øje at forsyne Gisselfeld Have med Planter; men efter at 
Hans Jacobsen i Midten af 30'erne i forrige Aarhundrede som 
ganske ung Mand havde faaet Planteskolen at passe, fik denne efter- 
haanden sit særlige Præg. Jacobsen, der var kommen til Pladsen uden 
særlige Forkundskaber, erhvervede sig ved sin Virken som Plantør 
og ved sine flittige Studier meget omfattende Kundskaber paa Bota- 
nikens Omraade, og da han var Samler om en Hals, fik han bragt 
mange sjældne Træer og Buske til Veje og snart forsynede han andre 
Planteskoler, botaniske Haver og Herregaardshaver nmåed Plantema- 
teriale af sin Rigdom. En Vandring gennem Planteskolen, især i 
Følge med Jacobsen, var overordentlig lærerig, og endnu huser Plante- 
skolen, selv om den har mistet noget af sit Præg, store Planteskatte. 


Dr. F. BØRGESENS Have i Hellebæk udmærker sig særlig ved 
sine smukke Samlinger af Naaletræer — her findes f. Eks. et Eksem- 


BOTANISKE HAVER 393 


plar af Sequoia sempervirens, et Træ, som ellers ikke vil trives paa 
Friland hos os — ved sine prægtige Stenhøje og sine Samlinger af 
Smaabuske. Stenhøjene rummer meget rige Samlinger af saavel 
Primler og Stenbræk som højnordiske sjældne og vanskelig dyrke- 
lige Arter af Cassiope, Juncus og Diapensia samt tillige mange Fri- 
landsbregner og Orchideer. Blandt Smaabuskene indtager Cotoneas- 
ter, Berberis og Rhododendron den mest fremragende Plads hvad 
Årtsantallet angaar; af den sidstnævnte Slægt findes 64 Arter, et saa 
stort Antal Arter kan sikkert ingen anden nordisk Have for Tiden 








pr" en did - v . 
Fig. 148. Fra Dr. Børgesens Have, Hellebæk. 
opvise. Nævnes kan ogsaa, at der af Daphne dyrkes 9 og af de ny- 
zelandske, buskagtige Veronica 5 Arter. 


Mag. art. M. LORENZEN, Stettemark, Holte, har i sin Have an- 
lagt storslaaede Stenpartier med rige Samlinger af Alpeplanter. Mag. 
Lorenzen samler endvidere Buske og mindre Træer, af de første 
findes 564, og af Træer 254 Sorter og Arter. Af egentlige urteagtige 
Alpeplanter dyrkes i denne Have ca. 700 Arter, Hybrider og Variete- 
ter, deraf 100 Sawzifraga og 100 Primula, dertil 70 Smaabuske og 
Dværgbuske. Paa Stenhøjene er der endvidere 24 Anemone-Årter og 
forskellige finere Løgvækster, medens der andetsteds i Haven er plan- 
tet 100 Arter og Sorter af Crocus. 


Lolland besidder nuomdage to Privatarboreter. Det ene findes paa 
tal] 


23 


354 DANMARKS HAVEBRUG OG GARTNERI 


KNUTHENBORG, i hvis over 500 hekt. store Park flere sjældne 
Træer er plantede, ofte i et betydeligt Antal Eksemplarer. Denne Park 
er under Overgartner J. K. Jørgensens Ledelse. 


Det andet lollandske Arboret findes paa HARDENBERG. Her er 
der mere Il: …t an paa et større Artsantal; det mindre Areal, som her 
raades over, begrænser ganske naturligt Individantallet. 

De vigtigste Træer i Hardenberg-Arboretet af nogen betydelig 
Størrelse er plantede omkring 1840, andre for omkring 40 Aar siden. 
Endelig er der i Tiden fra midt i 90'erne og op til den nyeste Tid 
indkøbt mange sjældne og smukke træagtige Planter, saavel Naale- 
træer som Løvtræer, bragt til Veje af den for Samlingerne saa stærkt 
interesserede Overgartner Chr. Jørgensen. 

I Samlingen findes en over 100-aarig Corylus colurna, 11m høj, 
en ca. 80-aarig Liriodendron tulipifera, 17m høj, 40-aarige Pinus 
peuce, Magnolia Soulangeana, Prunus lusitanica og T huyopsis dola- 
brata; en 210-aarig Tilia platyphylla, 26m høj, en 25-aarig Picea 
omorica, 6Gm høj og af samme Alder en Liquidambar styraciflua, 
6m høj, endvidere Cedrela sinensis og Gymnocladus canadensis. Af 
Naaletræer findes ca. 200 Arter og Former, fordelte paa 23 Slægter, 
deraf har Abies alene 31 og Pinus 38 Arter og Former. Af Løvtræer 
findes op imod 850 Arter og Former, fordelte paa 177 Slægter. Som 
særlig rigt repræsenterede Slægter kan nævnes Cratægus med 33, Pru- 
nus med 25, Quercus med 39, Spiræa med 64 og Acer med 37 Arter 
og Former. 


LITTERATUR-FORTEGNELSE 


Vigtigste Literatur: Otto Sperling: Hortus Christianæus 1642. — Joh. de Buch- 
wald: Specimen medico-practico botanicum, 1721. — Skrifter som udi det kjøben- 
havnske Selskab af Lærdoms og Visdoms Elskere ere fremlagte (Rottbøll) 1770. — 
J. W. Hornemann: Om den Kgl. bot. Haves nærværende Forfatning (i Athene) 
1813. — F. Thaarup: Nogle Bidrag til det Kbh. Universitets bot. Haves Historie (i 
Telegraphen) 1826. — Fr. Weber: Hortus Kiliensis, 1822, — L. Engelstoft og E. C. 


Verlauff: Udsigt over Kjøbenhavns Universitetsbygnings Historie, 1836. — F. 
Thaarup: Materialer til den danske Stats Havekulturs Historie 1843. — F. 
Didrichsen: For hundrede Aar siden (i Naturhistor. Tidsskrift) 1869. — Beretning 
om Univervitetets paatænkte nye Botaniske Have, 1870. — Ratjen: Geschichte der 
Universitåt zu Kiel 1870 — Joh. Lange: Fortegnelse over de i Veterinær- og Land- 
bohøjskolens Have og i Forsthaven .... dyrkede Frilandstræer .... 1871. — Joh. 


Lange: Erindringer fra Universitetets Botaniske Have ved Charlottenborg (i Bota- 
nisk Tidsskrift) 1875. — H. Matzen: Universitetets Retshistorie 1879. — J. C. Ja- 
cobsen og Tyge Rothe: Beskrivelse af Væxthusene i Universitetets botaniske Have, 
1879. — Birket Smith: Dr. Otto Sperlings Selvbiografi, 1885. — Holger Hansen: 
Den botaniske Have ved Amalienborg (i Historiske Meddelelser om Kjøbenhavn, 
3. Bind. — Den Kongelige Veterinær- og Landbohøjskole 1858—1908, Festskrift (Alfr. 
Bruun: Haverne) 1908. — J. Reinke: Der ålteste Botanische Garten, Kiels, 1912. — 
Forskellige Aargange af Badens Universitetsjournal, Engelstoft: Universitet og 
Skole-Annaler, Selmers Akademiske Tidender, Lindes Meddelelser; Goos, Matzen, 
Munch-Petersen: Aarbog for Kjøbenhavns Universitet; Chronik der Universitåt zu 
Kiel. Have-Tidende. — Dansk Hauge Tidende. — Tidsskrift for Havevæsen. — 
Gartner-Tidende. — Akter fra Rigsarkivet (Sjæll. Miss. 374 (1770), 1441 (1777) 
615 (1779), 239—41 (1788), Partikulærkammerets Ordreprotokol 1770. 





Resumé of the section »Botaniske Haver« from 
Svend Bruun og ÅAæel Lange: Danmarks 
Havebrug og Gartneri til 1919, Kbh. 1920. 


| soes gardens in the sense we now apprehend the notion were 
not known until the year 1545, when the university of Padua 
founded such an institution. Other universities followed suit, thus gar- 
dens were laid out in Bologna 1568 and in Leiden 1577. It will be 
seen, that the universities had the leadership in this regard, but later 
on royal as well as imperial, municipal and private gardens were 
founded. ' 


FHE FIRST-BOTANICAL. GARDEN; OF THE COPENHAGEN 
UNIVERSITY 
(HORTUS MEDICUS) 

In the year 1600 a new residence for a professor was to be erected 
close to the university of Copenhagen, and on this occasion the king 
Christian IV endowed to the university a site on which was to be laid 
out a botanical garden. This garden had no gardener, neither had it 
any income; the professor who was assigned to the lodging was bound 
to keep the garden in proper order. No alteration took place in these 
respects until Rasmus Casper Bartholin in 1696 presented the 
university with 1200 Rigsdaler, the interest of which should be used 
to buy seeds and to pay a gardener. 

We do not hear much of the garden during the coming Z/4. of a 
century; in 1719 the plants of the gardens have been transplanted 
under the guidance of Johannis de Buchwald and arranged 
alphabetically; after the great conflagration of Copenhagen in 1728 
the garden is reduced in size because of the regulation of the adjoining. 
streets, and about the middle of the 17th century we hear that a gar- 
dener is attached to the institution for very modest wages. 


DHE SECOND. BOTANICAL: GARDEN OF THE GOPENHAGEN 
UNIVERSITY 
(OEDERS GARDEN) 
The garden was really too small for the purpose, and the king, 
who took a certain interest in the garden, and who in 1769 haåd 
endowed it with 2500 Rigsdaler, promised to enlarge it with some 


adjacent gardens. This plan was not carried out, but the king resolved 
instead that a much larger botanical garden, which had been founded 
by Oeder under protection of the late king (Frederic V.) in another 
part of the town (at Amalienborg) should be given to the university. 
This garden measured about 11000 squarefeet ground, it was well 
furnished with plants and in a good order. Rottbøll, professor of 
botany and medicine at the university, had to take the charge of the 
garden and as curator was appointed J. W. Kæsemacher, a skilful 
gardener, who had spent some years in the Amsterdam-garden. 

Only a few years the garden was allowed to remain here. Neither 
was the soil so good as it ought to be, nor the situation owing to lack 
of shelter. 


THE THIRD BOTÅANICAL GARDEN TOFETHE COPENHAGEN 
UNIVERSITY 
(CHARLOTTENBORGHAVEN) 

The king therefore presented the university with a new site for its 
garden namely the garden of Charlottenborg. Furthermore he re- 
bought the Amalienborg-garden for 6000 Rd. The removal began in 
1778, and in 1787 the professor Rottbøll and the privy counsellor 
Holmskjold who had arranged the new garden, were able to 
report to the king, that the buildings had been completed as proposed. 

On this new site the botanical garden, which had a size of 3 Td. 
Land (1/2 hektare), stayed for about 100 years, although the trustees 
of the institution soon were aware that it really was too small. Already 
in 1803 the wealthy privy counsellor Classen offered a fine site, rather 
large (4 hekt.) in the eastern suburb of the town, where the garden 
would have good conditions to thrive; but as the expenses by the 
transfer would be too enormous, the plan was abandoned. Now the 
garden was under the necessity of enlargement on the spot and really 
in 1811 it succeeded in obtaining an adjacent garden, and in the year 
1844 again it obtained an extension; after this the garden measured 
about 4, Td. Land (2 hekt.), in which size it remained unaltered 
until its removal in 1871. 

After the dealh of Kæsemacher in 1780 Niels Bache had 
the curatorship until 1793, when F. L. Hollbøll, the most skilful 
curator of the garden, was appointed to the post. He remained in the 
situation until his death in 1829 and was succeeded by O.J.N.Mørch 
who died in 1842. After Mørch A. Weilbach was curator during 26 
years, and his successor was Th. Friedrichsen. The directorship 
was in the earlier time of the garden in the hands of a commitee con- 
sisting of Rottbøll and Holmskjold; later on four persons had their 
seat in the commitee until 1817. Till this year the institution had 
partly been under the university and partly under royal protection 


3 


but was now fully laid under the university, and from this juncture 
the directorship has been attached to a single person. Famous bota- 
nists as E. Viborg, M. Vahl and J. W, Hornemann have had 
their seat in the directory. Hornemann was the first solo-director, 
in 1841 he was followed by J. Fr. Schouw who again in 1852 was 
succeeded by Liebmann. After his death in 1856 Joh. Lange be- 
came the director of the garden. 

The garden was well sheltered in its situation in the old royal gar- 
den, several planis were thriving well here and the garden was on 
the whole of a certain snugness. But it had several defects. Chiefly it 
was still too small and the desire to get it removed, was permanently 
increasing. Furthermore the soil was partly too gravely and bad, and 
partly excessive rich in mould; the houses that had been partly rebuilt 
in 1843 were not spacious enough. In 1856 Curator Weilbach there- 
fore very opportunely published a note in one of the principal news- 
papers (Fædrelandet) about the necessity of a renaissance of the 
garden on a new and more spacious site. But about 20 years should 
elapse, before a new garden was fulfilled, and Weilbach should not 
live to see his plan realized. 


THE FOURTH BOTANICAL GARDEN OF THE COPENHAGEN 
UNIVERSITY 
(THE PRESENT GARDEN) 

Being fully aware of the bad circumstances for the garden the 
university in the year 1857 and after the desire of the government 
took the initiative to form a committee that should make investiga- 
tions how and where to form a new garden. This committee consisted 
of Johan Lange the director of-the garden, Jap. Steenstrup, 
and J. G. Forchhammer, professors of the university. With diffe- 
rent alterations and augmentations it worked until the garden in 1874 
was finished on its present seat and it had during these years several 
difficulties to conquer. Very soon it was evident to the committee, that 
the best situation would be some part of the fortifications of Copen- 
hagen, that were destined to be demolished within a few years. 

In 1871 this seat was finally accepted for the purpose, and the 
same year 25000 Rigsdaler were granted to preliminary labours. The 
committee that now consisted of professor Johan Lange, professor 
A.S. Ørsted, the university-bursar Gjede, the brewer J. C. Jaco b- 
sen and the curator of the garden Th. Friedrichsen, had already 
in 1870 issued a statement: ,,Beretning om Universitetets paatænkte 
nye Botaniske Have" mostly intended as an information for the mem- 
bers of the parliament, and in which therefore was given a brief narra- 
tive of the history of the garden and the special reasons why a new 
garden was desirable. A plan (fig. 132) that followed this report showed 


E: 


some details of the intended garden. This report caused a lively dis- 
cussion in the garden-periodical: ,,Tidsskrift for Havevæsen"- about 
the site of the garden, the application of the ground, the construction 
of the houses and the design of the garden on the whole. 

It was from some hand viz. the lecturer in horticulture at the 
veterinary and agricultural college J. A. Dybdahl proposed to aban- 
don the present plan and to lay the garden close to the veterinary & 
agricultural college; but at all events he found it necessary to have 
an expert in landscapegardening to enter the committee. 

The government did not wish to alter the site of the garden, but 
on the other hand it willingly entered on the proposal to add some 
new members to the committee and in accordance herewith the 
landscapegardener H. Flindt and the royal gardener ThygeRothe 
were summoned as counsellors, the former should work out a new 
plan and the latter was charged in connection with J. C. Jacob- 
sen to give new plans for the houses. 

In the years 1871—74 the garden was fulfilled in correspondence 
with the new plans at cost of 642,000 Kroner, and the 1 of june 1874 
the garden was delivered in the hands of the directory of the garden, 
and the committee consequently dissolved. The garden was opened 
to the public 9. Oktober 1874. 

The garden in form an irregular square, had by its foundation an 
area of 92/4 hektares. It consists of a larger lower part holding the 
$000 square-meter large lake, towards the east and southeast the 
ground rises rather abruptly against the boulevard and the obser- 
vatory, situated outside the garden at the highest point of the mount 
(13 m.); towards the south and west the ground-elevation is more 
even and soon the surface is quite level; on this plateau the museum, 
the lodges for the curator and foremen and the laboratory are situated. 

The trees and shrubs are disposed all over the garden after a 
systematically point of view, and from the beginning the peren- 
nials were dispersed in the same way. In the western corner of the 
garden a little grove was planted to illustrate the north-american 
silva. In the centre of the garden the rockery was laid out on twin- 
hills attaining a hight of 13—14 m., from whence a splendid view can 
be had over the garden and the surrounding town. The greenhouses 
cover an area af 2400 sq. m. and have a volume of 12000 mZ, and the 
glass-area is 3100 m”; a greater part of the houses has two layers of 
glass. The highest point of the palm-house has an elevation of 19 m. 
In front of the houses is a basin with a fountain, flanked by two span- 
roofed houses. Also a water-lilyhouse, propagating houses, pits and 
forcing-frames are found in the garden. The warmingapparatus of 
the houses is based upon steamheating; the annual amount of coal 
consumed is about 400 tons. 

The committee that had arranged the garden had proposed the 





5 


museum to be built at the same time as the greenhouses, nevertheless 
its erection was poestponed until 1875 when the parliament granted 
112,000 Kroner for the purpose; a sum which really was too limited, 
for which reason the museum had to be built on a smaller scale as 
intended, to immense disadvantage and drawback for the institution 
during its existence. 

Johan Lange, who had been director for the garden about 20 
years and had been the chairman in the committee during its latter 
years, was in 1875 succeeded by F. Didrichsen, who held the 
office until 1885, when Eug. Warming took his place. Warming 
came to the garden just at a time when the botanists of Denmark 
earnestly were wishing to get a laboratory for plant-anatomy and 
plant-physiology, and these efforts resulted in the erection of the 
botanical laboratory. This was built in the western corner of the 
garden, on which occasion the garden in itself was diminished in area 
with "., hekt. and at the same time the northamerican arboretum had 
to be demolished. The building was fulfilled in 1890 at the cost of 
about 200,000 Kroner. 

Now professor Warming devised a scheme for a biological garden, 
and in order to get room for this the area of the students” garden 
was reduced to half its size. 

The polytechnicum that in the end of the 80ers had been built 
close to the botanical garden and was inaugurated in 1890 was socn 
getting too small for its purpose and the institution therefore wanted 
to extend upon the domain of the botanical garden. In compensation 
for the 0,3 hekt. the polytechnicum wished, the garden should have 
on rent in 50 years an adjacent municipal area of 0,7 hekt. extent. 
The garden entered upon the proposal and hereby it was enabled to 
get some improvements on its own ground. Wiz: the perennials could 
be removed to a better quarter of the garden instead of being spread 
about in the lawns. At the same time the garden gave a freer ad- 
mittance for the public as formerly. Prior to these alterations the cu- 
rator Th. Friedrichsen took his leave in 1902 after having been 
in activity for 34 years; he was succeeded by Axel Lange. And by 
the expiration of 1911 prof. Warming retired and was followed by 
C. Raunkiær. 

The staff of the institution consists of: the director managing the 
whole establishment with exception of the physiological laboratory 
that is under the guidance of another university-professor (at the 
present time professor W. Johannsen) with his assistant. At the 
botanical laboratory is appointed an assistant and a door-keeper. The 
museum and library has a museum-superintendent, a librarian, an 
ammanuensis, a conservatrice and a museum-official; the plants in 
the garden are revised by the garden-ammanuensis. The garden itself 
is under the guidance of the curator with the help of 3 foremen and 


6 


2 subforemen, 3 stokers, 4 guards and 20 gardeners and workmen; 
the number of work-days during the year amounts to 7000. 


In the previous lines four different gardens have been mentioned, 
but of course these gardens may be regarded as a single garden — 
the garden of the university and while we above have described 
the topographical, economic and personal circumstances of this insti- 
tution we shall now shortly regard on what lines the garden has been 
working, of what use it has been and how the plant-collections have 
increased or altered during the days past. 

In its earlier days the garden was a mere students” garden in which 
on the first line indigenous plants were cultivated, and furthermore 
it was a pleasure-garden for the professor occupying the residence. 
Ole Worm brought several foreign plants to the garden for instance 
Rheum rhaponticum, Scorzonera hispanica and Mimosa pudica. J. 
de Buchwald in 1720 issued a book with a description of a little 
more than 200 plants: ,,Specimen medico-practico-botanicum" which 
book the year after was translated into german by his son B.J.Buch- 
wald; however, the translation does not agree fully with the original 
not even in respect to the plants mentioned. Corresponding with the 
plants described, dried specimen of the species are mounted on white 
pages of the book. Allthough we can not take as granted, that all the 
plants embodied have been cultivated in the garden (thus Viscum al- 
bum and Lathræa squamaria have hardly been grown), we dare sup- 
pose, that the greater part of them have been to be seen there, and 
presumably other plants not mentioned have also been cultivated. Of 
plants not indigenous embodied in the book shall be mentioned: An- 
tirrhinum majus, Cannabis sativa, Cnicus benedictus, Cucumis sati- 
vus, Cyclamen europæum, Juniperus sabina, Lilium candidum, Phy- 
salis alkekengi and Veratrum album. 

The size of the collection of plants while the university had its 
garden at Amalienborg is not known, the garden took possession of 
the large collections, that Oeder had brought together, these embraced 
in 1763: 1500 species. In the Charlottenborg-period the number of 
species increased from 2855 (in 1784) to 9500 (in 1857) and now the 
garden can boast of 12000 species distributed over 23570 genera. The 
composition of the collection has altered during times past, changing 
after the taste of man or after the interest that the leaders of the in- 
stitution have paid in the different sort of plants. We know, that Ho l- 
bøll in 1803 bought 25 different Ericas and that he the same year 
cultivated 40 species of Mesembrianthemum; in the time of Mørch 
(1839) the garden had 75 species of the latter genus and at the same 
time 35 Acacias, 50 Pelargoniums, 10 Begonias, 17 Primulas and 27 
Saxzifragas, while the collection now (1918) embraces 28 Mesembrian- 





I 


themums, 13 Ericas, 25 Pelargoniums, 15 Acacias, 64 Primulas, 65 Be- 
gonias and 100 Saxzifragas. In 1834 the garden had only 3 tropical 
Orchids, now the number is 180. 

The garden has always had in its possession specimens of plants, 
that could be regarded ås prominently noteworthy on account of 
their age, their size or their rarity. The centre-plant in the palmhouse 
a Phoenix spinosa has in 1807 been brought to the garden and has 
now attained a height of 18 m. The Livistona chinensis (figured on 
page 326) was in 1919 cut down because it was getting too big, it had 
a total weight of 615 kg and carried the year mentioned not less than 
15 kg of fruits. In 1904—06 an Arenga saccharifera flowered and 
subsequently died as is the nature of this plant; it had an age of 75 
years. Several times Agave americana has been brought to flower and 
in 1836 Mørch had this species in flowering state exhibited to the 
public. He had bought the plant in Skaane (Sweden) and it was car- 
ried to Malmø by 8 horses and from that place sailed to Copenhagen. 
Of rarities in the open air, the Eriolobus trilobata (figured on pag. 328) 
ought to be mentioned. 

Although the garden now and then has received collections from 
travellers or collectors in foreign countries, still it is keeping or in- 
creasing its stock mainly through its correspondence with botanical 
gardens abroad, especially by exchange of seeds. The garden yearly 
receives 2000—3000 lots of seeds and distributes about 11000 (average 
of 5 years) half of these are distributed to gardens connected with 
schools in Denmark. j 

In the year 1795 Holbøll started a kind of registration of the 
plants cultivated in the garden, a scheme that has been followed since 
then and which will enable us to tell the history of each plant in the 
garden. 

The garden is still what it has been from the beginning a scientific 
garden and in accordance herewith the greater part of it is reserved 
to the students in the forenoon. On the other hand the garden must 
also be regarded as a promenade park, and after 1 oclock the public is 
allowed to pass all over the garden. Figures tell that about 1,400,000 
persons pass the gates during the year; although several of these only 
hasten through the garden to and from their working place, surely 
the garden is by many people regarded as a blessing for this part of 
the town. 

The ,,biological garden" where the plants have been arranged after 
characters and qualities, is especially destined to serve public instruc- 
tion. Just to the same purpose the palm-house is opened for the public 
each day (1—5) in the summertime. Special collections have during 
the years been exhibited for the public, thus for instance in 1915 a 
collection of Begonias was to be seen here, and in 1918 a collection 
of Commelinace's and Årace'es was arranged in the hothouses. 


8 


The garden has during its existence been the seat for experiments 
planned out by the botanists or by the garden-curators. Thus curator 
Mørch tried to feed silkworms with substitutes for the leaves of 
Morus alba and found, that Ulmus americana gave the best result. 
Curator Weilbach planned out experiments with heavy glass for 
greenhouses and with air circulation in greenhouses in the purpose 
to give the plants more natural conditions. Dr. Samsø Lund and 
Kiærskou have carried out investigations upon the cabbage species. 
In the latter years professor W. Johannsen, C. Raunkiær and 
C. H. Ostenfeld have planned different experiments. 

Although we here speak of the garden itself, a few notes about 
the institutions connected with it, will be of some value. The museum 
embraces collections of plants in alcohol, samples of wood and seeds 
and first of all the herbaåriums. These containing c. 475,000 sheets 
consist of 3 main divisions namely the universalherbarium, the 
danish and the arctic herbarium. The library to which admittance 
can be had 3 times the week, contains c. 20.000 volumes. 


SORØ ARBORETUM 


About the expiration of the past century an arrangement took 
place between the Sorø forests and the botanical garden of Copen- 
hagen in accordance with which the garden should rear trees and 
shrubs to be planted in the Sorø forests on grounds that could be 
spared. Since 1903 different conifers and a large collection of Cratæ- 
gus have been planted near ,,Filosofgangen" while a selection of Acer 
has been put down in ,,Feldskov"”. 


THESBOTANICAF GARDEN OFÆKIE EB) 


The university of Kiel erected in 1665 by the duke Christian 
Albrecht, received in 1669 from this prince a botanical garden 
and the planning out of this was intrusted to the medicinal professor 
Johan Daniel Major. The work he had undertaken lasted about 
4 years full of trouble and vexations for the poor Major. In 1773 
another medicinal professor Nikolaus Pechlin had to overtake 
the directorship. The garden soon declined and it seems that it did 
not exist after the year 1684. 

Å new hortus medicus was shortly after 1720 erected near the 
university-building. The apothecary Christiani pledged himself 
against the payment of 10 Rdl. the year to cultivate the ground and 
grow different medicinal plants and furnish the professors with mate- 
rial for their lectures on botany. 

In 1802 a new hospital with surrouling garden was erected in the 


") Until 1864 Holstein was united with the danish kingdom. 


2) 


town build by professor Georg H. Weber from voluntary contri- 
butions. This garden was in 1807 put under the guidance of his son 
Friederic Weber. The curator Peter Håhle was appointed from 
1806 until 1831. Twice the garden was enlarged and had in 1822 an 
area of about 2/, hekt. (4%/, Td.) The greenhouses were built partly 
at the start in 1802 partly in 1816 and 1820. The collections were 
rather large, in 1822 about 6000 species. Ferdinand Nolte 
succeeded Weber as director of the garden and about 1855 C. Dahle 
was curator there. 


ROYAL BOTANICAL GARDENS IN DENMARK 


In 1606 the king Christian IV. bought a number of gardens 
situated outside the town and here in the years 1610—24 he erected 
the famous castle Rosenborg. The garden to this castle was rather 
spacious and the king found it desirable here to make a botanical 
garden, and in consequence herewith such a one was laid out and 
in the year 1638 put under the guidance of dr. O. Sperling. This 
botanist was a very industrious and energetic man and soon brought 
together a large collection of plants; his ,,Hortus Christianæus" 
of 1642 that contains 1519 names of plants bears witness of this, and 
in this respect his garden surpassed (he garden of the university. The 
curator of this part of the Rosenborg garden David Konig has not 
been capable to manage the large collection, and Sperling expresses 
his discontent herewith. A selection of plants mentioned in ,, Hortus 
Christianæus" is given on page 337; the names mentioned in brackets 
give the names now used. Several of these plants were to be grown 
under glass and we also know that Sperling has had greenhouses for 
this purpose. In 1644 Sperling feli into royal disgrace and as his 
successor did not påy any special interest in the botanical garden, 
this soon decayed. i 

HnedericavøÆkKkmskor Denmarks hadstherntentonktolereetka 
splendid quarter of the town and in connection herewith he founded 
1752 a hospital and a botanical garden. The latter consisted of two 
parts measuring 11.106 (| al. and 24.539 O al. divided by a broad 
street. The famous botanist G. C. Oeder was designed to govern 
this garden and the first curator was Kristian Hansen. About 
1760 the smaller garden was finished; the greater one never came 
to fulfilment although a wall had been built all around the ground. 
Greenhouses have been in the garden and in 1763 it was opened to 
the public. As it turned out to be too expensive for the king to keep 
this garden the smaller piece of ground was presended to the uni- 
versity. (See further about this garden on page 1). 


10 


THE FORESTIAL-BOTANICAL GARDEN IN CHARLOTTENLUND 
AND THE GARDEN AT THE AGRICULTURAL COLLEGE 


Before the year 1863 no instructions were given at the agricul- 
tural college to horticulturists and foresters. The former received 
instruction where they were apprenticed, the latter took lessons at 
the polytechnicum and were examined by a commission with Jonas 
Collin in the chair. This commission proposed in the year 1838 to 
lay out a smaller arboretum close to the. metropolis and in conse- 
quence herewith an area of 8 Td.…. (c. 11 acres —< 4 hekt.) in the 
southwestern corner of Charlottenlund forest "was destined for the 
named purpose. This garden was rearranged in 1863 at which epoque 
the forestry-studium was transferred to the agricultural college, and 
since the year 1869 the forest-garden has been put under the same 
college. 

The old veterinary-college had a small garden where medicinal 
herbs were grown, but-this garden has been of no significance. In 
1858 when the veterinary college was removed and enlarged to an 
agricultural college, a garden was laid out around the buildings. This 
garden was planned out by A. Bentzien, while the selection of 
plants was made by professor Joh. Lange assisted by R. Rothe 
and Aug. Weilbach. In 1863 when the course for horticulturists 
was put under the institute, the garden was enlarged with an orchard. 

Owing to the frowns of fortune the two gardens here mentioned 
have been limited in area; thus in 1895 the forest garden in Char- 
lottenlund had to make over ”/, Td. area to the railw ays and in 1892 
the agricultural college was enlarged, and the garden must yield the 
necessary area 

Since 1869 the two gardens have been under the same guidance; 
this charge was firstly put in the hands of the lecturers of botany 
and horticulture, later on under the latter alone and in 1892 the gar- 
dener of the institution took over the curatorship; as curator Bruun 
in 1902 became a professor he was at the same time designed to 
be director (eforus) of the garden. 


BOTANICAL GARDENS AT GYMNASIUMS AND MUNICIPAL 
SCHOOES 


AtSorø-academy a garden existed in the time of Buser (about 
1625), 1822 a new garden ,,Plantagehaven" was laid out close to the 
curator's lodge. In 1837 not less than 1422 species were cultivated in 
these grounds. 

At Herlufsholm an area of 3, Td. was laid out im the year 
1890 to a pinetum, designed by Tra ustedt. Here more than 100 
species and varieties of conifers were grown about 1896. The soil is 


11 


rather bad, but nevertheless the collections are of great interest. To 
the college also belongs a students” garden containing 550 species. 

In Aarhus a municipal-garden was established in 1905. Its pur- 
pose is to furnish the schools of the town with material for the 
instruction in botany. 300 different species are cultivated and about 
14000 shares of plant-material are distributed in the year. 


PRIVATE BOTANICAL GARDENS 


Different persons have shown so much interest in botany that they 
have formed their gardens partly as botanical collections. Klavs 
Urne m"Lund (the "16th century) and Johann Ludvig Hol 
stein at Ledreborg (the 18th century) have to be mentioned in this 
respect, and Lotze, apothecary in Odense had in the middle of the 
19th century large collections. An arboretum at Aalholm was star- 
ted in the end of the 18th century and was partly renewed about 
1830. In our days interesting arboretums are to be seen at Knuthen- 
borg and Hardenberg. Private collections of alpine plants and 
shrubs are kept by mag. Lorenzen, Holte and dr. F. Børgesen, 
Hellebæk. 


REE ER GE UREANDF BIOLOGYFOFFARGEIG 
ELOWERENGEPEANTS 


14. 


EFB æres. 


Morten P. Porsild. 


19210) 


Reprinted from »MEDDELELSER OM GRONLAND« Vol. XXXVII. 


Arbejder fra den botaniske Have i København. Nr. Y 


Hitherto, the following papers have been published in "Med- 


delelser om Grønland”: 


ss 


VolssoemeviE 


. Ericineæ (Ericaceæ, Pirolaceæ). 


1. Morphology and Biology. By EuG. WarMING  p.1—71. 
2. The Biological Anatomy of the Leaves and of 
the Stems. By HENNING EILER PETERSEN... p.73—138. 


. Diapensiaceæ. Diapensia lapponica L. By HENNING 


FEER PETERSEN ne FR SR BESS MEE p. 139—154. 
Empetraceæ. Empetrum nigrum L. By A. MENTZ p. 155—167. 


. Saxifragaceæ. 


1. Morphology and Biology. By EuG. WARMING p. 169—236. 
2. The biological Leaf-anatomy of the Arctic spe- 


cies of Saxifraga. By OLAF GALLØE....... p. 237—294. 
5. Hippurid aceæ, Halorrhagidaceæ and Callitrichaceæ. 

By ENE LE SED EEN er p. 295—332. 
6. Ranunculaceæ. By KNUD JESSEN.........…. p. 333—440. 
7. Lentibulariaceæ (Pinguicula). By Fr. HEIDE... p. 441—481. 

Volker 
8. Rosaceæ. By KNUD JESSEN ....…... ERR p. 1—126. 
9. Cornaceæ. By CARSTEN OLSEN..........….... p. 127—150. 
10. Caprifoliaceæ. Linnæa borealis. By O. HAGERUP p. 151—164. 
11. Primulaceæ. By Fr. J. MATHIESEN ......….…. p. 165—220. 
12. A List of Arctic Caryophyllaceæ, with some 


synonyms by CE Os renere DE p. 221—227. 


3. Caryophyllaceæ. By Euc. WARMING......... p. 229—342. 





LIBRARY 


NE VK 
97 =- 
HAL sr an 


f the order Liliales several species, belonging to differ- 
ent families, advance to the arctic limit of forest 
occasionally even crossing it, not attaining, however, a wider 
distribution in the arctic territory where they do not belong. 
We find also that they in the alpine regions do not, or only 
occasionally, cross the limit of forest, cf. remarks in SCHROE- 
TER: Pflanzenleben p. 3534—5. 

The principal of those species are the following: 

1) Veratrum album L. and the closely related V. viride 
Ait. extend respectively in the river-valleys of Northern 
Eurasia and Northern America to the coasts of the Arctic sea. 

2) Streptopus amplexifolius (L.) D.C. is an E. Asiatic 
—N. American forest plant; the fruit is a berry, and it has 
probably, through the agency of birds, been brought to the 
subarctic copses of the southernmost part of Greenland. 

3) Lloydia serotina (L.) Sweet — closely related to 
Gagea — advances on both sides of Bering's strait farther 
into arctic territory than any of the others. Thus it is found 
at Cape Lisburn and extends even as far as the purely arctic 
New Siberian Islands. 

4) Allium schoenoprasum L. (including Å. sibirieum L.) 
is found in the island of Kolgujew, in the river-valleys of 
N. Asia up to the Arctic sea, and on the north coast of 
Alaska. 


346 MortEN P. PoRrSsILD. 


5) Iris sibirica L. (and related species?) extend to the 
coast on both sides of Bering's strait. 

As to the structure and biology of the majority of the 
above-mentioned species the reader is referred to KIRCHNER, 
Loew & ScHROETER's: "Lebensgeschichte” where — on 
Alpine material — these questions have recently been treated. 





Widely distributed in the arctic territory are only 


the two following species of the genus 


Tofieldia: 


1) T. palustris Huns., circumpolar, common from the 
subarctic territory far into the arctic, probably without 
reaching the purely high-arctic regions, however. 

2) T. coccinea Ricxa., N. Asiatic, N. American, found 
besides in the northernmost two-thirds of Greenland, every- 
where rare or perhaps overlooked; numerous of its Asiatic 
occurrences are only known from previous erroneous deter- 
minations, corrected by OstENFELD (Fl. Arctica p. 32). 

A third species: 7. calyculata WAHLENB. is distributed 
throughout nearly all the mountain regions of Central Europe, 
whence it often descends into the lowland, whilst it only in 
a few places crosses the Alpine limit of forest: the variety 
glacialis REICHENB. 

The structure and biology of 7'. calyculata is extremely 
well-known, and the same applies partly to T'. palustris. An 
account of this, based upon Alpine material, is found in 
Loew & KIRCHNER's: "Lebensgeschichte” p. 229 ff., a copious 
literature being quoted here, too. Cp. also the description of 
the closely related Narthecium ossifragum Hvuns. ibid. p. 244 
fi., and RAUNKIÆR p. 138ff. The following observations, 


adds 


Liliales. 347 


having all been carried out upon arctic, living and alcohol 
material, chiefly from Greenland, will therefore serve mostly 
as a complement, especially with regard to the least known of 


the species: 7'. coccinea. 





Ocecurrence in nature. Fofieldia palustris and T. coc- 
cinea grow on moist spots among heath- and bog vegetation, 
more rarely on fresh plantless moraines. They occur most 
frequently here as firm cushions or cakes which are held 
together by the vigorous secondary roots (Fig. 1, A). They 
attain their highest development when growing among bog- 
mosses. Here the leaves become longest, and the floral shoot 
tallest, but on the other hand, the stalks are able to creep 
better and more widely, and consequently we do not get 
here as large and firm cakes as otherwise. The habitats are 
covered by snow during winter. 

The shoots are short-jointed, flat, frequently erect, 
more rarely, through want of space, obliquely ascending and, 
in vigorous specimens, each shoot bears from 6—8 leaves. 
During several years the shoot remains on a purely vegetative 
stage until finally ending in ån inflorescence. Some of them 
remain on the vegetative stage. Structure of shoot cp. Fig. 1, 
A—D. 

The leaves are all uniform, apart from a slight difference 
in length, the lowermost being the smallest, evergreen, two- 
rowed ensiform, equitant with a long sheath. Below the 
green leaves some withered ones (often black-spotted by fungi) 
are found, and at the base setaceous, vascular strands are 
seated originating from older marcescent leaves. Scale-leaves 
are not found. The lateral shoots are formed in the uppermost 
axils of the leaves, normally only one or two are developed. 


348 





sål 


B. 


MortEN P. PORSILD. 


Foss 
A, B, C. Structure of shoot in 7”. palustris (Finse, Norway). 


Habit drawing of two flowering shoots which only cohere by the 
aid of the roots. b, the scape with its sheath-formed bract. 

The main-shoot, I, has two withered leaves (1—2) and six fresh 
(3—8). In the axil of 2 a shoot: II, is to be seen, and in the axil of 
3 another one II, of which 4 leaves are visible. Magnified, detailed 
drawing of it in the following (C). 

Shoot with its subtending leaf: f, and with 3 leaves marked I, 2, 
3,4. 1 and 2 turn their back to the mother-axis. 

T. coccinea. Foliage shoot with withered remains of leaves, from 
unfavourable habitat. (Dove Bay, E. Greenland; leg. A. Lundager, 
le 1631908) 

Germination in T'. calyculata. (Drawn by E.W.) 


349 


Liliales. 


In T. coccinea there is as a rule 9(—11) vascular bundles 
(Fig. 2), in T.palustris 11(—13). In the first-mentioned the 
leaves are often more flaccid (Fig. 1, D), frequently partly 
decumbent, especially in exposed places, whilst they, in the 
latter, always are rigidly erect. 

The structure of the leaf is in all essentials identical 
in the two species. The sheath is constructed like a normal 
leaf, above this the leaf is isolateral, has an equal right and 
left side, an upper and lower margin (Fig. 3). The epider- 
mis consists, except on the upper 
(inner-) side of the sheath, of ra- 
ther small cells, the walls of which 
are slightly undulating, highly 
thickened, porous and covered 
by a well-developed cuticle which 





also spreads to the guard-cells of 
the stomata (Fig. 3 B,.D). The 


Nordre 
Stromfjord, Greenland. 


Fig. 2. T, coccinea. 


stomaåta are arranged longitu- 
dinally, their slit is quite diminu- 
the 
beneath them is quite small. The 


tive, intercellular  cavity 
base of the sheath consists of 
hyaline parenchyma ; alittle higher 
up, on the morphological lower 
side, we find the chlorophyll 


tissue consisting of several lay- 


a—e show sections in various 


height through a foliage 
leaf. The xylem of the vas- 
cular bundles is dark. The 
dotted line shows the bound- 
ary between the chlorophyll 
tissue and the aqueous tis- 
sue; a, showing the basal 
part of the sheath, has no 
chlorophyl. 
(Drawn! by MB SES) 


ers of cells, whilst the upper side is constantly non-chlorophyl- 
lose. Above the sheath thechlorophyll tissueis homogeneously 
developed on both sides, the non-chlorophyllose part occupy- 
ing the centre of the leaf, extending right to the tip of the 
leaf (Fig. 2, 3, A, B). In contradistinction to Narthecitum 
(vide RAUNKIÆR) no air-chambers are found in the chloro- 


phyll layer on transverse sections. 


DS 
(95) 


XXXVII. 


350 MortEN P. PoRrSsiILD. 


The cells in the chlorophyll tissue are parallelepipedic 
and frequently no intercellular spaces are seen in transverse 
sections; in longitudinal sections, however, quite small ones 
are to be seen. The cells of the chlorophyll tissue contain 
chlorophyll grains filled with starch. In the isolateral part 
the vascular bundles approach by twos to each other, 
and the uppermost one is entirely connated of two (Fig. 2 6, 
3 A). In transverse sections the large, crescent-shaped 
coverings of bast are to be seen. 





am 
BASS MÅ SS SSSUNINT, SENDER 
sa DÅ AGNES ANES ang 





Fig. 3. Anatomy of leaf. 

A,B,C. T. palustris; from Jakobshavn, W. Greenland. 

A. Transverse section of a foliage leaf above the equitant part. The 
hatched am is the chlorophyll tissue; the epidermis and the 
central aqueous tissue are light. 

B. A part of the same section, more highly magnified; ep. Epidermis 
with thick cuticle and a stoma with very minute air cavity. Chloro- 
phyll tissue almost without any intercellular spaces, the cells filled 
with starch; m aqueous tissue. 

C.. Epidermis with two stomata, longitudinally placed. The walls of 
the cells are highly thickened, slightly undulating. 

D. Transverse section of a leaf of 7'. coccinea (Danmarks Ø, E. Green- 
land leg. N. Hartz). (Drawn by E. W.) 


The scape of inflorescence is as a rule leaf-less, yet 
in vigorous specimens 1—2 leaves may be found, the upper- 
most one at least having the character of a bract (Fig. 1). In 
T. coccinea 2—3 stem-leaves are nearly always present 
(Fig. 4 4), the uppermost one having, in this case, less of 
the character of a bract. 


Liliales. SØDE 


The inflorescence in 7. palustris is nearly always a 
giobular head (Fig. 1 4), more rarely a subcylindric spike. 
In T. coccinea the elongated spike is the normal form, the 
globular the rarer one (Fig. 4 A, B), but it varies according 
to the quality of the habitat. In 7. coccinea, moreover, 





Fig. 4. T. coccinea. Nordre Stromfjord, Greenland. 

A. Habit-drawing of a weakly flowering specimen growingin Sphagnum. 

B. Inflorescence of a more richly flowering specimen. 

C. Single flower, shortly before the hermaphrodite stage. 

D, E, F. Almost ripe fruits in various positions; D and E the same 
specimen; note the asymmetry. 

G. Transverse section of the ovary. 

H. Uppermost part of the inflorescence, showing the downwards- 
directed carpotropic curvature, characteristic of the species. 
(Drawn by Thorbjørn Porsild). 


single isolated flowers are frequently to be found below the 
spike, just as in 7, calyculata and in other species of the genus. 


23+ 


352 Morten P. PorsiILD. 


The flowering season is the middle of the summer, 
consequently the species do not belong to the earliest 
flowering. In an average year they will, on the latitude 
of Disko, be out from the end of July, and then flowering 
specimens are to be met with all through the rest of the sum- 
mer, because the floral shoots from the same tuft are not 
all of them developed simultaneously. 

Under each flower a small 3-dentated bractlet is 
present which often envelops the peduncle. In 7. palustris 





Fig. 5. T, palustris (Norway.,) 
A. Part of inflorescence; note the small, 3-dentated bractlet. 
B. Flower åt the beginning of the hermaphrodite stage: the stamens 
arise from the spoon-formed perigonial leaves. 
Stamen in a perigonal leaf when the flower bursts. 
Anther with open valves. 
Carpels in the stage of pollination. 
A single carpel, more highly magnified, showing pollen-grains 
on the stigma. (Drawn by E.W.) 


Bk SES 


it is always shorter than the peduncle (Fig. 5 4), in T. cocci- 
nea just as long as this or longer (ABROMEIT). In the latter 
a fairly well-developed "calyculus” is as a rule present, a 
bi-symmetrical whorl of 3 small leaf-formations of which 
one is turned outwards, the other two transversal (Fig. 6). 
The morphological interpretation is uncertain. In 7. palu- 
stris it is normally absent, but may also, according to 
ÅBROMEIT, occur occasionally. 


Liliales. 353 


Diagram. EiIcHLer's diagram of 7. calyculata shows, 
besides the zygomorphy in the calyculus, a slight irregularity 
in the mutual size of the carpels. This also applies to the arctic 
species, especially to 7'. coccinea (Fig. 6), where it appears 
to be even more strongly developed than in E1cxaLeRr's 
diagram. This difference is undoubtedly connected with the 
carpotropic movements (see below). 

In T. palustris the perianth is purely white, or with 
a greenish rather than a yellowish tinge, in 7. coccinea 
it is purely white on the inner side, deep purplish on the 
exterior, especially along the median line 
of the leaves. Also the upper part of the 
scape and the carpels are of a beautiful 
purple-red colour. This bi-colouring is in 





live specimens very conspicuous, and when 
the species grow together they are easily Fig. 6. 
distinguished from each other by the colour Floral diagram 
of T'. coccinca. 
Note the 
red pigment often disappears entirely, the asymmetry in 
the carpets. 


É (Drawn 
hence the numerous confoundings and erro- by M. P. P.) 


of the flowers alone. But in herbariums the 
flowers in both species becoming yellowish, 


neous determinations. 

The biology of the flowers (Fig. 5 B—E, 4.C) has 
been studied by H. MuLLEeRrR in the Alps on T'. calyculata 
which he found proterogynous, while 7'. palustris was almost 
homogamous. In spite of a greater abundance of honey- 
secretion in the latter he found a greater number of insects 
visiting the former. Greenlandic flowers of 7. palustris and 
T. coceinea were almost homogamous with a slight indication 
of proterogyny. Visiting insects I have never seen there. In 
T. palustris MULLEerR draws the stamens as freely projecting 
in the flower, in Greenlandic living material the filaments 
were curved downwards into the cavity of the perigonial 


354 MortEN P. POoRSsILD. 


leaf (Fig. 5 B, b, 4 C), this was especially prominent in the case 
of T. coccinea. During calm weather the cavity of the peri- 
gonial leaf is filled with pollen. When the flower has been open 
for some time the filaments, as well as the perigonial leaves, 
start curving upwards, and then the introrse anthera cannot 
avoid touching the stigmas. Thus cross-pollination is possible, 
but self-pollination the rule. 





Fig. 7. T'. palustris (Disko, Greenland). 


A. Hibernated inflorescence with upwards-directed capsules, cha- 
racteristic of the species. 

B. Single, complete fruit of the same. 

C.. Almost ripe inflorescence gathered late in autumn. 

D. Fruit, almost ripe, but not yetopen. (The perigonial leaves removed). 


Note the almost perfect symmetry, as against 7'. coccinea. 

E. Hibernated capsule with walls partly fallen off, typical for the 
species. Some seeds are still seated on the placentas. 

F and G. A "normally” opened fruit (a rare case in the species). 7. in 
moist condition. G. dry, seen from above. 

Haseeds (Drawn bys FPS) 


Afterthe pollination the inflorescence, which hitherto 
has been slightly nodding, straightens itself up, and the 


Liliales. 355 


scape becomes rigid. The head or spike is stretched a little 
(Fig. 7 A). The perigone withers, but persists for a long time 
round the carpels (Fig. 7 C). In 7. palustris the capsule is 
ovate, often a little brownish in colour and bent upwards 
(assm 7. calyculata). (Fig. 7 A,.C), whilst it in 7. eoccinea is 
shorter, subglobose and bent downwards (Fig. 4 H). The 
lowest fruits in the inflorescence bend right downwards 
parallel with the scape, the uppermost obliquely downwards 
or the top one only horizontally. This carpotropic movement 
begins immediately and is continued during the ripening of 
the fruit. 

Theripeningofthe fruit is very late. Even on Disko, 
that lies far to the south of the Polar limit of the species, it 
is very difficult, even after favourable summers to find ripe 
fruits in the autumn, and if the unripe fruits are brought 
home the ripening is not continued. Exceptionally I have, 
however, found single, quite ripe fruits of 7'. palustris late in 
the autumn; the dehiscing is septicidal, forming a slight open- 
ing so that the seeds might be shaken out (Fig. 7 G). I have 
had no chance of seeing 7'. coccinea late in autumn. The 
majority of fruits in 7'. palustris, and probably also in the 
other, do not thus attain to ripen their fruits before the snow 
comes. Consequently the ripening must take place under 
the snow. I have looked through a large material of hibern- 
ated fruits of both species and only succeeded in finding a 
single ”"normally” opened fruit. Instead, the very thin cap- 
sule-walls had gone to pieces, and the seeds had got out 
through the holes which had come into existence in that 
way. Often the capsules may be quite ”skeletonized” so that 
only the backs of the carpels are left (Fig. 7 E). 

SERNANDER (,,S9pridningsbiologi” pag. 354) has gathered 
T. palustris with seeds in the capsules in spring at the time 
when thesnow was beginning to melt, and he therefore includes 


356 MortEN P. PorsiLDn. 


them among the winter-standers. To å certain extent this 
is right, as it does not attain to ripen its fruits till the snow 
comes, and consequently may happen to spread its seeds on 
the snow. But the white, paper-thin capsules of TFofielda are 
to a less extent adapted to this than for instance, the Luzula 
species, the brown capsules of which are saturated with a sub- 
stance which makes them proof against the various attacks 
of the first winter and preserves their hygroscopicity. 

What brings about the skeletonizing of the Tofieldia 
capsules, I do not know. Ifit had been effected by mechanical 
wear of drifting snow in the spring-time there would always 
be found some which had been lying in sheltered places and 
remained intact. The apertures are irregular, as if the thin 
parchment-like capsular membranes had, through iterated 
freezing or exsiccation, become so fragile that even the slight- 
est touch was enough to cause the apertures. 

The production of seeds is no doubt abundant in 
both species, judging by the number of well-developed ovules 
and by the occurrence of the species in nature. Vegetative 
propagation is practically excluded, because even the strong- 
est stream issuing from melting snow would hardly be able 
to break asunder the cakes which are made up of the strong 
roots felted together. Only in loose growing mosses a libera- 
tion of the lateral shoots takes place when the rhizome dies 
away at the back. 

The seeds (Fig. 7 H) in T7. palustris are small, yellow, 
a little triangular and slightly curved, with few wrinkles 
faintly marked on the testa. They weigh 0.03 mg. In T. 
coccinea, of which I have gathered but a few seeds in hibern- 
ated capsules, they were of about the same size and appear- 
ance. 

Germination. The first stages have been seen by 
KLEBSs (quoted in "Lebensgeschichte”) and later on by War- 


Liliales. 357 


MING in T. calyculata (Fig. 1 E). After having emerged from 
the testa the cotyledon is geniculate bent, and at a very early 
stage a whorl of roothairs is produced at the base of the 
young root. The later stages are not known, but they might 
agree with those in Narthecium described by BucHENau. 





Fig. 8. 
A. Seedling of T. palustris at the end of the second period of vegetation. 
B. 3—4 years old seedling of the same species. Both from Disko, 
Greenland. 
C. Retarded inflorescence, extricated from an almost withered spec- 
imen of T. coccinea; g. small vegetative lateral shoot. Nordre 


Stromfjord, Greenland. (Drawn by Th. P.) 


Seedlings (Fig. 8) are easily found in nature, but it is 
very difficult to find the very first stages because they are 
so small. The earliest formed foliage leaves wither rapidly, 
and the young plants pass through a period of strengthening, 
which lasts for many years, before they attain to flowering. 








BIBLIOGRAPHY. 


ÅBROMEIT, J.: Botanische Ergebnisse der von der Gesellschaft f. Erd- 
kunde.... ausgesandten Gronlandsexpedition. Phanerogamen. 
Bibliotheca Botanica 42. 1899, 

Loew, E. und KircHaNErR, O.: Tofieldia im KircaNER, LOEwW u. 
SCHROETER: Lebensgeschichte der Blitenpflanzen Mitteleuropas 
Bjert trane EEN Een TS worn extensive bib- 
liography is found. 

MiLLER, H.: Alpenblumen, ihre Befruchtung durch Insekten und ihre 
Anpassungen an dieselben. Leipzig 1882, 

RAUNKIÆR, C.: De danske Blomsterplanters Naturhistorie I. Kjøben- 
havn 1895—99. 

SCHROETER, C.: Das Pflanzenleben der Alpen. Zurich 1908. 


SERNANDER, R.: Den skandinaviska florans spridningsbiologi. Upsala 
1901. 


Besides, I have made freely use of notes and drawings by Pro- 
fessor E. WARMING. 








TYP, BIANCO LUNO. KBHVN. 





| ORCHIDACEAE 
"TENIANAE YUNNANENSES 


VON 


FR. KRANZLIN 


WOLFENBUTTEL 


Sonderabdruck aus Fedde, Repertorium XVII (1921), pp. $99—112 


Arbejder fra den botaniske Have i Kobenhavn. Nr. 95 





Ren eo VEN, ÆRE Å — 


. Fr. Krånzlin: Orchidaceae Ténianae Yunnanenses. 99 


XXXI. Orchidaceae Ténianae Yunnanenses ”. 
Von Fr. Krånzlin (Wolfenbiittel). 
Arbejder fra den botaniske Have'i København. Nr. 95. 


Herr Dr. C. Christensen in Kopenhagen hatte die Freundlichkeit, 
mir die Orchidaceen anzuvertrauen, welche Herr Siméon Tén gesammelt 
hat. Es sind im ganzen 41 Arten, von denen 13 neu sind; leider sind wieder 
6 davon Habenarien. Die im vorigen Jahre erschienene Arbeit des 
Herrn Dr. R. Schlechter tiber die Orchideenflora des japanisch-chinesischen 
Florengebietes, welche in Feddes Repertorium Beiheft IV veråffentlicht 
ist, war in erster Linie zu bertcksichtigen, sodann die Flora of Hongkong 
and Kwantung von Dunn and Tutcher. Das erstgenannte Werk mubte 
ich wegen einer ganzen Anzahl Meinungsverschiedenheiten stårker beruck- 
sichtigen. Der Verf. hat geglaubt, in Arbeiten, die ich vor 20—25 Jahren 
geschrieben habe und denen zum groBen Teil sehr dirftiges Material 
zugrunde lag, eine Anzahl Fehler aufdecken zu miissen, was an und fur . 
sich berechtigt, låblich und fur die Wissenschaft nur fårderlich ist. Ob- 
gleich ich Kontroversen dieser Art grundsåtzlich aus dem Wege gehe, 





1) Alle hier aufgezåhlten Exemplare sind Originale des Bot. Mus. Kopen- 
hagen. Fedde. 


fx 


100 Fr. Krånzlin. 


hat die hier vorliegende Arbeit mich doch dfter, als mir lieb war, genåtigt, 
meine friiher vorgetragene Ansicht zu verteidigen, was mir, wie ich hoffe, 
gelungen ist, ohne den Boden der Sachlichkeit zu verla«sen. 

1. Cypripedilum yunnanense Franchet in Journ. de Botan. VIII 
(1894) 231. — Yunnan. In silvis La pa ho, Pe yen tsin. (Siméon Tén 
sine no. !) 

2. Spiranthes sinensis (Pers.) Ames Orch. II (1908) 53. — Yun nan. 
Pe yen tsin. (Siméon Tén no. 1256!) g 

,Nomen vernac. Long pao tschou, quod significat. (Long) Draco 
(pao) amplectens (tschou) columnam." 

3. Epipogum aphyllum Swartz, Summa veget. Scand. (1814) 32. =— 
Yunnan. Ta( ?) tsin teon. Prope unum fontem. (Siméon Tén no. 1332) 


4. Neottia camtschatea Rchb. f. Fl. Germ. XIII. XIV 146 t. 478. — 


Yun nan. Ex silvis. Ta tsin teon. (Siméon Tén no. 1331!) 
5. Enipactis Mairei Schlechter in Fedde, Repert. Beiheft 4 (1919) 
55-et 148. — Yun-nan."Pe yen tsin, im 'silvis. "(Simeon Ten sine no): 
6. Epipactis setschuanica Ames et Schlechter in Fedde, Repert. 
Beiheft 4 (1919) 56 et 149. — Yun nan. Pe tsao lin, in silvis. (Siméon 
Tén no. 1383!) 


Die Bliten sind etwas kleiner als in der Diagnose angegeben und ihrer 


Farbe nach sind sie ,,flavi'", wåbrend die der Wilsonsehen Exemplare 
,brownish''" sein sollen, sonst konnte ich keinen Unterschied finden. 

7. Epipactis discolor Krånzl., spec. nov. — Radices copiosae, tenues, 
satis longae. Caulis ad 30 cm altus, tenuis, flaccidus, fractiflexus, basi 
cataphyllis ochreatis, brevi-acutatis, ringentibus vestitus, excepta in- 
florescentia ubique glaber. Folia subdisticha, infimum fere orbiculare, 
sequentia oblonga ;v. ovato-oblonga v. oblongo-lanceolata, apiculata 
v. acuta v. (suprema). acuminata, in bracteas decrescentia, margine et 
in venis subtus ciliolata, superne opaca, subtus pallidiora (unde nomen !), 
2,5 ad 7 cm longa, 2 ad 3 cm lata. Inflorescentia racemosa, obscure se- 
cundiflora, vix 10 cm longa, utplurimum 10-flora, rhachis glandulis durius- 
culis obsita, scabra, bracteae lanceolatae, acuminatae, infimae 2 cm longae, 
flores duplo superantes, superiores multo minores, illis aequilongae, ovaria 
fere sessilia, minute scabriuscula. Sepala late ovata, acuta, dorso carinata, 
carinis minute serrulatis, ceterum glaberrima, 7 mm longa, basi 2 mm 
lata. Petala textura teneriore, ovata, acuta, non carinata, glabra, aequi- 
longa et-lata. Labelli hypochilium profunde naviculare, in fundo pa- 
pillosum et carina mediana praeditum, epichilium subquadratum, antice 
retusum, basi callis 2 semiglobosis haud confluentibus praeditum, venis 
3 percursum. Flores ,,rubri". — Yun nan. Peyen tsin (an recte ?)'in 
silvis. Flores Junio. (Siméon Ten sine no.!) — Adsunt specimina 
4 satis bene exsiccata. 

Die Pflanze åhnelt entfernt einem schwåchlichen Exemplar unserer 
Epipactis rubiginosa, alle (4) Exemplare hatten das gleiche Aussehen. 
Von neuerdings publizierten chinesischen Årten ist zweifellos Ep. yunna- 
nensis Schlechter die zunåchststehende und es ist nicht absolut aus- 


ut ark je sl ERE: 


Orchidaceae Ténianae Yunnanenses. 101 


geschlossen, daB diese Art hier doch eines Tages in jene aufgehen wird, 
zunåchst sind jedoch folgende, in der Schlechtersvhen Diagnose nicht 
erwåhnte Abweichungen festzustellen: Erstens schlaffe, deutlich im 
Zickzack gebogene Stengel; zweitens die verschiedene Fårbung der beiden. 
Seiten der Blåtter, drittens die Wimperung am Rande wie unterseits 
auf den Hauptnerven, Meérkmale, die der Autor wohl erwåhnt håtte, 
wenn sie bei Ep. yunnanensis vorkåmen und die keineswegs fur Epipactis 
selbstverståndlich sind. Die Åhre ist armbliitig und sehr undeutlich ein- 
seitwendig; die Bluten sind noch kleiner, nur 7 mm lang, die Sepalen 
haben auf dem Riicken einen kurz gezåhnelten Kiel, sonst sind sie glatt, 
die Farbe ist dasselbe trube Rotbraun wie bei unserer Ep. rubiginosa. 
Das Epichilium der Lippe hat am Grunde zwei sehr-hervorragende Buckel, 
welche nicht miteinander verschmelzen, und drei ziemlich undeutliche 
Nerven; leider war dieser Teil bei allen Bliten nicht gut zu sehen, die 
Exemplare waren schon uber ihre Hohe etwas hinaus. Schliehlich bluht 
Ep. discolor im Juni und Ep. yunnanensis- im August. Die anderen 
Schlechterschen Arten kommen nicht in Betracht. — Leider hat der 
Sammler nieht gesagt, ob die Bluten Vanillinduft haben, der unsere Ep. 
rubiginosa bekanntlich auszeichnet; auch beim Ep. yunnannensis scheint 
er zu fehlen. 

8. Orchis Chusua D. Don Prodr. Fl. Nepal. (1823) 23. — Yun nan. 
Ex vertice montis Pe trao lin. (Siméon Tén no. 1377!) 

9. Orehis puberula King et Pantl. in Ann. Roy. Bot. Gard. Calcutta 
VIII (1898) 304 t.403.—Yun nan. Ta tsin teou. (Siméon Tén no. 1326! 
Fl. 28 Septembri. 

Die Bliten sollen nach Angabe des Sammlers ,,paulo flavi' sein, 
wie er in seinem etwas wunderlichen Latein sagt, gemeint ist wohl ,,pallide 
flavi', King und Pantling sagen: the flowers are white", bilden sie aber 
ganz hellrosa ab mit einzelnen roten Punkten auf dem Labellum. Im 
ubrigen stimmen Exemplare, Text und Abbiidung so gut wie nur måglich. — 
Die Art ist in der Schlechterschen Zusammenstellung noch nicht erwåhnt. 

10. Anthogonium gracile Wall. ex Lindl., Cat. no. 7398 ex Lindl., 
Gen. et Spec. Orch. (1840) 426; non Lindl. — Anthogonium corydaloides 
Schlechter in Fedde, Repert. Beihefte Bd. IV (1912) 66 et 230... Yun nan. 
Konty. (Siméon Tén no. 1302!” Exeunte Augusto. 

Uber die Autorschaft dieses Namens lauten die Lesarten verschieden, 
geht man bis auf die Quellen zurick, so findet man, dak Lindley stets 
Wallich als Autor nennt, so in dem Introduction to the Natural System 
2. ed. (1835) 341, wo der Name zuerst vorkommt, wie in den Genera and 
Species und in Ubereinstimmung hiermit steht auch im Index Kewensis 
Wallich als Autor der Gattung sowohl wie der Art. Bentham hat in den 
Genera Plant. III, 515 in striktem Widerspruch hierzu Lindley zum Autor 
gemacht und ihm ist Hooker in der Flora of Brit. India III, 823 gefolgt, 
selbstverståndlich auch Grant, Orchids of Burmah S. 138 und schlieblich 
Schlechter in seinem ,,Orchideologiae Sino-Japonicae Prodromus'"' S. 66, 
was alles zusammen nichts an der oben festgestellten Tatsache åndert, 


102 Fr. Krånzlin. 


bæ= 


daB Lindley die Autorschaft von vornherein abgelehnt hat. Die neu É En 


aufgestellte Art A. corydaloides Schlechter ist, soweit sich aus den be- 
sonders hervorgehobenen Merkmalen ergibt, nichts weiter als die Stammart.… 


Schon Griffith, der die Pflanze in den Notulae III, 383 gut beschreibt, 


sagt geradezu: ,,flos fumarioideus purpureus. Der rechte Winkel, den 
das Ovarium mit dem Perigon bildet und der nach Schlechter eines der 


Merkmale seiner Art sein soll, findet sich auf Griffith's Abbildung in den 


Posthumous Papers III t. 345. Diese ist, wie die meisten Abbildungen 
jenes Werkes, roh in der Technik aber charakteristisch und um das andere 
,Merkmal'' zu erwåhnen — auf ein etwas tiefer geteiltes Labellum gleich 
eine neue Art aufzustellen, geht denn doch nicht an; das hieBe, die Haar- 
spalterei zum Prinzip 'erheben. Hooker war seinerzeit ganz im Recht, als 
er in Flora of Brit. India VI, 823, A.Griffithir Rchb. f. und die in Griffiths 
Werken beschriebene Art zu 4. gracile Wall. zog und die A. corydaloides 
Schlechter ist gleichfalls nichts anderes. Das Verbreitungsgebiet reicht 
ubrigens noch viel weiter, denn die Pflanze ist auch von Java bekannt 
(Lobb no. 349! in herb. Petropol.). Reichenbach hat in der Bonplandia II 
(1854) 90 (= Walp. Ann. VI, 471) ein A. Griffithir aufgestellt, zu dem er 
aber in den Annalen hinzufigt: ,,Monet ill. Lindley nil nisi ipsissimum 
A. gracile Wall.,” eine Bemerkung, der er nicht widersprach. 

11. Cynosorehis gracilis Krånzl., Orch. Gen. et Spec. I (1898) 488 et 
in. Engl. Jahrb. XXXVI (1905), Beiblatt 82, 27: — Amitostigma gracile 
Schlechter in Fedde, Repert. Beiheft TV (1919) 93. — Yun nan. Peye tsi. 
-Koan ie myao. (Siméon Tén no. 1178!) 


Meinen Untersuchungen und dem, was ich nach nochmaliger Prifung 


des Materials gesagt habe, habe ieh nichts hinzuzufugen noch etwas zurick- 
zunehmen. Sollte sich trotzdem an frischem Material nachweisen lassen, 
dak die Blute eine //N gestaltete Narbenflåehe besitzt, so wåre damit 
ihre Zugehårigkeit zu Gymnadenia erwiesen, dann aber wåre es wohl- 
getan, alle Orchidaceen, die wir jetzt als echte Cynosorchis- Arten ansehen, 
auf eine derartige Narbenflåche hin noch einmal zu revidieren. Der leicht 
umgebogene Rand der Antherenkanåle findet sich åhnlich bei Hemipilia, 
es ist die Andeutung einer Bursicula, aber noch keine vollståndige und . 
derartig deplazierte Staminodien, wie sie nach Schlechter hier vorkommen 
sollen, wåren ein Unikum; ich halte diese Organe fir Narbenfortsåtze. — 
Von den l.c. beigebrachten Zitaten ist eins falsch: Gymnadenia tryphiae- 
formis Rchb. f. Otia Hamburg (1878) 51; dort steht nåmlich das gerade 
Gegenteil, denn diese Art ist von Reichenbach selbst zuriickgenommen, 
publiziert ist sie in Journ. of Botany XIV (1876) 209. Die Stelle in den 
»Otia" lautet: ,,Haec omnia a viro ill. (Miquel) scripta .... me seduxerunt, 
ut eandem plantam (G. gracilis Miq.) sub nomine G. tryphiaeformis de- 
scripserim.”" Daraufhin durften ,,Otia'' nicht zitiert werden, das richtige 
Zitat wåre in Engl. Jahrb. 1. supra c. zu finden gewesen (abgesehen vom 
Ind. Kew.). Auf eine weitere Discussion tuber die Gattung Amitostigma 
Schlechter mich hier einzulassen, habe ich keine Veranlassung. 
12. Platanthera Susannae Lindl., Gen. et Spee. Orch. (1835) 295. 


SKREG mør 


k 


AD s Lynn” på 







tå 


CÅKGRLES, 





nl) ordet 


SNEDE Men 


SÅ Ud SUSE AE 


Saaler ES EEK ED NEJ UREEEE ge 
mes 0 Orchidaceae Ténianae Yunnanenses. 103 
Huc ma judice Pectetlis Henryi Schlechter in Fedde, Repert. Beiheft 4 
— (1919)121. —Yun nan. Peyen tsin. Ex silvis. (Siméon Tén no. 1326!) 
; Die mir vorliegenden Exemplare sind in Abmessungen und sonstigen 
= Merkmalen ein genaues Mittel zwischen Pl. Susannae und -der Pflanze, 
owelche Dr. Schlechter jetzt Pecteilis Henryi genannt hat. Das am besten 
erhaltene hat auBerdem ein bisher nirgend erwåhntes Merkmal, die 13 cm 
langen Sporne sind genau von der Mitte an gegenlåufig nach oben gebogen, 
so daB die Spitze nahe der Mindung liegt; auch sind bei demselben Exemplar 
die Unterschiede zwischen Laubblåttern und Stengelscheiden nahezu 


" verwischt und die Seitenzåhne des Labellums kirzer als ich sie je beob- 


achtet habe. Gleichwohl kann ich mich nicht entschlieBen, hierauf hin 
eine neue Art zu machen, so groB die Versuchung auch ist und von j: her 
war, in Pl. Susannae eine Gruppe von Årten zu sehen, denn die anderen 
Exemplare von demselben Standort zeigen alle diese Merkmale ab- 
geschwåcht oder undeutlich. — Eine Zwischengattung als Bricke zwischen 
—… Habenaria Willd. und Platanthera L. C. Rich. ist nicht måglich. Ich stutze 
mich hierbei auf die Ansicht Sir Joseph Hookers, der sich mit mir lange 
uber diese Frage (gelegentlich meiner Dissertationsschrift uber Habenaria) 
unterhalten hat. Auf Grund der Untersuchung zahlreicher indischer Arten 
. miisse er die Wichtigkeit der Antherenkanåle ablehnen, da sie in allen 
Graden vorkommen vom vålligen Fehlen bis zur vollståndigen Åhnlich- 
keit mit denen von Habenaria. Die Narbenfortsåtze wolle er gelten lassen, 
es fånden sich aber nicht selten zwei buckel- oder bruståhnliche Vor- 
wolbungen auf der Narbenflåche, die ebenfalls von ganz schwachen An- 
fången bis zu stark entwickelten Protuberanzen variierten, doch gab er 
schliehlich allerdings zu, dab Narbenfortsåtze und Narbenflåchen zwei 
recht wesentlich verschiedene Dinge seien. Es handelt sich also um Merk- 
måle, die man von jeher kannte und wenn Finet daraufhin seine Gattung 
Hemihabenaria aufstellte, die sich nicht durchzusetzen vermochte, so gilt 
genau dasselbe von der ganz einseitig auf Pl. Susannae und nur auf diese 
zugeschnittenen alten Rafinesqueschen Gattung Pecteilis. 

13. Platanthera praeustipetala Krånzl., spec. nov. — Tuberidia 
ellipsoidea, 1,5 ad 3 cm longa, 6 ad 7 mm crassa. Caulis cum spica 25 ad 
30 cm altus, basi tantum foliatus. Folia 2 v.3 anguste linearia, acuminata, 
— utplurimum 10 cm longa, 1,5 ad 2 mm lata, folium 1 multo minus in scapo. 
Spica plerumque 10 interdum 12 cm longa (tortilis ?), secundiflora, floribus 
15 ad 18, bracteae lanceolatae, acuminatae, 4. mm longae, ovarium cur- 
vulum, brevi-rostratum, 3 mm longum. Sepala ovato-oblonga, obtusa, 
concava, dorsale modice carinatum, omnia 3 mm longa, basi vix I mm 
lata, textura tenera. Petala sublongiora, e basi late ovata subito in acumen 
lineare, crassiusculum v. subcartilagineum, quasi praeustum contracta, 
4,23 mm longa, basi 1,53 mm lata ibique trinervia. Labellum in quarta 
basilari integrum, subquadratum, deinde tripartitum, lobi lineares, laterales 
quam intermedius paulo breviores et angustiores, 3 mm longi, intermedius 
3,5 mm longus, calcar dimidium usque filiforme, deinde leviter inflatum, 
obtusum, 4 mm longum. Gynostemium latum quam altum, glandulae 


104 Fr. Krånzlin. 


conspicuae, massulae grosse granulatae. Flores pallide flavi. — Fl. excunte 
Septembri. — Yun nan. Pe tsao lin. Ex silvis. (Siméon Tén no, 1384!) 

Im ganzen Åufbau erinnert die Pflanze mehr an manche Peristylus- 
Årten, eine Verwandtschaft, an die man angesichts des Spornes und 
besonders der Såule nicht denken darf. Auch gewisse Herminium-Årten 


konnte man zum Vergleich heranziehen. Die Bliten stehen alle nach einer 


Seite und erinnern sehr an Herminium, soweit nur das ÅuBere in Be- 
tracht kommt. Das Labellum hat an der Basis einen flach ausgehåhlten 
quadratischen Teil, an den sich die drei wenig verschiedenen Lappen 
ansetzen, der Sporn ist annåhernd so lang als die Lippe. Sehr apart sind 
die Petalen, welche aus breiter Basis ziemlich rasch in eine sehr viel 
schmalere, dunklere, etwas knorpelige Spitze versehmålert sind; auch dies 
kommt, wenn auch selten, bei Herminium vor. Von diesem Merkmal 
habe ich den Speziesnamen entlehnt. Die Blåtter, zwei ziemlich tief unten 
stehende und ein håher stehendes, kleineres, wurden nahebei den Aus- 
druck ,,filiformia'" rechtfertigen, sie sind indessen etwas zu flach. 

14. Platanthera Henryi (Rolfe) Krånzl., Orch. Gen. et Spec. I (1899) 
632 — Pl. Henryi Rolfe in Journ. Linn. Soc. XXXVI (1903) 55 et R. 
Schlechter in Fedde, Repert., Beihefte Bd. 4 (1919) 111. — Habenaria 
Henryi Rolfe in Kew Bull. (1896) 202. 5 

Herr R. A. Rolfe hat iibersehen, daB ich bereits die Umtaufung 
seiner Habenaria Henryi hatte vornehmen miissen und Schlechter ist 
ihm darin gefolgt. 

15. Platanthera minor Rolfe in Bot. Ztg. (1878) 75. — Yun nan. 
Pe tsao lin. (Siméon Tén no. 1390!) 

16. Platanthera ? Båakeriana Krånzl., Orch. Gen. et Sp. (1898) 633. — 
Yun nan. Ta tsien teon. (Siméon Tén no. 1334!) 

Das einzige vorhandene Exemplar hatte nur beschådigte Bliten, ich 
habe mich genåtigt gesehen, wesentlich nach dem Habitus zu bestimmen 
und das ist gerade in dieser Gruppe von Platanthera ziemlich nichtssagend. 

17. Platanthera obcordata Lindl. in Wall. Cat. (1828) et Gen. et Sp. 
Orch. (1835) 290. — Yun nan. Siao tsin ho. (Siméon Tén no. 1213!) 

Von dem sehr abweichenden Habitus abgesehen ist die Pflanze eine 
typische Platanthera, besonders was die Struktur der Såule angeht. Ein 
zwingender Grund, gerade hier eine neue Gattung (Phyllomphax Schlechter) 
mit den dadurch unvermeidlichen Umtaufungen aufzustellen, liegt keinen- 
falls vor. 

Platanthera Florenti Franch. et Sav., Enum. pl. Jap. II (1879) 32. — 
Als ich 1897 an dem I. Bd. meiner Orchid. gen. et spec. schrieb, habe 
ich, wenn schon mit grobem Bedenken und ? diese Art als Synonym zu 
Pl. minor ØRchb. f. gestellt. Material hatte ich nicht zur Hand und den 
von Franchet angewendeten Vergleich mit Pl. bifolia L. C. Rich. nahm 
ich leider ernster, als er es verdient. Die Pflanze ist inzwischen nochmals 
als Pl. listeroides Takeda in Tokio Bot. Mag. XXIV (1910) 109 beschrieben 
worden und es ist sehr schade, dak dieser auBerordentlich gut geprågt. 
"Name nicht beibehalten werden kann. Die Pflanze hat zwei fast gegen- 


Orchidaceae Ténianae Yunnanenses. 105 


ståndige basale Blåtter und ist den Bluiten nach eine Platanthera, damit 
ist die Åhnlichkeit zu Ende, im ibrigen ist sie von den Blåttern angefangen 


"bis zu den Einzelheiten der Bliten vållig verschieden von unserer 


europåischen Pl. bifolia, wåhrend sie in der Tat an Listera ovata erinnert. 
— Ich nehme also die von mir 1897 geschriebene (1899 publizierte) 
Zusammenziehung mit Pl. minor Rchb. f. zuruck. 

18. Habenaria Delavayi Finet in Rev. Gen. Bot, XIII (1901) 527. — 
Yun nan. Teyen tsin. (Siméon.Tén no. 1216!) 

19. Habenaria glaucifolia Bureau et Franchet in Journ. de Botan. V 
(1891) 152. — Yun nan. Nithon. (Siméon Tén no. 1422! 1425!) 

20. Habenaria diceras Schlechter in Notes Roy. Bot. Gard. Edinb. 
XXIV (1912) 101 t. 78. — Yun nan. Ni thon pr. Pe tsao lin. (Siméon 


"Ten no. 1381!) 


21. Habenaria Miersiana Champ. in Hook. Kew Journ. Bot. VII 
(1855) 37. — Yun nan. Peyen tsin. (Siméon Tén sine no. !) 

22. Habenaria lacertifera Benth., Fl. Hongkong. (1872) 362. — 
Hong kong. Happy valley. (Feilberg sine no. !) 

23. Habenaria atramentaria Krånzl., spec. nov. — ( Peristyloideae.) 
Tuberidia globosa, 1,5 cm diametro.  Caulis gracilis, strictus, leviter 
flexuosus, cum spica 30 ad 36 cm altus, basi foliatus. Folia 3 v.4 pro caule 
parva, lanceolata, acuminata, 4 ad 5 cm longa, 1 ad 1,2 cm lata, 2 ad 3 cm 
inter se distantia, additis foliolis ad 5 parvis, bracteiformibus, acumina- 
tissimis, spica ad 13 cm longa, %, totius plantae occupans, multiflora, 
cylindracea, laxi- aut densiflora, bracteae lanceolatae, acuminatae, in- 
feriores flores, superiores ovaria aequantes, 1,2 ad 8 mm longae, ovaria 


" non rostrata, ad 8 mm longa. Sepala oblonga, obtusa, concava, carinata, 


lateralia cum apiculo vix prominulo ante apicem ipsum, 3,5 mm longa. 
Petala sublatiora, aequalia, oblonga, obtusa, non apiculata. Labellum 
e basi integra mox in lobos 3 lineares divisum, quorum laterales fere fili- 
formes cum intermedio breviore latiore triangulo, obtuso cruciati, sub- 
reflexi, omnes apice ipso obtusi, intermedius 3 mm longus, laterales 3,5 ad 
4 mm longi, calcar filiforme, 5 mm longum apicem versus leviter incrassa- 
tum, acutum. Gynostemium, minimum in fundo calcaris subabscondi- 
tum, apices processuum stigmaticorum nitiduli, vix (sub lente valida) 
eonspicui, antherae canales nulli. Tota planta sicca (more Brachycory- 
thidis) nigra. Flores pallide (paulo) flavi. — Fl. Septembri. — Yun nan. 
Peyen tsin. (Siméon Tén no. 1261!) 

Wenn ein Bastard zwischen einer Habenaria und einer Brachycorythis 
zumal in China uberhaupt måglich wåre, so multe er habituell so aus- 
fallen, wie die hier beschriebene Art. Und zu diesem Gesamtbild stimmt 
auch ganz gut das aubergewohnlich winzige Gynostemium, dessen Narben- 
fortsåtze, oder richtiger deren Spitzen nur mit Zuhilfenahme der 30fachen 
VergråBerung zu finden sind. Nach den rein technischen Einzelheiten 
gehårt die Pflanze in eine leider schon sowieso an Mikrospezies reichen 
Gruppe der Habenaria acuifera, Hancockii und åhnlicher. Nicht håufig 
ist wenigstens in diesem Formenkreise das Auftreten eines winzigen Kiels 


106 Fr. Krånzlin. 





und einer minimalen Spitze auf den seitlichen Sepalen, des ferneren sind. 
die drei Abschnitte des Labellums und ihre Långenverhåltnisse etwas 
abweichend von denen der verwandten Arten. — Alles in allem eine un- 
erfreuliche Art, die nicht aufstellen zu missen ich viel vergebliche Muhe 
aufgewendet habe. - 
24. Habenaria peyentsinensis Krånzl., spec. nov. — (Plantagineae.) 
Tuberidia ellipsoidea v. ovalia, 1,5 cm longa, villosa. Caulis cum floribus 
12 ad 18 cm altus, strictus, Folia plerumque 3, dissita, latissime ovata, 
infimum fere orbiculare, sequentia paulo angustiora, omnia apiculata, 
albido-marginata, 2 ad 5 cm longa, 2 ad 3 cm lata, vagina in scapo I acu- 
minata, 3 cm longa. Spica pauci- et laxiflora (5- ad 8-flora),- bracteae 
lanceolatae, pedicellum cum ovario aequantes, 1,5 ad 2 cm longae, su- 
premae breviores. Sepalum dorsale ovatum, acutum, cucullatum, 8 mm 
longum, basi (expansum) fere aequilatum, sepala lateralia deflexa, ovata, 
acuta, obliqua v. asymmetrica, 1 cm longaåa, 5,5 mm lata. Petala simplicia, 
erecta, cum sepalo dorsali non conglutinata, linearia, subfalcata, 6 mm 
longa, vix 1 mm lata. Labelli lobi laterales rhombei, margine exteriore 
minute denticulati, lobus intermedius linearis, e basi paulo latiore sensim 
angustatus, apice subito acutatus, totum labellum basi cuneatum, 1 cm 
longum et inter lobos laterales latum, calcar leviter curvatum clavatum, 
apicem versus leviter inflatum, obtusum, 1,8 ad 2 cm longum. Anthera 
crassa, 3,5 mm alta, canales antherae 2,5 mm longae, recurvae, processus 


stigmatici erassiusculi, recti, aequilongi, staminodia pro flore magna, 


triangula, crasse papillosa. — Flores albi. — Fl. ineunte Septembri. — 
Yun nan.. Peyentsin. (Siméon-Tén no. 1352!) 

Ein Exemplar dieser Pflanze mit abgebrochenem Blitenstand wurde 
jeder fur Platanthera viridis Lindl. halten, so sehr gleich sind die Blåtter 
und der ganze Habitus. Zieht man die Merkmale alle in Betracht, so 
erhålt man ein Bild, das an Hab. Richardiana. R. Wight gemahnt, aber 
bei dieser Art stehen 4 bis 6 sehr spitze Blåtter ziemlich dicht am Grunde 
desStengels; die uibrigen Teile, besonders die Blåtter der Blite stimmen so 
ziemlich. Weder diese Art noch eine der Verwandten hat Staminodien 
von dieser GråBe und dieser runzligen rauhen Oberflåche. Es ist somit 
zunåchst sicher, dab Hab. Richardiana nicht vorliegt und eine ins einzelne 
gehende Untersuchung fårdert noch mehr Unterschiede zutage. Unter 
den Habenarien von Schlechters Prodom. Orchid. Sino-Japonicae ist 
keine die gut mit diesen Merkmalen hier ubereinstimmt. 

25. Habenaria bihamata Krånzl., spec. nov. — (Diphyllae.) Tu- 
beridia mihi non visa. Folia 2 humistrata, suborbicularia, brevi-api- 
culata v. subrhombea, 2,5 cm longa et lata (in uno specimine 5 cm longa 
lataque). Scapus tenuis 20 ad 25 cm (48 cm) altus, brevissime pilosus, 
foliolis bracteiformibus paucis valde dissitis praeditus, spica 7 (17?) cm 
longa, obscure distichantha, pluri- ad multiflora, laxiflora, bracteae 
lanceolatae, acuminatae, 3 ad 4 mm longae, ovarium cum rostro 7 mm 
longum. Sepalum dorsale ovatum, obtuse acutatum. 4 mm longum, basi 
2,5 mm latum, lateralia triangula, subfalcata, acuminata, 4,5 mm longa, 








EEN SILKE SEE Se ERE 
RE SEER 


: "Orchidaceae Føbiande RER DER: FOR 


” energice in ovarium reflexa. Petala cum ser alo dorsali arctissime con- 
" glutinata, illique aequilonga, triangula, acuminata, in dimidio postico 
" membranacea. Labelli basin usque partiti partitiones laterales filiformes, 
— arctissime reflexae, apice cornuum instar recurvatae, 4 mm longae, partitio 
=å antermedia linearis, arctissime, deflexa-latior quam laterales, apice obtusa, 
"5 ad 6 mm longa, calcar striete dependens per duas tertias filiforme, deinde 
— inflatum, apice obtusum, 6 mm longum, lobo intermedio labelli parallelum 
— et aequilongum, canales antherarum semierecti, processus stigmatici 
—… longiores, deflexi, antice incrassati, leviter bipartiti. Flores rubri. — 
— Fl. Julio. — Yun nan. Peyentsin. (Siméon Tén sine no.!) 

g Eine typische Art der Sektion Diphyllae und sehr nahe verwandt 
— mit Hab. diceras Schlechter. Auffållig ist zunåchst die rote Farbe der 
—… Bluten. Der Sammler, dessen Latinitåt etwas angealtert zu sein scheint, 
—. sagt aber ausdricklich ,,Fl. rubus! Die in ( ) gesetzten MaBe beziehen 
— sich auf ein wesentlich gråkeres Exemplar, welches bei den anderen lag, 
"aber leider von der Fåulnis derart mitgenommen war, daB eine Unter- 
— suchung der Bliiten zu nichts gefiihrt håtte; soweit es sich ohne die Bliiten 
aufzukochen ersehen liek, glichen sie denen der drei kleineren Exemplare. 
É Den Abmessungen -der Sepalen und Petalen nach kånnte man an Hab. 
+ diplonema Schlechter denken, aber dem widersprechen die AusmafBe 
— des Labellums, welche vållig verschieden sind. Ferner stimmen die der 
 Såule durchaus nicht. Trotz allér Abweichungen doch eine sehr åhnliche 
— "Art. Die Abbildung von Hab. diceras in den Notes R.B. G. Edinb., plate 
= LXXVIII, zeigt einen sebr viel dichteren Bliitenstand, die von Hab. di" 


— plonema eine viel schwåchere Pflanze; viel anzufangen ist mit beiden 
— Abbildungen nicht. 

2 26. Habenaria Simeonis Krånzl., spec. nov. —(Chlorinae — olim 1898.) 
—… Tuberidia ellipsoidea, interdum subcylindracea, 2,5 ad 4 cm longa, 1 ad. 
x 1,5 em crassa. Caulis cum inflorescentia 20 ad 30 cm altus, strictus v. 
;: leviter flexuosus. Folia in tertia inferiore caulis plerumque 3, longe vagi- 
— nantia, compressa, oblonga v. oblongo-lanceolata, acuminata, in vaginas 
— caulis illaeque in bracteas -decrescentia, basilaria saepius recurvata, ad 
— 6 cm longa, ad 2 cm lata, vaginae caulis 3 acuminatissimae, spica sub- 
— eylindracea interdum capitata, pluri- ad multiflora, 4 ad 7 cm lønga, 3 em 
E diametro, bracteae florum anguste lanceolatae, longissime acuminatae, 
— carinatae, margine et antice in dorso fimbriatae, 1,5 ad 1 cm longae. 
— Pedicelli cum ovariis 1,5 cm longi, ovaria rostrata. Sepala petalaque 
… forma et magnitudine vix diversa, late oblonga v. ellipsoidea, antice 
z 


rotundata, sepala lateralia semideflexa, paulo tantum majora subobliqua, 
dorsale cucullatum, illud et petala 5 mm, lateralia 6 mm longa, sep. dors. 
et petala 3 mmslata, lateralia 3,5 mm. Labellum trilobum, basi ipsa sim- 
plice, lobi laterales parvi, lineares, paulum- infra basin oblique inserti, 
2,5 mm longi, lobus intermedius 8,5 (totum labellum 9 mm) longus, duplo 
latior quam laterales, calcar filiforme a dimidio apicem versus vix in- 
" crassatum, 2,8 ad 3 cm longum; Gynostemium valde reclinatum, pro- 
cessus stigmatici arcte deflexi, crassiusculi canales antherarum elongati 


Sa UELSA nos UNS 


108 Fr. Krånzlin. 


protensi, recti, rostellum breve, triangulum, pronum. Flores flavi.:— 
Fl. exeunte Augusto. — Yun nan. Pe yen tsin. (Siméon Tén no. 1255 :) 

Eine wenn auch nur sehr unbestimmte Ahnlichkeit mit dieser Art 
hier zeigt die Abbildung von Hab. viridiflora KR. Br. in R. Wight's Icon. 
pl. Ind. or. V, t. 1705. Ich betone nochmals, dab nur der allgemeine Ein- 
druck mit diesem Vergleich wiedergegeben werden soll, bei der genaueren 
Untersuchung verfliichtigt sich alle und jede Ubereinstimmung. = Diése 
Årt ist schwer zu deuten und dal alle von Hooker in Fl. Brit. Ind. VI, 
150 hierzu einbezogenen Årten wirklich zu ihr gehåren, ist mir sehr zweifel- 
haft: so z. B. die von Griffith in Notulae IIT, 269 beschriebene und in den 
Icon. Pl. Asiat. III, t. 342 abgebildete Pflanze. Ohne sehr reiches Material 
ist diese Frage nicht zu låsen; Reichenbach, der die Orchideen aus Rob. 
Browns Prodromus bekanntlich zum Gegenstand einer besonderen Studie 
gemacht hat, hat gerade diese Frage nicht beruhrt. Nun fehlt freilich 
in den beiden bisherigen Aufzåhlungen chinesischer Orchidaceen, der von 
R.A. Rolfe und der von Schlechter, Hab. viridiflora R. Br., was aber nicht 
viel besagen will, denn auf Vollståndigkeit machen diese Arbeiten keinen 
Anspruch. Mag nun Hab. viridiflora R. Br. eine Art, sein, oder, was mir 
das wahrscheinlichere scheint, ein Knåuel von Arten, so muBte ich bei 
dieser neu aufgestellten mich mit der Stammart auseinandersetzen, in 
deren Nåhe sie im System gehårt. 

Herr Dr. Schlechter ibt an der von mir aufgestellten Gruppe der 
»»Chlorinae'' eine etwas abfållige Kritik. Dal diese Gruppe viel zu wunsehen 
ubrig låkt, wukte ich bereits vor zirka 25 Jahren und habe es klar 
und deutlich auf S. 397 meiner damaligen Bearbeitung der Orchidaceen 
gesagt. Die Sache ist also nicht gerade neu. — DaB ich mit dem Material 
von heute und dem, was ich mittlerweile gelernt habe, jetzt vieles anders 
sagen wurde, mag Herr Schlechter mir glauben oder auch nicht — nihil 
curo. Wenn ich somit fur die Kritik die Prioritåt fur mich in Anspruch 
nehmen darf, so uberlasse ich ihm um so bereitwilliger die der an dieser 
und anderen Stellen von ihm beliebten Form und Ausdrucksweise, denn 
die ist in der Tat nen. 

27. Habenaria erassilabia Krånzl. spec: nov. —  Peristyloideae- 
Tradescantifoliae.) Tuberidia ellipsoidea, ultra 3 cm longa, 1,2 cm crassa. 
Cataphylla mox grandescentia, infrafoliacea 3 v. 4, ochreata, ample va- 
ginantia, folia 4 (v. interdum plura?) fere basilaria, oblonga, acuta, ap- 
proximata, membranacea, satis tenera, 5 ad 8 cm longa, 1,8 ad 3,2 cm 
lata, plurinervia. Scapus cum inflorescentia 30 ad 36 cm longus, foliolis 
bracteiformibus 4 longe acuminatis praeditus, spica multi-satis laxiflora, 
ad 15 cm longa, dimium fere totius plantae efficiens, bracteae longe acu- 
minatae; ovato-lanceolatae, 1,3 ad 1 cm (supremae) longae, ovaria brevi- 
rostrata aequantes.  Sepala ovata, acuta, tenui-membranacea, 4 mm 
longa, basi 3 mm lata. Petala late ovata, obtusav.retusa, fere subbilobula, 
sepalis aequilonga, crassiora quam sepala. Labellum basin usque tri- 
partitum, lobi laterales ovati, falcati, acuti, lobus intermedius ligulatus, 
obtusus, totum labellum textura firma, satis crassa, partitiones omnes 





Orchidaceae Ténianae Yunnanenses. 109 


inter se et sepalis petalisque aequilongae, calcar e basi filiformi ampliatum, 
paulum incrassatum, acutum, incurvum, 6 mm longum. Gynostemium 
breve, anthera pro rata parva, canales breves, rostellum amplum, pro- 
cessus stigmatici quam canales longiores. Flores pallide flavi. — Fl. 
éxeunte Augusto. — Yun nan. In silvis pr. Pa yen tsin. (Siméon 
TF én no. 1333!) 

Soweit die Frage ohne Vergleichsmaterial klarzustellen ist, wird 
man Hab. Hancock Rolfe als die nåchstverwandte Art ansehen miissen, 
sie hat allerdings auch Anklånge an die mehr åstliche Hab. tradescanti- 
folia Rchb. f. von den Fidji-Inseln, deren sie noch mehr åhneln wiirde, 
wenn die Blåtter etwas håher am Stamm hinaufgerickt wåren. 2 

28. Peristylus goodyeroides Lindl., Gen. et Sp. Orch. (1835) 299. — 
Yun nan: Peyentsin. (Siméon Tén no: 1217! 1218!) 

29. Satyrium nepalense Don.:, Prodr. Fl. Nepal. 26 (1802—03!. — 
Yun nan. Nithon. (Siméon Tén no. 1421!) 

var. Wightiana Hook. f. in Fl. Brit. Ind. VI (1894) 168. — Yun nan. 
Pe tsen lin. (Siméon Tén no. 1416!) 

30. Satyrium microcephalum Krånzl.; spec. nov. — Tuberidia ob- 
longa v. ovata, ad 3 cm longa, villosa. Caulis cum spica 15 ad 22 cm altus, 
cataphylla infrafoliacea 2, laxe vaginantia, supra retusa. Folium unicum. 
ovatum v. oblongum, obtuse acutatum v. obtusum, brevi-vaginatum, 
lamina 2 ad 5 cm longa, 1,8 ad 2,5 cm lata, foliola in scapo-l v. 2, lanceolata, 
acuta, spica brevis, subcapitata, pauci- (6- ad 8-) flora, 2 ad 2,5 cm longa, 
1,5 ad 2 cm diametro, bracteae infimae reflexae, ovatae, acuminatae, 
9 mm longae, supremae erectae, minores, ovarium brevij-rostratum, pro 
tlore crassiuseulum, 4 ad 5 mm longum, 2 mm crassum. Sepala petalaque 
paulo minora oblonga, concava, obtusa, basin usque libera, 3 mm longa, 
vix 1,2 cm lata, omnia margine nusquam, extus tantum sub lente valida 
minute papillosa. Labellum subglobosum, profunde cucullatum, extus 
carinatum, apice paulum erosulum, haud explanandum, 4 mm longum 
et basi latum, calcaria filiformia v. leviter curvata, ovarium subaequantia. 
Gynostemii stigma quadratum, rostellum trilobum, lobus intermedius 
major, dilatatus, laterales multo minoses. Flores flavi. — Fl. exeunte 
Septembri. — Yunnan. In silvis Ni thon. (Siméon Tén no. 1397!) 

Eine zierliche, kleine Pflanze, welche als neu zu erkennen leicht 
genug war. AuBer dem Habitus ist an der Pflanze sonst beinahe nichts 
von besonderem Interesse zu bemerken, dieser ist allerdings auffallend 
genug und auch bei dieser Art vållig abweichend von dem Typus des 8. 
nepalense Don, den die såmtlichen bisher aus China bekannten Satyrium- 
Årten mehr oder minder stark variieren. Bemerkenswert ist auch der 
Standort ,,in silvis". Ich hatte 5 Exemplare zur Verfugung, welche ab- 
gesehen von ihrer wenig verschiedenen GråBe alle einander glichen. 

31. Hemipilia flabellata Bur. et Franch. in Journ. de Botan. V (1891) 
152. — Yunnan. Peyen tsin, in silvis. (Siméon Tén sine no.!) 

Als Abweichungen von der Originaldiagnose wåre zu vermerken: 
1. Der Schaft hat nicht mehrere sondern nur ein einziges deckblattåhn- 


TUD) Fr, Krånzgsin, 


liches Blåttchen. 2. Das Labellum ist in UmriB finfeckig mit vorn aus- 
gerandeter Vorderseite. 3. Jede Spur von Zåhnelung des vorderen. Randes 
fehlt. Alle anderen Merkmale stimmen vortrefflich, vor allen Dingen ist 
,flabellatum" die einzig mågliche Bezeichnung fur das Labellum, welches 
auch (wie sonst die ganze Pflanze) in ihren Abmessungen gut stimmt, 
Ich wage nicht, auf diese Abweichungen hin eine nov. spec. zu machen. 
Wie enorm die Anzahl der Orchideen anschwillt, davon ist Hemipilia ein 
Beispiel, Schlechter zåhlt 11 gut unterscheidbare Arten auf und man E 
kann bereits die Unterabteilung der Cordifoliae von den anderen ab- 
trennen und H. cordifolia Lindl. war lange Zeit die einzige bekannte ARE 
Bentham kannte zu Anfang der achtziger Jahre nur 2 Arten. 







32. Hemipilia silvatica Krånzl., spec. nov. — Tuberidia ellipsoidea, 
interdum stipitata, ad 2 cm longa, cataphyllum 1 acutum sub folio late 
ovato-acuto, 1,5 ad 3 cm longo, 3/4 ad 2 cm lato. Scapus tenuis, omnino 
nudus cum spica4 ad 12 cm altus, tenuis, spica secundiflora, 2- ad 12-flora, 
floribus haud resupinatis, bracteae lanceolatae, acuminatae, ovaria sub- 
semiaequantes, 6 mm longae, ovaria rostrata, 9 ad 10 mm longa. Sepala 
oblonga, obtusa, dorsale concavum, vix 3 mm longum, lateralia erecta, i 
paulo ultra 3 mm longa. Petala elliptica, apice rotundata, tenerrima, å 
2 mm longa, I mm lata. Labelli lobi laterales parvi, oblique divergentes, 
apice rotundati, apiculati, lobus intermedius e basi cuneata obtriangulus, 
antice -retusus, medio apiculatus, totum labellum 5 mm longum, lobus 
intermedius antice 3 mm latus, lobi laterales 2 mm longi, calcar pro flore 
amplum, obtusum, 3 mm longum, sepalis aequilongum. Gynostemium 
breve, globosum, rostellum lineare, antheram non aequans. Flores albi. — 
Fl. exeunte Septembri. — Yun nan.  Nithon, prope Pe tsoo lin. 
(Siméon Tén no. 1396!) 


Eine kleine, sehr zierliche Art, die wohl H. cruciata Finet zunåchst 
steht, aber keinenfalls identisch ist. Die Exemplare variieren sehr an 
GroBe, die kleinsten hatten nur 4—5 cm Hohe, mit 2 Bliten, die gråkten 
12—13 cm Håhe mit ungefåhr 12 Bliten. Diese kleinsten åhneln im 
Habitus ungemein gewissen winzigen Platanthera-Arten des japanisch- 
chinesischen Gebietes. 


33. Herminium Ténianum Krinzl., spec. nov. — Tuberidia globosa 
1,5 ad 2 cm diametro, cataphyllum 1 infrafoliaceum, laxe vaginans acutum, 
Folium 1 oblongum v. oblongo-lanceolatum, acutum, 4 ad 10 cm longum, 
1,5 ad 2 cm latum, addito foliolo I minore acuto medio in scapo, totus 
caulis cum spica 18 .ad 24 cm altus, spica densiuscula, multiflora, 8 ad 
10 cm longa, bracteae lanceolatae, acuminatae, ad 5 mm longae, ovaria 
superantes. Sepalum dorsale late oblongum, obtusum, 3 mm longum, 
2 mm latum, lateralia similia, minora, subaequilonga, vix 2 mm lata, 
obtusa. Petala latissime ovata v. potius rhombea dicenda, obtuse acutata 
apice subcartilaginea, 2 mm longa et basi lata. Labellum maxima pro 
parte simplex, antice inlobulos 3(quorum intermedius vix longior) aequales, 
obtusos divisum, callus nasiformis, subelavatus, proclivis, apice obtu- 





'” Gi: 


ARGO ride Ål e Dee skt 


> 
== 
= 
E 
E- 3 
i 
åd 
, 


Orchidaceae Ténianae Yunnanenses. 111 


sissimus, dense minuteque papillosus in medio disco, totum labellum 2,5 mm 


longum et latum. Gynostemium latissimum. Flores pallide flavi. — Fl. 
exeunte Septembri. — Yun nan. Pe tsao lin in silvis. (Siméon Tén 
no. 1387!) ) ; 


Diese Art ist die zweite oder måglicherweise die dritte einer Gruppe 
von Herminium, welche auf der Lippe mit einem hornartigen Callus ver- 
sehen sind. Dr. Schlechter zieht in seinem Orchid. Sino-Japon. Prodromus 
unter dem Namen H. coeloceras (Finet) Schlechter das von mir 1903 auf- 
gestellte H. unicorne mit Peristylus coeloceras Finet (1901) zuammen. 
Diese Zusammenziehung kann berechtigt sein, es spricht aber doch manches 
dagegen. Zunåchst bedeutet coeloceras ,,Hohlhorn”. Wenn in meinen 
H. unicorne die Protuberanz der Lippe hohl gewesen wåre, håtte ich dies 
bemerkt und es in meine Diagnose aufgenommen; ich habe die Pflanze 
augenblicklich nicht zur Hand, entsinne mich ihrer aber noch ganz genau. 
Hiermit ist nichts gegen die von Schlechter eingefiihrte nov. comb. gesagt, 
aber die Zugehårigkeit meiner H. wnicorne måchte ich zunåchst wenigstens 
bezweifeln. Die Beschreibung dieser Art hier zeigt Punkt fir Punkt Ab- 
weichungen von der von H. unicorne, angefangen von den Blåttern bis 
zur Lippe, deren Protuberanz wie eine warzige Nase aussieht. 

34. Herminium angustifolium Benth. ex Hook. f. Fl. Brit. Ind. VI 
(1890)129.—Yun nan Pany tien(Pe yen tsin). (Siméon Tén no. 1174! 


1221!) 

35. Herminium Limprichtii Schlechter in Fedde, Repert. Beiheft 4 
(1919) 42 et 101..— Yunnan. Ta tsintao. (Siméon Teén no. 1324! 
1325!) 

36. Coelogyne elegantula Krånzl., spec. nov. — Planta terrestris, 


radicibus villosis obsita. Pseudobulbi subglobosi, magnitudine nucis 
Avellanae, mox in caulem 4 ad 5 cm longum, tenuem, cylindraceum atte- 
nuati, cataphyllis paucis arctissime vaginantibus obsiti, bi- v. trifoliati. 
Folia angustissima, linearia v. lineari-lanceolata, acuminatissima, ad 
15 cm longa, 5 ad 6 mm lata, scapus 12 ad 15 cm longus, apice pauci- 
florus, nudus, flores 1 ad 4 (interdum forsan plures ?”), bracteae sub anthesi 
nullae, cicatrices tantum vidi, pedicelli cum ovariis 1,5 cm longi. Sepala 
petalaque vix minora oblongo-lanceolata, acuta, 2,2 v. 2 cm longa, 4 mm 
lata, dorsale et petala, conniventia, sepala lateralia subdeflexa. Labelli 
lobi laterales semiobovati, obtusi, lobus intermedius cuneatim dilatatus, 
antice profunde - bilobus, utrinque obtusus, carinae inter lobos laterales 
4 elevataåae, margine ciliatae, in lobum intermedium decurrentes, totum 
labellum 1,8 cm longum, inter lobos laterales circ. I cm latum, lobi laterales 
1.2 em longi, intermedius 5 ad 6 mm longus, antice 5 mm latus. Gyno- 
stemium satis curvatum 1,1 cm longum. Flores ,,rubri'" sec. collectorem. — 
Fl. Octobri. — Yun nan. Panvy tien (anreote?). (Siméon Tén 
no. 1186!) 

Im ganzen erinnert die Pflanze trotz des nicht gewimperten Labellums 
an C. fimbriata Lindl. Die Scheinknollen sind auffallend klein, nahezu 


kugelig und endigen in einen langen Halsteil, der dann 2 oder 3 auffallend 


112 Fr. Krånzlin. 


lange, schmale Blåtter und einen gleichfalls schlanken Blitenschaft trågt. 
Die Bliten haben schon vor der Blutezeit keine Deckblåtter mehr, ihrem 
Bau nach sind es typische Coelogyne-Bluiten, wie sie sehr oft vorkommen, 
aber sie sind ,,rot" und das ist bei dieser Gattung auBerordentlich auf- 
fallend. Die Farbe der getrockneten Bliten widerspricht der Notiz des 
Sammlers nicht, rote, besonders rosafarbige nehmen stets den Farbenton 
an, ae die — leider spårliehen — Bliten zeigten. 

Arundina chinensis Bl., Bijdr. (1825) 402. — Yun nan. Peyen 
lg See: Tenso. 135EN 

38. Spathoglottis Fortunei Lindl. in Bot. Reg. XXXI (1845) t. 19. — 
Yun nan. Pany tien. (Siméon Tén no. 1249!) 

39. Dendrobium nobile Lindl., Gen. et Sp. Orch. (1830) 79. — Yun nan. 
Pe yen tsin. (Siméon Tén sine no.!) Nomen vern. Che fou.  . 

40. Liparis bootanensis Griff., Not. III (1851) 278, Ic. Pl. Asiat. III 
t: 287. — Yun nan. Peéyen tsin.: in silvis.. (Siméon Tén sine no.!) 

Die Abbildung in King und Pantling, Orch. Sikk. Himal. I, 30 t. 40 
stimmt bis auf Kleinigkeiten im Habitus und gånzlich in den. Bliten. 
L. ovyphylla Schlechter in Fedde, Rep. Beiheft 4, 63 hat zweiblåtterige 
Ståmme und der Blutenstand ist kurzer als die Blåtter, auBerdem sind die 
Bluten gråBer, sonst scheint sie åhnlich zu sein. — In demselben Um- 
schlagbogen lag noch eine ganz andere Art von Liparis. 

41. Liparis Téniana Krånzl., spec. nov. — Pseudobulbi ovoidei ad 
3 cm longi, basi I ad 1,5 cm crassi, cataphyllis quibusdam emarcidis tuni- 
cali. Folia 2 rarius 3, additis foliolis 1 v. 2 in ima basi caulis, oblongo- 
lanceolata, acuta, membranacea, trinervia, 6 ad 12 cm longa, 1,5 ad 
2,5 cm lata, cauli adpressa, dimidium caulis cum inflorescentia 20 ad 36 cm 
alti subaequantia, scapus ceterum nudus, spica 6 ad 12 cm longa, pauci- 
ad pluriflora, sparsiflora (flores 8—15), bracteae lanceolatae, acuminatae, 
3 ad 5 mm longae, pedicelli cum ovariis 1 cm longi. Sepalum dorsale late 
lineare, obtusum, lateralia late oblonga, subobliqua, obtusa, petala 
linearia, obtusa, omnia 8 mm longa, sepala lateralia 2,5 mm lata. Labellum 
cuneato-obcordatum, antice utrinque minute obtuseque denticulatum, 
& dimidio deflexum, basi complicatum, ad insertionem callis 2 acutis, 
subulatis praeditum, totum labellum vi expansum 8 mm longum, antice 
5 ad 6 mm latum. Gynostemium generis. Flores ,,valde violacei'' sec. 
collectorem. — Fl. Augusto. — Yun nan. Siao tsin ho. (Siméon Tén 
ne. 1219!) 

Eine Art, deren Aufstellung mir schwere Be iEnken macht. Sie gehårt 
zu den,, Bituberculata'', womit sehr wenig gesagt ist und violette Bliten sind 
auch kein sehr gutes Merkmal aber die beiden verhåltnismåBig kurzen, 
dem Stengel angedrickten Blåtter von sehr diinnhåutiger Textur geben 
mit allem anderen zusammen doch ein besonders Bild ab. Ich hatte"im 
ganzen 5 Exemplare von gleicher Entwicklung und nur in der GråBe 
verschieden zur Verftgung. 





ER BESERREU GE URKPÆNDEBTOEOG YO EÅEGETG 
HEEOWEREN GER EÆNTS 








ED: 


Scrophulariaceae. 
By 


Fr. J. Mathiesen. 


1921. 


Reprinted from »MEDDELELSER OM GRONLAND« 
Vol. XXXVIT. 


LIBRARY 


Arbejder fra den botaniske Have i København. Nr. 96. 


Hitherto, the following papers have been published in "Med- 


delelser om Grønland”: 


MolskssesevI:: 


. Ericineæ (Ericaceæ, Pirolaceæ). 


1. Morphology and Biology. By Euc. WARMING p. 1—71. 
2. The Biological Anatomy of the Leaves and of 
the Stems. By HENNING EILER PETERSEN.. p.73—138. 


. Diapensiaceæ. Diapensia lapponica L. By HENNING 


BILER PETERSEN es SDR SAS eN p. 139—154. 


. Empetraceæ. Empetrum nigrum L. By A, MENTZ p. 155—167. 
. Saxifragaceæ. 


1. Morphology and Biology. By EuG. WARMING p. 169—236. 
2. The biological Leaf-anatomy of the Arctic spe- 


cies of Saxifraga. By OLAF GALLØE....... p. 237—294. 
5. Hippurid aceæ, Halorrhagidaceæ and Callitrichaceæ. 
Bys FA GNE BE SED EEN p. 295—332. 
GÆR an un ease y KNUD JESSEN eee p. 333—440. 
4. Lentibulariaceæ (Pinguicula). By Fr. HEIDE... p. 441—481. 
Vol. XXXVII: 
8. Rosaceæ. Bye ke ESSEN p. 1—126. 
VE Corner Br ARSEN EEN p. 127—150. 
10. Caprifoliaceæ. Linnæa borealis. By O. HAGERUP p. 151—164. 
JP rim ulacee By ER JS MATEESEN ERE REE p. 165—220. 
12. A List of Arctic Caryophyllaceæ, with some 
synonyms byt Ore NEED p. 221—227. 
13. Caryophyllaceæ. By EuG. WARMING......... p. 229— 342. 
LÆ Entates EB ye OR BEN BP OS p. 343—358. 





Preface. 


E the present paper, partly on the basis of notes on the 
subject found in previous publications, and partly on 
the basis of my own investigations, I have given a descrip- 
tion of the ramification, shoot-structure, flower-biology and 
anatomy of some Arctic and Subarctic Scrophulariaceae; in 
addition to which, under each species, the geographical 
distribution has been briefly mentioned, and, as far as the 
literature on the subject rendered it possible, the nature of 
the habitat. 

Besides the herbarium and alcohol material belonging 
to the Botanical Museum of the University of Copenhagen, 
I have, through the courtesy of Mrs. THEKLA Resvorr, Dr. 
phil., with respect to several of the species, had an opportunity 
of investigating some excellently preserved material, collected 
on the mountains of Norway. 

The subject-matter of the present paper was worked 
out in the Plant-anatomical Laboratory of the University 
of Copenhagen, and I wish to express my grateful and heart- 
felt thanks to the Director, Professor C. RAUNKIÆR, for his 
advice and help during the work. 

The majority of the figures illustrating the structure of 
the flowers, are drawn by Professor WARMING, to whom also 
my most sincere thanks are due, partly on account of the 
interest he has taken in my work, and partly because he so 


362 Fr. J. MATHIESEN. 


generously placed his notes on the Arctic Scrophulariaceae 
at my disposal. Fig. 20, B, C and D, Fig. 40 and Fig. 44. 
which are published here for the first time, have been re- 
drawn after Professor WARMING's originals, in order to be 
reproduced according to the method employed here. The 
remainder of the figures are drawn by myself. 

Mr. M. Porsirn, mag. sc., Director of the Danish Arctic 
Station, Greenland, has very kindly read through great parts 
of my manuscript, and made several additions of a biological 
nature, besides having also kindly permitted me to use his 
notes on the geographical distribution of Arctic plants. For 
the help thus rendered to me I herewith beg him to accept 
my best thanks. 


The following species have been investigated:— 


Veronica fruticans Crantz. 
— alpina L. 
— officinalis L. f. glabrata Fristedt. 
Castilleita pallida (L.) Kunth. 
Euphrasia arctica Lange. 
Bartschia alpina L. 
Pedicularis lapponica L. 
-- sudetica Willd. 
— euphrasioides Steph. 
= Sceptrum carolinum L. 
— capitata Adams. 
— hirsuta L. 
— lanata (Willd.) Cham. & Schlecht. 
— flammea L. 
— Oederi Vahl. 


Scrophulariaceae. 363 


Principal literature. 


ÅBROMEIT, J., 1899: Bot. Ergebnisse der ... unter Leitung von Dr. 
von Drygalski ausgesandten Gronlandsexpedition. B. Samen- 
pflanzen (Phanerogamen). (Bibliotheca botanica. 42. Stuttgart). 

ÅNDERSSON, G. and H. Hesserman, 1900: Bidrag till kånnedomen 
om Spetsbergens och Beeren Eilands kårlvåxtflora, grundade 
på iakttagelser under 1898 års svenska polarexpedition. (Bihang 
till K. Svenska Vet.-Akad. handlingar. Bd. 26. Afd. III. No. 1). 

ÅURIVILLIUS, C., 1883: Insektlifvet i arktiska lånder. (Nordenskiådld : 
Studier och forskningar fåranledda af mina resor i håga norden. 
VI. Stockholm). 

ÅXELL, S., 1869: Om anordningarna får de fanerogama våxternas 
befruktning. Stockholm. 

Brytrt, A., 1906: Håndbog i Norges flora. Kristiania. 

BrunNDIN, J. A. Z., 1898: Bidrag till kånnedomen om de svenska 
fanerogama årternas skottutveckling och åfvervintring. Dis- 
sertation. Upsala. i 

BørGeEsEn, F., 1895: Bidrag til Kundskaben om arktiske Planters 
Bladbygning. (Botanisk Tidsskrift. Kjøbenhavn. Bd. 19. — 
Resumé frangais: Journal de botanique. Vol. IX. 1895). 

CLEVE, ASTRID, 1901: Zum Pflanzenleben in Nordschwedischen Hoch- 


gebirgen — Einige åkologische und phånologische Beitråge. 
(Bihang till K. Svenska Vet.-Akad. handlingar. Bd. 26. Afd. 
INFANOES): 


Dusen, P., 1901: Zur Kenntnis der Gefåsspflanzen Ostgrånlands. 
(Bihang till K. Svenska Vet.-Akad. handlingar. Bd. 27. Afd. 
III. No. 3). 

EAsTWO0OoD, ÅLICE, 1902: A Descriptive List of the Plants collected 
by Dr. F. E. Blaisdell at Nome City, Alaska. (Botanical Gazette. 
Vol. 33). 

EKgSsSTAM, O., 1897: Einige blutenbiologische Beobachtungen auf Novaja 
Semlja. (Tromsø Museums Årshefter. 18. 1895). 

— , 1899: Einige blutenbiologische Beobachtungen auf Spitz- 

bergen. (Tromsø Museums Årshefter. 20. 1897). 

FREIDENFELT, 1904: Der anatomische Bau der Wurzel in seinem Zu- 
sammenhange mit dem Wassergehalte des Bodens. (Bibliotheca 
botanica. 61. Stuttgart). 

GoEBEL, K., 1897: Morphologische und biologische Bemerkungen. 
7. Ueber die biologisehe Bedeutung der Blatthohlen bei Tozzia 
und Lathraea. (Flora. Bd. 83). 


364 Er. J. MATHIESEN. 


Groom, Percy, 1897: On the Leaves of Lathraea squamaria and of 
some allied Scrophulariaceae. (Annals of Botany. Vol. XI). 

HABERLANDT, G., 1897: Zur Kenntniss der Hydatoden. (Pringsheims 
Jahrbucher. Bd. 30). 

Hartz, N., 1894: Botanisk Rejseberetning fra Vest-Grønland 1889— 
1890. (Medd. om Grønland. 15. Hefte. Kjbh. 1898). 
— , 1895 (I): Østgrønlands Vegetationsforhold. (Ibidem. 18. 
Hefte. Kjbh. 1896). 
— , 1895 (II): Fanerogamer og Karkryptogamer fra Nordøst- 
Grønland, c. 75—70? N. Br., og Angmagsalik, c. 657407 N. Br. 
(bide met FE etter ED GE 
— and Car. Kruuse, 1911: The Vegetation of Northeast Green- 
land, 695255 lat; n:——759 lat: mn, H(Ibidem "307 Fefte Ko HOLLY 
HEINRICHER, H., 1898—1910: Die grunen Halbschmarotzer. I—VI. 
(PringsheimsfJaåbrbucher BARS ES GESTA GS FAR 
HOLLSTEIN, O., 1907: Beitråge zur vergleichenden Anatomie der Stengel 
und Rhizome von dicotylen Alpenpflanzen. Dissertation. Goått- 
ingen. 

Horm, TxH., 1885: Novaia-Zemlia's Vegetation, særligt dens Phane- 
rogamer. (Dijmphna-Togtets zoologisk-botaniske Udbytte. Kjbh. 
1885). 

HoveLacQuE, M., 1888: Recherches sur Vappareil végétatif des Big- 
noniacées, Rhinanthacées,-: Orobanchées et Utriculariées. Paris. 

HucHeDe, 1907: Veronique et Gratiole: Thése. Paris. 

JUNGNER, J. R., 1894: Klima und Blatt in der Regio alpina. (Flora. 
Bd%/9)' 

JØRGENSEN, E., 1919: Die Euphrasia-Arten Norwegens. (Bergens 
Museums Årbok. 1916—17. Naturvidenskab. række. No. 5). 

KERNER v. MARILAUN, Å., 1898—1900: Pflanzenleben. Bd. I—II. 
Leipzig u. Wien. 

KserLman, F. R., 1882 (I): Sibiriska nordkustens fanerogamflora. 
(Vega-expeditionens vetenskapliga iakttagelser. Bd. I. Stock- 


holm). 
— , 1882(II): Fanerogamfloran på Novaja-Semlja och Wajgatsch. 
(Ibidem). 


— , 1882 (III): Om tschuktschernas hushållsvåxter. (Ibidem). 

— , 1882 (IV): Asiatiska Beringssunds-kustens fanerogamflora. 
(Ibidem). 

— , 1883: Ur polarvåxternas lif. (Nordenskidld: Studier och 
forskningar foranledda af mina resor i håga norden. VII. Stock- 
holm). 

— and A. N.Lunnstrom, 1882: Fanerogamer från Novaja-Semlja, 
Wajgatsch och Chabarova. (Vega-expeditionens vetenskapliga 
iakttagelser. Bd. I. Stockholm). 

Knurtix, P., 1899: Handbuch der Blutenbiologie. Bd. II, Teil 2: Lobe- 
liaceae bis Gnetaceae. Leipzig. 


Scrophulariaceae. 365 


Kocx, E., 1895: Uber die systematische Bedeutung der anatomischen 
Charaktere der Scrophulariaceen. Dissertation. Erlangen. 
KRUUSE, CHR., 1898: Vegetationen i Egedesminde Skjærgård. Medd. 

(om Grønland. 14. Hefte. Kjbh. 1898). 

— , 1905: List of the Phanerogams and Vascular Cryptogams 
found on. the coast, 757—66720” lat. N. of- East Greenland. 
(Ibidem. 30. Hefte.. Kjbh. 1911). 

— , 1906: List of Phanerogams and Vascular Cryptogams found 
in the Angmagsalik District on the East coast of Greenland 
between 65730” and 66220” lat. N. (Ibidem). 

— ,1911: Rejser og botaniske Undersøgelser i Øst-Grønland mellem 
657307 og 67220” i Årene 1898—1902 samt Angmagsalik-Egnens 
Vegetation. (Medd. om Grønland. Bd. XLIX. Kjbh. 1912). 

LANGE, J., 1871: Bemærkninger om. frøenes form og skulptur hos 
beslægtede arter i forskellige slægter. (Botanisk Tidsskrift. 
Kjbh. Bd. 4). 

— , 1880: Conspectus Florae Groenlandicae. (Medd. om Grøn- 
land. 3. Hefte. Kjbh. 1880). 

— , 1887: Tillæg til Fanerogamerne og Karsporeplanterne i Consp. 
Flor. Groenl. (Ibidem. 3. Hefte. Fortsættelse. Kjbh. 1887). 
Leist, 1889: Ueber den Einfluss des alpinen Standortes auf die Aus- 
bildung der Laubblåtter. (Mitth. der naturforsch. Gesellschaft 

von Bern). 

Linndnman, C. A. M., 1887: Bidrag till kånnedomen om skandinaviska 
fjellvåxternas blomning och befruktning. (Bihang till K. Sven- 
ska Vet.-Akad. handlingar. Bd. 12. Afd. III. No. 6). 

LUNDAGER, Å., 1912: Some Notes concerning the Vegetation of Ger- 
mania Land, North-East Greenland. (Medd. om Grønland. 
BUSSEN EEGDRETOEJ 

MULLER, H., 1881: Alpenblumen, ihre Befruchtung durch Insekten 
und ihre Anpassungen an dieselben. Leipzig. 

NATHORST, Å. G., 1883: Nya bidrag till kånnedomen om Spetsbergens 
kårlvåxter och dess våxtgeografiska fårhållanden. (K. Svenska 
Vet.-Akad. handlingar. Bd. 20. No. 6). 

NORMAN, J. M., 1895: Norges arktiske Flora. II. Kristiania. 

OstENFELD, C. H., 1901: '""Phanerogamae and Pteridophyta” in 
<Botany of the Færåes”. Vol. I, Kjbh. 

— , 1908: The Land-Vegetation of the Færåes. (Ibidem. Vol. ILI). 

— , 1915: Plants collected during the First Thule Expedition 
to Northernmost Greenland. (Medd. om Grønland. Bd. LI. 
Kbh ToTS) 

— and AÅ. LUNDAGER, 1910: List of Vascular Plants from North- 
East Greenland (N. of 769N. lat.), collected by the Danmark-Ex- 
pedition 1906—1908. (Medd. om Grønland. Bd. XLIII. Kjbh. 
ILSALDE 

SEKSVEE 24 


366 Fr. J. MATHIESEN. 


Porrius, B. R., 1903: Blombiologiska iakttagelser. (Acta soc. pro 
fauna et flora fennica. 25). 

Porsirn, M. P., 1902: Bidrag til en Skildring af Vegetationen på Øen 
Disco etc. (Medd. om Grønland. 25. Hefte. Kjbh. 1902). 

— , 1910: The Plant-Life of Hare-Island off the coast of West- 
Greenland. (Ibidem. Bd. XLVII. Kjbh. 1911). 

— , 1912: Vascular Plants of West-Greenland between 71% and 
dos Ny lats (Ibidem BAK EL 92) , 

— , assisted by A. ERLING Porsirn, 1920: The Flora of Disco 
Island and the adjacent coast of West-Greenland from 66%— 
TASÆNEMatteteree(Ibidem Bb ME VISES DD 

RAUNKIÆR, C., 1907: Planterigets Livsformer og deres Betydning for 
Geografien. Kjbh. 

RESVOLL, THEKLA R., 1917: Om planter som passer til kort og kold 
sommer. (Archiv for mathematik og naturvidenskab. Bd. XXXV. 
Kristiania). 

RosENVINGE, L. KorDnerur, 1892: Andet Tillæg til Grønlands Fanero- 
gamer og Karsporeplanter. (Medd. om Grønland. 3. Hefte. 
Fortsættelse. Kjbh. 1887). 

— , 1896 (I): Nye Bidrag til Vest-Grønlands Flora. (Ibidem. 
lime er te Ek DDS 98] 

— , 1896 (II): Det sydligste Grønlands Vegetation. (Ibidem 15. 

: Hefte. Kjbh. 1898). 

SCHERFFEL, Å., 1888: Die Drusen in den Håhlen der Rhizomschuppen von 
Lathraea squamaria L. (Mitth. a. d. Bot. Inst. zu Graz. V. 1888. 
Jena). 

ScHROTER, C., 1908: Das Pflanzenleben der Alpen. Zurich. 

SILEN, F., 1905: Blombiologiska iakttagelser i Kittila Lappmark. 
(Medd. af soc. pro fauna et flora fennica. 31). 

Simmons, H. G., 1906: The Vascular Plants in the Flora of Elles- 
mereland. (Report of the second Norwegian Arctic Expedition 
in the "Fram”, 1898—1902. No. 2. Kristiania). 

— , 1913: A Survey of the Phytogeography of the Arctic American 
Archipelago, etc. (Lunds Universitets Årsskrift. N.F. "Afd. 2. 
BdSÆNOST9)' 

SKOTTSBERG, C., 1901: Einige blutenbiologische Beobachtungen im 
ark. Teil von schwedisch Lappland 1900. (Bihang till K. Svenska 
Vet.-Akad. handlingar. Bd. 27. Afd. III. No. 2). 

SyYLveEn, N., 1906: Om de svenska dicotyledonernas fårsta forstårk- 
ningsstadium etc. I—II. (K. Svenska Vet.-Akad. handlingar. 
Bd. 40. No. 2). 

WAGNER, Å., 1892: Zur Kenntniss des Blattbaues der Alpenpflanzen 
und dessen biologischer Bedeutung. (Sitzungsber. der kaiserl. 
Akad. der Wissenschaften in Wien. Matem.-naturw. Classe. Bd. 
CIFADEDET)! 


Scrophulariaceae. 367 


WARMING, EuG., 1884: Om Skudbygning, Overvintring og Foryngelse. 
(Naturhistorisk Forenings Festskrift. Kjbh.). 
— , 1886: Om Bygningen og den formodede Bestøvningsmåde af 
nogle grønlandske Blomster. (Oversigt o. d. Kgl. Danske Vid. 
Selsk. Forhandl. Kjbh.). 
— , 1888: Om Grønlands Vegetation. (Medd. om Grønland. 12. 
Hefte. Kjbh. 1888). 
— , 1890: Biologiske Optegnelser om grønlandske Planter. 3. Scro- 
phulariaceae. (Botanisk Tidsskrift. Kjbh. Bd. 17). 
R.v. WETTSTEIN, R., 1896: Monographie der Gattung Euphrasia. Leipzig. 
VOLKART, 1899: Untersuchungen uber den Parasitismus der Pedicu- 
laris-Arten. Dissertation. Zurich. 


368 Fr. J. MATHIESEN. 


Veronica fruticans Crantz (V. saxatilis Scop.). 


Alcohol-material from Greenland (Julianehaab, leg. Litrt- 
ZEN, 3. 7. 1887; Præstefjældet, leg. E. WARMING, 2. 8. and 
6. 8. 1884) and Northern Norway (Kåfjord, leg. E. WARMING, 
15. 7. 1885). — Herbarium-material from Greenland, Iceland, 
the Færåes and Fennoscandia. 

rt SPAS EET LEG POP EAN GENE ES OPUS EET EP RE GE 
MULLERULSSLMpE 2 67 WARMING 1688 pp. 30 4 and ks 790) 
p:203-RHO0SENVINGE HEL 92 ED 160859 GT Ep pE S EAN 
WAGNER, 1892, pp.8 and 21; Hartz, 1894, pp. 9 and 57; 1895 (I), 
pp: 273,;289;11895(I1),"p:7335; HARTZ" and KRUUSE, 191 Eppy352) 
357, 359 and 409; Kocxu, 1895, p.117 et seq. and p. 127; NoRrRMAnN, 
1.895 Ep FAP EK Nora ONE pE GE Pors VO ED 2302 
PESH ORE DENE OD TP MØN ES AVE and he VER G2- 
SyæVvEN 900 Ep EO KRUUSE ELDO SEP LD 90 GE p SP AS EOS RE 
part IV pp. 202, 207, 243, 261 and 262, besides many notes in the 
preceding parts; ScHROTER, 1908, pp. 221 and 656. 

Ån evergreen, nanophyllous?, sympodial under-shrub- 
chamæphyte which has a main root that dies away early 
(according to SYLVEN), and develops adventitious roots 
fairly abundantly; the winter-buds are raised at most a 
few cm above the surface of the ground; older, vigorous 
specimens form small tufts (Kruuse, 1906, p. 248). The 
shoot-development extends over two years; during the 
first year a horizontal or obliquely-ascending part is de- 
veloped, only a few cm long and furnished with a few 


(3—7) pairs of foliage-leaves; the next year the shoot 
1 C. RAUNKIÆR: Om Bladstørrelsens Anvendelse i den biologiske 
Plantegeografi (Bot. Tidsskrift, Kjbh., Bd. 34, 1916). 


Scrophulariaceae. 369 


usually continues its growth in a vertical direction, larger 
leaves are developed, and the growth can be terminated by 
a few-flowered raceme without a terminal flower. Special 
bud-scales do not occur; during the winter-rest the point of 
the axis is protected by the uppermost, not yet expanded pair 
of foliage-leaves, which are in con- 
tact with each other by their hairy 
margins. After the fruit hasripened, 
the stem dies as far down as to some- 
what above the boundary line be- 
tween the lst and 2nd year's growth ; 
the perennial basal portions bear 
the innovation shoots. Principal 
buds proper do not occur, but the 
uppermost buds appear generally 
to be the most advanced, and it 
is evidently especially these buds 
which produce the flower-bearing 
axes, while the lower ones often 
produce only small, weak, few- 





leaved shoots which — as recorded 

by Wammunc (1890, p. 203) and (8 Free tiger 
as also shown in Fig.l — may  2.8.1884). (About nat. size.) 
be somewhat runner-like and fur- 

nished with only a few small leaves. The specimen illustrated 
in Fig. 1 is rather scantily branched; each of the floral shoots 
has only two real ”innovation-buds”, and of their parent- 
shoots the one to the left has also had two, of which one 
has developed into a vegetative shoot; the parent-shoot to 
the right is somewhat more richly equipped: in addition to 
the lowermost quite small shoot it bears two opposite floral 
shoots, and in the axils of the next pair of leaves two more 
shoots, of which the one (cut-off) was floral. Such a difference 


370 Fr. J. MATHIESEN. 


between shoots from two opposite leaves, that the one 
becomes vigorous and floral, and the other vegetative only, 
is very common; it appears to be the rule that the shoot 
which turns outwards towards the periphery of the tuft, 
becomes more vigorous. It happens not rarely, however, that 
the short, vegetative shoots are later on instrumental in the 
formation of floral shoots, in that the latter arise as lateral 
shoots upon them; in such a case the part of the parent- 
shoot above these lateral shoots dies away, as is the case 
in the floral shoot. The development of the lateral shoots 
usually takes place in the second year of the parent-shoot, 
and consequently simultaneously with the flowering. 

The Leaves are rather thick, entire or slightly serrate; 
on the median rib and along the margin there is a sparse 
covering of non-glandular hairs; such are also found on the 
stem and in the floral region, viz.,-on the calyx and ovary. 
In the Botanical Garden in Copenhagen the leaves on the 
new innovation-shoots, developed during the last summer, 
remain green and fresh during the winter; on the flowering 
shoots, in my herbarium-material, the leaves from the previ- 
ous year were withered in some cases; most often, how- 
ever, they were green. 

As a rule, the basal portion of the shoots takes root 
abundantly, and is drawn down by the roots to the surface 
of the ground; in the individual illustrated in Fig. 1, which 
has a somewhat more erect growth than is generally the 
case, adventitious roots are only scantily produced; the roots 
are developed at the earliest in the second year of the shoots. 

The basal portions of the shoots have growth in thick- 
ness and may live several years; an individual from Greenland 
(Ikalik) had a prostrate stock, 4mm thick, which showed 
16 annual rings. 

The Flower-biology has been investigated by H.Mir- 


Scrophulariaceae. Bl 


LER (the Alps) and E. WARMING (Greenland and Northern 
Norway). The flowers are large and conspicuous; also in the 
Årctic regions, as a rule, they are of a pure and bright colour; 
but according to M. Porsirn, in northern Greenland a white- 
flowering variety may commonly occur. The ultimate dia- 
meter of the corolla is very constant, being 10—12 mm in 
material from the different localities; I found it to be parti- 
cularly large in some material from Iceland, viz. 14 mm (in 
the Alps, according to Muller, only 6—7 mm). ""The corolla- 
tube and the throat is white, and furnished with a wreath 
of hairs; then follows a dark, brown or reddish ring and 
then the deep-blue limb. The corolla-tube is 17/,—21/, mm 
in length (according to Muller 3 mm). As in the Alps the 
filaments taper very much at the base, are about 3 mm in 
length and, as the pistil is of the same length, the stigma 
and anthers stand at the same level. The flowers are homo- 
gamous. In fully expanded flowers self-pollination appears 
to be able to take place only with difficulty, as the stamens 
diverge so much laterally and the anthers are thereby re- 
moved from the stigma on the straight outstretched style; 
but self-pollination might have taken place at an earlier 
point of time, for I have seen. a flower (from Greenland at 
67? N. lat., Aug. 6) which, perhaps on account of gloomy 
and rainy weather, was only slightly expanded and in which 
the one anther was open, so that the pollen fell out of it 
down upon the stigma which stood close to it and which 
appeared to be fully ripe. The ovary is covered with small 
upwardly directed, adpressed hairs, and the style is 2—3 
times longer than the ovary. Ripe fruit is produced in West 
Greenland at leastupto 709 N. lat.” (E. WarmiInG, 1890, p. 203). 

V. fruticans is homogamous also according to ÅAXELL 
and H. Mirer. The style, however, appears to me to be 
always slightly longer than the anthers; the outspread posi- 


< 
+ 


72 Fr. J. MATHIESEN. 


JO 


tion of the latter is distinetly seen in Fig. 5, C; the figure 
is drawn from living material from the Botanical Garden in 
Copenhagen. 

According to M. Porsirn, near its northern limit in Green- 
land it flowers late and even sets fruit; but it is only in 
favourable, dry autumns, without too much frost, that its 
seeds ripen. Unripe fruits live through the winter, but perish 
without developing further. The structure of the shoots ex- 
plains, as in V. alpina (TH. ResvorLt), the late flowering. 
There was, however, ripe fruit to hand from all localities. 
KRUUSE (1906, p. 248) in the Angmagsalik-district notes 
abundant ripe fruit”. 

Geographical Distribution according to LANGE: 
Greenland, the Ural Mountains, Lapland, Fmmark, Norway, 
Iceland, Great Britain, the Alps and the Pyrenees. Besides 
this it is recorded from the Færåes. Its certainly known north 
limit in West Greenland is 70717' on the continent (Vaj- 
gattet), from 70%—67” it occurs in isolated specimens only, 
and not until further southwards is it common; consequently, 
in Greenland it belongs to the southern types (M. PorsiILn). 
In East Greenland it occurs as far north as the Scoresby- 
Sound-district (Hartz and KRUUSE, 1911). 

According to WARMING (1888) and ROsENVINGE (1896 
(II)) im West Greenland the species grows in willow-copses, 
birch-copses and on "herb-slopes”, and in places whence 
snow melts early it can ascend as high up in the moun- 
tains as 750 metres (LANGE, 1880: in grass-covered, open 
places, in rocky clefts). M. Porsirn informs me in con- 
firmation of this that its natural habitat is the edge of the 
willow-copse, there it grows on warm, sunny slopes which 
are not too dry; the species thrives but badly when over- 
shadowed; it needs snow-covering throughout the winter 
but must be early freed from snow; at its north limit it 


-= 


Scrophulariaceae. 315 


does not at any rate belong to the real snow-flora. In the 
Scoresby-Sound-district it is found "in particularly well 
sheltered luxuriant, humid, herby slopes with high snow- 
cover in winter,” but here it is very rare (Kruuse, 1905, 
p. 175). In the Angmagsalik district, where it is somewhat 
more common (Kruuse, 1906, p. 248), besides growing on 
the herby-slopes, it is 
also noted from the 
"steps of steep rocks 
above the slopes.” Ac- 
cording to NORMAN, 
in Northern Norway 
the species is found 





on all kinds of stony 


(7% 










let 


set 


substrata rather than 










on grassy ground, and 5 
åt very much prefers OSSE NSSS 
3 « > DD 

the sunny side, especi- SR 8 






ø 
OQ 
E 


å : SER st UW 
ally the side facing ROOS 
directly south) and SER SES SØEN 











occurs only very rarely 


<Q 
ON 
> 

(DN 








on the indifferent 
Fig. 2. Veronica fruticans. 

SE HR Transverse section of an adventitious root 

sides”. In the Alps (about 239/,). (Greenland.) 

it is recorded from 


"Schneeblåssen” (ScHROTER, p. 656), spots where the wind 


(eastern and western) 


sweeps the snow away in winter time. 

Anatomy. The Root. The epidermis of the adventi- 
tious roots dies away early; the outermost layer of the cortex 
is developed as an exodermis with cuticularised walls. Even 
before any secondary growth has taken place in the stele, 
a cork-cambium is developed in the layer under the exo- 
dermis, which is instrumental in the formation of a few- 


314 Fr. J. MATHIESEN. 


layered cork (cf. Fig. 2). The walls of the primary cortex 
attain a rather considerable thickness; this is especially the 
case as regards the endodermis; both in this and in the 









Q w O 
CH 
HU See] 





Fig. 3. Veronica fruticans. 
Transverse section of the stem (about 299/,).  (Greenland.) 


other layers, radial walls are formed during the secondary 
growth of the root. 

In the wood of the 2-year-old root illustrated in Fig. 2, 
an annual ring is formed outside the central part which was 
developed during the first year; the limit between them is 


Scrophulariaceae. 375 


easily discernable, since they are separated by a zone in 
which the vessels and wood-fibres are intermixed with non- 
lignified, axially elongated parenchymatous elements. 

The Stem. In the lower persistent parts of the stems, 
a cork-cambium commences activity during their second 
year; it appears in the outermost layer of the cortex and 
forms a few-layered cork (Fig. 3). The cells of the primary 
cortex are rather thick-walled and show division by radial 
walls; they contain chlorophyll-grains. The endodermis has 
very distinct Casparian dots (as in the root). In the 
pericycle small groups of hard-bast cells are found. The 
figure shows a portion of the transverse section of a stem 
with apparently two growth-rings; between the two rings 
consisting of vessels and wood-fibres there is a zone inter- 
mixed with thin-walled and non-lignified cells; however, in 
reality the stem is 3 years old; during the first year only 
the innermøost zone of the xylem with the small scattered 
vessels being developed. According to whether the shoots 
during the first year succeed in becoming more or less 
vigorous, so also does the thickness of the xylem of the 
first year vary; shoots may sometimes be found in which 
a continuous wood-ring has been developed as in the follow- 
ing years. 

In the upper part of the shoot which dies away, no 
cork-formation takes place, the cortical cells are less thick- 
walled than in the lower part and richer in chlorophyll and 
Casparian dots are less distinctly developed. Hard bast is 
wanting or is scantily present in the pericycle, and the wood- 
ring is quite narrow. From all the axial organs medullary 
rays are quite absent. 

The Anatomy of the Leaf has been investigated by 
KocH and HucHEenk; my investigations entirely bear out 
the conclusions arrived at by them. The epidermal cells of 


376 i Fr. J. MATHIESEN. 


both the upper and lower surface have undulating lateral 
walls, those of the epidermal cells of the lower surface are 
however more strongly undulating; under a higher magni- 





Fig. 4. Veronica fruticans. 
A, Epidermis of the upper, and B of the lower surface of; the leaf. 
C and D, The uppermost and lowermost layer of the mesophyll respec- 
tively, shown in surface view. E, Epidermal cells from the upper sur- 
face of the leaf, more highly magnified. F, Å non-glandular hair from 
the mid-rib. G, Transverse section of the leaf (A and B about ?%,; 
CEED G abo SEERE about Greenland) 


fication the lateral walls are seen to be as shown in Fig. 
4, E: pores and nodose thickenings alternate. The outer walls 
have fine cuticular striations. Stomata are almost equally 


Scrophulariaceae. BMET 


distributed on both surfaces of the leaf, I found them how- 
ever (contrary to Wagner's statement) to be slightly in the 
majority on the. lower surface; the absolute number per 
square unit proved, however, to vary somewhat, but I did 
not succeed in finding any fixed rule as regards this point. 
The stomata are on a level with the surface of the leaf. 

One of the non-glandular hairs occurring on the leaf- 
margin and the under-side of the midrib is shown in F ig.4,F; 
they are 2—4 celled, thick-walled and have cuticular warts 
on the surface.. Glandular hairs with a one-celled stalk and 
two-celled head occur in great numbers and are equally 
abundant both on the upper and the lower surface of the 
leaf (Fig. 4, A and B). A transverse section discloses 2—3 
layers of short and broad palisade-cells with rather large 
intercellular spaces; the spongy parenchyma consists of ovate 
to slightly branched cells; all the cells of the mesophyll are 
abundantly filled with chlorophyll-grains. Palisade tissue and 
spongy parenchyma are shown in surface view in Fig. 4, 
C and D. The mid-rib has on its under-side a thin layer 
of stereom. 


Veronica alpina L. 


Alcohol-material from Norway (Troms&, leg. E.WARMING, 
21. 7. 1885; Muggrubskampen, Rérås, leg. TH. RESsVOLL, 
29. 7. 1918) and Greenland (Nunatsuk, 11. 8. 1885; Ivigtut 
and Dronning Louises Ø, leg. P. EBERLIN, 21. 8. 1883 and 
8. 8. 1885). — Herbarium-material from Greenland, Iceland, 
the Færåes and Fennoscandia. 

Jr ASEE BESES OS OP SANGE SS OD VORES SUEDE 2 
MULLER, 1881, p. 270; Linnman, 1887, p. 81; WARMING, 1888, pp. 31, 
35, 39, 75, 87, 93 and 142; 1890, p. 204; RosENvInNGE, 1892, p. 685; 
1896 (II), pp. 128, 161, 168; WAGNER, 1892, pp. 9 and 20; HArTz, 1894, 
pp. 9, 49, 50 and 57; 1895 (I), pp. 137, 170, 179, 266, 271, 288 and 304; 


1895 (II), p. 335; HArTtz and Kruuse, 1911, pp. 346, 359, 364, 409, 417 
and 423; JuNGNER, 1894, p. 275; KocxH, 1895, pp. 117 et seq. and 


378 Fr. J. MATHIESEN. 


128; NorMaAnN, 1895, p. 452; KERNER, 1898, p. 350; Knurx, 1899, p. 
171: Greve, 190LSpp-12 16; 25540 57 Fandt so) Ras woo EO? 
pp 292" PORSILD I 1902 pp TORE Fan de 209 EON ONE DEPOT OD 
p:140: KRUUSE, 19057 pp 1705 900 Sp PAS ELSE par EVE PPS 
196, 202, 229, 230, 242, 247, 255, 261, 262, besides many notes in 
the preceding parts; SyLvEn, 1906, p. 80; Hucxaende, 1907, p. 73, fig. 
IV AS fr VEG SCHROTER, 1908 pp 221226 1468 kande 493 Er 
RESVOLL, 1917, p. 208: 

Nanophyllous sympodial chamæphyte with primary root 
which dies away early (SYLVEN), and abundant development 
of adventitious roots from the stem-hbases, this in conjunction 
with the fact that the older shoots gradually die away, de- 
termines the vegetative reproduction, enabling the plants 
to form lax, but large tufts (Kruuse, 1906, p. 248). The 
winter-buds either rest upon, or are slightly raised above, 
the surface of the ground. 

The Shoot-development has been described by TH. 
RESVOLL; it is a process of two years duration, as in the 
foregoing species; the first-year's part of the shoots is either 
erect or obliquely ascending or quite horizontal, as in TH. 
ResvorLL's Figs. 60 and 61; most often only 1—2 cm long and 
bearing a few small leaves. The next year the growth of the 
shoot is continued in a vertical direction, while essentially 
larger leaves are being formed; the shoot is frequently ter- 
minated by an inflorescence; the flowers are already formed 
during the autumn of the first year (1. c.). Special bud- 
scales do not occur, the end of the axis is only pro- 
tected by the uppermost pair of leaves. After the fruit has 
ripened, the axis dies away to slightly above the "innova- 
tion-buds”; the latter occur, however, often rather far down 
on the shoots, even in the axils of the very first pair of 
leaves, which causes the branches of the sympodia to be- 
come very short, and the stems crowded. 

Of the two shoots in the axils of two opposite leaves, 
the one may be far more vigorously developed than the 


Scrophulariaceae. 319 


other; this is no doubt usually the case, and then it appears 
to be the shoot which is turned towards the periphery of 
the tuft which becomes the more vigorous and floriferous. 
When the plant grows in damp moss, the internodes of the 
horizontal, first-year portions of the shoots — as mentioned 
by TH. REesvorL, and as I myself had an opportunity of 
verifying in the mountains of Norway — may become elon- 
gated, so that the shoots become almost runner-like; in this 
case the plant is capable of spreading considerably. 

The longevity of the branches of the sympodium is 
greatly restricted; a growth in thickness of the axial organs, 
continuous for years as in V. fruticans, does not take place. 

Adventitious Roots are developed in the second 
growth-period of the shoots, they arise in the neighbourhood 
of the nodes. The foliage-leaves are either entire or slightly ser- 
rate; I am not prepared to say whether the leaves occurring 
at the base of the shoots remain green throughout the winter, 
in the following summer they are at any rate always found 
im a withered condition. 

The Flower. H. Mircer, Linnman, E. WARMING and 
KERNER have in the works cited above described the structure 
and biology of the flower, which according to these authors 
agree in the Alps, Scandinavia and Greenland, nor have I 
been able to find any differing features. "The small, dark blue 
flowers are at first only 2.5—3 mm in diameter, but may 
ultimately become 5—5.5 mm. They are protogynous-homo- 
gamous, and appear to be well-adapted to self-pollimation. 
While the corolla is still almost tubular or funnel-shaped 
and consequently only slightly open, the anthers may be 
open and lie close to the stigma, which may be seen to be 
covered with pollen-grains, many of which are germinating. 
Afterwards the anthers are slightly removed from the stigma 
by the filaments bending backwards, but not so decidedly 


380 Fr. J. MATHIESEN. 


as in V. saxatilis; the anthers are, indeed, not far from 
remaining parallel with the style, and are therefore con- 
stantly near the stigma”” (WAarmiInG, 1890). Tx. REsvoLL 
records self-pollination (at Rorås, in sched.); Fig. 5, A and 
B show that this may easily take place; moreover, in 
scarcely expanded flow- 
ers I too found the 
anthers open and the 
stigma covered with 
pollen. 

The ovary varies 
between the glabrous 
and  finely-hairy con- 
dition. 

LINDMAN remarks 
that the flowers open 
two at a time. They 


are, however, on the 





whole, not very con- 


Fig. 5. 4, and B, Flowers of Veronica 

alpina (Rorås, Norway, ”/,): A, In front- ie 

view; B, in side-view, half of the calyx — Visits are scarce. 

and of the corolla is removed. (About By reason of the 

5/,). C, A flower of Veronica fruticans. 
(Hort. bot. Hauniens.) (About ”/,.) 


spicuous, and insect- 


fact that there is no 
division of labour be- 
tween the vegetative and purely floral shoots, the flowering. 
period occurs late (Tx. ResvorL). In Greenland V. alpina 
sets fruit abundantly, and ripens its seeds normally; the 
latter are scattered in autumn; wherever in Greenland V. 
fruticans and V. alpina occur in the same locality, the 
latter is always the first to flower (M. PorsiLn). 
Geographical Distribution according to LANGE: 
East and west coasts of Greenland, Labrador, the Rocky 
Mountains, western North America, Siberia, Arctic Russia, 


Scrophulariaceae. 381 


Scandinavia to Lapland and Finmark, Iceland, Great Britain 
and the Alpine regions of Southern Europe. To these should 
be added the Færåes (OstENFELD, 1901, p. 56). In West 
Greenland the northernmost limit is found on the Nugsuak- 
peninsula (70742'); in 
East Greenland it 
passes over the Scores- 
by Sound. 

Habitat. In West 
Greenland V. alpina 
grows in copses, on 
"herb-slopes”,insnow- 

troughs (Snelejer) 
where the snow does 






not lie till very late 
in the summer, and 






ØS 


on sandy flats near (Å eQ. 
the shore (WARMING, KR S 

















(l) 

[> 

g) 
FISG IN 

mn 


as 
Sy 
BO 
Sta 


OGS Y 
1888; ROSENVINGE, BY == 
<S ES 
1896 (II)). In East — | SEE Er 
Greenland it is noted TT Å 
too from "herb- Fig. 6. Veronica alpina. 
slopes”, grassy-slopes Transverse section of an adventitious root 


y about 239/,). (Norway. 
and copses; besides, DRM | 


KRUUSE, in the Angmagsalik district, has found it growing 
on the steps of steep rocks above the slopes. It is always 
well covered with snow during its winter-rest (cf. also 
KRUUSE, 1905, p. 175 and Porsirn, 1920, p. 140). TH. REes- 
VOLL mentions the species as a common plant of the snow- 
troughs in the whole of Norway; according to NORMAN it is 
also found by mountain-streams, in birch-wood glades, on 
inundated river-banks, on the coast and on mountain-sum- 
mits. Å. CLEVE records: ”"Håufig und uppig ausgebildet in 
XXXVII. 25 


382 Fr. J. MATHIESEN. 


Sumpfwiesen, . . . Ferner håufig in der Moosmatte, dort 
noch mit Bluiten und Knospen am &/, und in rein vegetat. 
Individ. nåher an dem ewigen Schnee als irgend eine 
andere Blutenpflanze beobachtet.  Meidet dagegen ent- 
schieden trockene Hei- 
(87 den und Wiesen”. Tx. 
beses RESVOLL, also, states 
FO XZ Bee that, in unfavourable 
KODER SE localities, where the 
DEN EN ÅR snow lies till late mm 
AIF ie the summer, V. alpina 
2 en occurs as sterile speci- 

mens. 

Im the: Alps; as 
recorded by SCHROTER, 
the species is "einer 
der verbreitetsten Ra- 
senbestandteille == von 
der subalpinen bis zur 
nivalen Region (1500— 
3185 m in der Schweiz, 
1560—2870 m in Bay- 
ern) alle Bodenarten 





Fig. 7. Veronica alpina. bewohnend, indifferent 
Transverse section of a stem (about ?39/,). gegen Dungung und 
RER Humusgehalt”.  Be- 
sides this, it occurs in snow-troughs (Schneethålchen). 
Anatomy. The Root. The epidermis becomes cuti- 
cularised and persists a long time. The outermost layer of 
the cortex consists of thin-walled cells with cuticularised 
outer and lateral walls; the two succeeding layers within 
the cortex (Fig. 6), have, on the other hand, fairly thick- 
walled cells which unite mutually and with the exodermis 


Scrophulariaceae. 383 


without intercellular spaces; the walls show the reaction of 
cellulose. During the growth in thickness of the stele, the 
endodermal cells are stretched tangentially, and become sub- 
divided by 2—4 thin radial walls; the walls of the original 
endodermal cells show Casparian dots, but only faintly. The 
cortical cells contain starch; root-hairs occur in great num- 
bers. In the xylem portion I have not found indication 
of annual rings (the root figured is several years old) as in 
the foregoing species; consequently, the roots complete their 
development in the course of one year. The xylem is 
composed of vessels and wood-fibres. 

The Stem. As in V. saxatilis the xylem, in the first 
year of growth, when the shoots are but weak, frequently 
consists only of scattered vessels with intervening non- 
lignified parenchyma, and not until the second year is a 
continuous ring of vessels and wood-fibres formed, as shown 
in Fig. 7, which shows a portidn of a transverse section of 
the basal, persistent part of the shoot. I never found more 
than 3 years” growth in thickness, not even in shoot-bases 
apparently older. The vessels are comparatively numerous, 
and the wood-fibres are more sparingly present and also 
more thin-walled than in the foregoing species. In the cells 
of the long-lived epidermis tannin is found; cork-formation 
does not take place, neither in the stem nor in the root. 
The cells of the cortex contain chlorophyll-grains. The young 
shoots are beset with glandular hairs of the same type as 
those on the leaf. In the upper part of the stem, which dies 
away after flowering, the wood-ring is of course formed 
entirely during one growth-period, here it is thinner than 
im the lower part; the cortical cells are thinner-walled and 
richer in chlorophyll, and the epidermis contains no tannin. 

The Leaf. The epidermal cells of the upper surface 


have slightly, those of the lower surface more highly, un- 
25% 


384 Fr. J. MATHIESEN. 


dulating lateral walls; they are thin and without pores (Fig 
8, D). Stomata occur almost equally on both sides of the 
leaf, frequently they are slightly in the majority on the 





















erMe 


(7 






ore OLE :: | 
rd Va Y) e E i 
SODE 

2 LY 









Fig. 8. Veronica alpina. 
A, Epidermis of the lower surface of the leaf. B, Epidermis of the 
upper surface of the leaf. C, A non-glandular hair from the leaf-margin. 
D, An epidermal cell more highly magnified. E and F, The upper 
and lower layers respectively of the mesophyll, in surface view. G, 
Transverse section of the leaf (A and B about ?9%,; C, E, F and G 
about 115/,; D about ?75/,). (Greenland, Dronning Louises Ø, 8. 8. 1885.) 


upper surface; the stomata are on a level with the surface 
of the leaf. Both leaf-surfaces are furnished with glandular 
hairs of the usual type — a two-celled head on a one- 


Scrophulariaceae. 385 


celled stalk; non-glandular hairs (Fig. 8, G) usually occur 
only along the leaf-margin. 

The transverse section shows 2—3 layers of short and 
broad palisade cells with rather large intercellular spaces; 
the cells of the spongy parenchyma are somewhat more 
branched than in the foregoing species (cf. surface view, 
Eje 8 HA): 

Chlorophyll-grains are abundantly present in the whole 
of the mesophyll; they are also found in the epidermis of 


the lower surface. 


Veronica officinalis L. 
(f. glabrata Fristedt). 


In the collections preserved in the Botanical Museum 
in Copenhagen there is some material of Veronica officinalis 
collected in the Færåes, part of which belongs to the main 
form with hairy leaves, and part to f. glabrata Fristedt 
(Væxtgeografiska skildr. af Sådra Ångermanland, Upsala, 
1857). Of both forms there were individuals of normal size, 
as well as dwarf individuals. 

The Morphology of the species has been described several 
times (e. g. WARMING, 1884, p. 58; BrunDin, 1898, p. 83), 
most exhaustively by the first-named author, from whose 
description the following is quoted: '"The creeping and rooting 
shoots bear only foliage-leaves (evergreen); at the apex they 
are frequently bent slightly upwards in a curve, especially 
when the plant grows among moss; but gradually as the 
"adventitious roots are developed in ascending succession, the 
stems are drawn down to the ground. From the basal por- 
tion of the year's shoots there proceeds shoots which re- 
semble the parent shoot; in this species no lateral shoots 
are found which can be indicated as special "assimilatory 
shoots”, which are such only, without taking at the same 


386 Fr. J. MATHIESEN. 


time part in the vegetative propagation; from the leaf- 
axils which follow next, racemose inflorescences proceed. But 
alter a short pause during the flowering period, the parent 
shoot continues its growth at the apex and, under favour- 
able conditions, may also produce new lateral shoots, as also 
it may, it is true, 
continue its growth 
throughout the 
winter, as soon as 
thetemperature has 
reached a certain 
degree of warmth. 
I have, however, 
observed several 
cases im which the 
main shoot had been 
biologically arrested 
through flowering, 
and had died away 
as far down as 





below the inflores- 


cences.” 
Fig. 9. Veronica officinalis L. A dwarf speci- 
f. glabrata Fristedt. men of f. glabrata 
Kirkebåkamp, Stromo (The Færådes). SR É i 
(About 2/,). (Strom6, Kirkebå- 


kamp Hest EMEE 
OsTENFELD, 8. 6. 1895) is illustrated in Fig. 9; it is drawn 
almost twice the natural size. The individual did not flower; 
the few leaves from the previous year which are still remain- 
ing, are recognizable by their being shaded. The branching 
is seen to be abundant — these dwarf individuals therefore 
often form rather dense tufts. Uppermost in the figure the 
last-formed leaves are seen to bend over the apex of the 


Scrophulariaceae. 387 


shoot; the reason of this is unknown to me. Upon the 
oldest portion of the shoot-system which has been figured, 
adventitious roots are seen to be rather abundantly devel- 
oped. 

According to Raunkiær (1907, p. 46) Veronica offici- 
nalis is an active chamæphyte; the species is nanophyllous, 
and evergreen (see above). 

In "Field-notes on the Biology of some of the Flowers 
of the Færåes” (Botany of the Færåes. Vol. III, p. 1065) 
E. WARMING writes concerning Veronica officinalis: "The 
diameter of the flower is 7—8 mm. The corolla is pale-lilac 
with stripes of a darker colour. Homogamous. The anthers 
and the stigma occur at the same level, but as the stamens 
are spreading, insect-pollination appears to be necessary for 
the setting of fruit.” 

Geographical Distribution: Europe, Western-Asia, 
North-America. 


According to OSTENFELD (1901, p. 57) im the Færåées 
Veronica officinalis is "Rather common in low-lying regions 
on hill-slopes and on rocky-ledges; also occurs at high levels 
and there mostly as f. glabrata Fristedt ... Fl. beginning of 
July. Fr. August, but bears fruit sparingly; often fails 
altogether in the hills.” 

The specimen illustrated in Fig. 9 was investigated with 
respect to its leaf-anatomy. The epidermis of the upper 
surface (Fig. 10, B) had lateral walls, from straight to slightly 
undulating, and that of the lower surface (Fig. 10, 4), 
rather strongly undulating lateral walls. Glandular hairs 
with one-celled stalk and two-celled head occurred abundant- 
ly on both leaf-surfaces. In Fig. 10, C some epidermal cells 
are shown under higher magnifying power; the two smaller 


and thin-walled ones have borne glandular hairs (the small 


388 Fr. J. MATHIESEN. 


circles inside their outlines indicate the connection of the 
stalk of the gland with the outer wall of the cell), the lateral 
walls of the others are seen to be porose with nodose thick- 
eningsf between the pores; in the main form I found these 
thickenings to be somewhat more strongly developed; they 
were very pronounced in individuals collected in dry habitats 
in Denmark. Å transverse section of the leaf is shown in 


2 





BD 
rs gere 


SV 
SE 


Fig. 10. Veronica officinalis L. 
f. glabrata Fristedt. 
A, Epidermis of the lower, and B of the upper surface of the"leaf. C, 
Epidermal cells from the upper surface of the leaf, more highly magni- 
fied. D, Transverse section of the leaf. (Å and B about ?%/,; C about 
2%/; D about ”5/,). (Kirkebåkamp, Stråmå (The Færåes)). 


Fig. 10, D; in the upper part of the figure there are two 
layers of short and broad palisade-cells; the spongy paren- 
chyma consists of rounded or slightly branched cells. Any 
special difference in the structure of the mesophyll in the 
Færéese and the Danish plants, could hardly be demonstrated ; 
the description given by KocxH (1895, p. 134) also agrees 
with that given above. 


Scrophulariaceae. 389 


Fig. 10, A and B, which are drawn from epidermis-pre- 
parations of one of the leaves with the under surface turned 
upward, seen uppermost in Fig. 9, show the proportion 
between the number of the stomata on the upper and lower 
surface of the leaf to be about 2 to 3; in the normally orien- 
tated leaves of the same individual, there were, however, 
generally about 3 times as many stomata on the lower as 
on the upper surface. On the whole, I always found in all 
the individuals of Veronica officinalis investigated by me, 
by far the greater number of stomata on the lower surface 
of the leaf, viz., 3 to 5 times as many as on the upper sur- 
face — this feature, as well as the absolute number of the 
stomata per unit of area, may however vary in the different 
.leaves on the same individual. 

The non-glandular hairs, which are found in greater or 
fewer numbers on the leaves and stems of the main form, 
are multicellular, thick-walled, and have cuticular warts. 


CastilleiaY pallida (L.) Kunth. 


Alcohol material from Kola (the Voronej River, leg. 
BROTHERUS, 2. 7. 1887) and Arctic America (King Point and 
Herschell Island, leg. A. H. LinnstroMm, 1905—1906). Her- 
barium material from Arctic America (Port Clarence, King 
Point and Herschell Island (var. unalaschkensis Cham.), 
Labrador, the coast of Hudson Bay (Ranken Inlet, Churchill 
(var. septentrionalis (Lind.) Gray)), Lapponia imandrae. i 

Lit.: LANGE, 1880, p. 79; E. WarminGc, 1890, pp. 220—223, fig. 
34; RoSENVINGE, 1892, p. 687; P. Knurx, 1899, p. 193; Simmons, 1913, 
pp. 122 and 138. 

Spot-bound, nano-microphyllous, sympodial proto-hemi- 
cryptophyte with a slightly branched primary root of long 
duration. Only one shoot-generation reaches maturity in 
each growth-period. The perennial basal portions of the shoots 


390 Fr. J. MATHIESEN. 


live for several years, and can have growth in thickness; 
adventitious roots are developed but sparingly. The shoots 
are erect and hairy especially above in the floral part; if 
they are capable of flowering — which ordinarily appears to 
be the rule — they terminate in a raceme, the subtending 
leaves of which are large, the lower ones similar to the foliage- 
leaves, but often having long lobes. 

In the axils of the 4—7 radical scale-leaves (bud-scales) 
of the shoots "”innovation-buds” occur, but of these only 
1—2 are further developed; the uppermost of them appear 
on the whole to be the most vigorous, but none of them is 
a decidedly principal bud. The foliage-leaves differ greatly 
in shape and size in the different forms in which the plant 
occurs; in var. unalaschkensis they are large, ovate-lanceolate 
(50—60 mm long, as much as 17 mm broad); in var. septen- 
trionalis they are smaller, linear-lanceolate 30—40 mm long, 
3—4 mm broad); the principal form is intermediate. Fruits 
with ripe seeds occurred in my material from the locali- 
ties in Labrador and Hudson Bay; the fruit ripens in July— 
August. The seeds are small and light with a reticulated, 
pitted testa; this pattern is produced by the very thin outer 
walls of the outermost, large-celled layer of the seed-coat, 
sinking down into the cavity of each cell, while to the inner 
and lateral walls rigidity is given by a network of anasto- 
mosing flange-like thickenings. 

The structure and biology of the flower has been ex- 
haustively described by E. WARMING, (l. c.), from whom I 
quote the following (compare also the accompanying figure 
with explanation which has been taken from the paper in 
question): "The hairy calyx is deeply cleft into two lateral 
lobes, which again are cleft into a larger anterior and a 
smaller posterior lobe, both oblong. Here. the corolla is only 
of about the length of the calyx, tube-shaped and two-lip- 


Scrophulariaceae. 391 


ped. The lower lip is shorter, with three ovate obtuse lobes 
which are erect or slightly reflexed; at the base of each lobe 
the lip is inflated into a bipartite arch (4, B and G). The 
upper lip is erect, boat-shaped, and terminates in an entire 
apex; a little distance below the apex it is provided on 














Fig. 11.  Castilleta pallida (From Kola at the White Sea). 
A and B, A flower seen from the right side and from the front; the 
anthers (anth.) have grown out of the flower and are open; the stigma 
protrudes above them; behind it the upper lip (s) is seen; the middle 
part of the lower lip is marked m, one of its lateral lobes (the right 
one) /. The hairs are indicated only along the periphery. C and D, 
Parts of a young flower, seen from the left side and from the front, 
after having been artificially opened. The anthers have not dehisced ; 
m is the middle lobe of the lip, I one of its left lateral lobes. E, A flower 
from the same inflorescence as A, seen from the left side, but much 
vounger and smaller; even here the stigma is large. F, Ovary of A. 
G, Each stigma in the whole inflorescence to which this flower belongs 
is as much enclosed as it is here; the anthers (anth.) of the long stamens 
are open, those of the short ones are closed. H, The relative positions 
of the stamens and stigma in the lowermost flower (14 mm long) in 
an inflorescence; the two uppermost anthers are open and are situated 
just above the stigma; the next flower appears to be similar; the third 
one was almost a bud, but had its stigma protruding to the usual 
distance. (Drawn by E. WARMING). 


392 Fr. J. MATHIESEN. 


either side with a rather thin wing (D and G), which in its 
upper part may be erect or curved slightly inwards around 
the anthers and stigma, while in its lower part the edge is 
revolute (G) as in Pedicularis, but the revolute portion ("die 
Rolle”) is quite smooth here. As both lips are erect and 
close tightly, there is, between them, a very narrow entrance 
to the flower, through which, on the one hand, only very 
small flies and other small insects can creep in, and on the 
other hand, only butterflies and bees with long proboscides 
can gain entrance into the flower. The anthers of the long 
stamens open before those of the short ones (HM), and the 
anthers of the latter always remain enclosed, while those of 
the long stamens protrude more or less. Here also the style 
appears to vary in length; in some inflorescenses I found it 
protruding far, even in quite young flowers — sometimes 
so young that they must rather be called buds (E); and as 
the stigma here also appears to be developed early, these 
flowers must be called longistylous—protogynous. In other 
flowers I found the stigma to protrude less, but nevertheless 
to be higher than the anthers, while in others again the 
long stamens were so long that their anthers were in con- 
tact with the stigma, or even protruded above the latter 
(H).” (The author then goes on to say, that among flowers 
from the same. inflorescence there may be some difference 
as regards the relative length of the style, the stamens and 
the corolla which apparently cannot be referred to differ- 
ence in age); "in flowers such as A, B and E (from the 
same inflorescence) self-pollination will be able to take place 
only with difficulty, the stigma being throughout higher than 
the anthers and the flower standing erect, hidden behind the 
large bracts provided with marginal lobes. Nor will self- 
pollination be easy in a flower like G, but in flowers like 


Scrophulariaceae. 393 


H, self-pollination will be able to take place easily, as the 
pollen will almost inevitably fall upon the stigma.” 

To the above description I can add, that in all cases, 
I found in fully developed flowers, with the exception only 
of the flowers in two inflorescences from Kola, — which hap- 
pen to belong to the material upon which WARMING has 
based his description — the stigma projecting more or less 
beyond the apex of the posterior lip, usually as in Fig. 11, 
A and B, sometimes projecting even more, and in quite 
young flowers somewhat less; in several cases I verified the 
presence of germinating pollen upon the stigma. It has not 
been possible to demonstrate conditions indicative of hetero- 
styly. It does not appear to be uncommon for the pollen, 
at any rate in the anthers of the two longest stamens, to 
germinate even before the flower has entirely expanded, i. e. 
while it is still hidden by the large subtending leaf, and the 
corolla is surrounded by the lobes of the calyx. As the 
stigma, even at that time, protrudes beyond the lobes of 
the corolla, the distance between it and the anthers will, 
however, in all probability be so great that self-pollination 
will not be able to take place; in material from Kola there 
was a flower, like that figured in /, which had germinating 
pollen in the anthers. 

Both calyx and corolla have a dense covering of shorter 
and longer non-glandular and glandular hairs, similar in form 
to those described below under the anatomy of the leaf. 
According to E. WARMING (l. c.) the flowers are visited by 
small wasps (the genus Pteromalus); he found such in several 
flowers. 

The Geographical Distribution of the plant is, ac- 
cording to LANGE: Labrador, Canada, the Rocky Mountains, 
western Arctic North America, Kamchatka, East and Arctic 


394 Fr. J. MATHIESEN. 


Siberia, and Arctic Russia. This author records it also from 
Greenland, but as it has not been found since, it ought to be 
omitted from the flora of Greenland (RosEnNnvinGE, M. Por- 
sILD). According to Simmons the following localities from 
the Arctic North American archipelago can be added: Banks 
Land, Baring Land, Victoria Land. 

Anatomy. The Root: The epidermis dies away quick- 
ly. The primary cortex follows for a long time the growth 
in thickness of the stele by tangential elongation of the cells, 
accompanied by divisions in these by radial walls. During 
the secondary growth the cambium forms outwards a large 
amount of parenchyma, arranged in very regularly radiating 
rows, while the formation of sieve-tissue is extremely spar- 
ing. The cells of the parenchyma are entirely filled with 
starch. In the woody parts the single growth-zones are easy 
to distinguish, in an inner narrow portion of the annual ring 
the vessels being surrounded by thin-walled cells, while in 
the outer larger part the vessels are accompanied by wood- 
fibres. Also in the thin-walled cells in question starch occurs. 
Parenchyma-rays are absent. B 

The Stem: As shown in Fig. 12, A and B the ring of 
wood in the lower, persistent part of the year's shoots is 
considerably thicker than in the upper, perishable part. In 
that part of the stem, from which the section shown in 
Fig. 12 A is taken, the epidermis, together with the outer- 
most thick-walled cortical layer connected with it without 
intercellular spaces, has partly separated from the rest of 
the cortex; this happens especially at the nodes, thus a leaf 
occurred on the axis just above the plane of the section on 
the side turned upwards in the figure. At the base of the 
shoots slits are formed at some depth in the cortex (Fig. 12, 
B and C); this explains the fact that the cortex can so easily 


be rubbed off in flakes. The persistent basal portions of the 


Scrophulariaceae. 395 


shoots have growth in thickness, but probably only to a 
very limited degree. In Fig. 12 C two annual-rings are seen, 
the last-formed is very excentric, the fact often being that 
the most vigorous growth in thickness takes place on that 
side of the stem which is opposite to a developing bud. 








o WAS | 
ORE 
SHOP SER 





BW 


XV 







Fig. 12. Castilleta pallida. 
AÅ and B, Transverse sections of the stem of the year's shoot; A is 
taken from half-way up the stem, B from near the base. C, Trans- 
verse section of a 2-year-old, persistent stem-base. D, Å portion of 
Å, more highly magnified. (King Point.) (4, B and C about ”%/,; D 
abon ra 


Fig. 12 D represents a part of A more highly magnified, the 
epidermis and the outermost layer of the cortex is thick- 
walled; the cortex is few-layered, its cells contain chloro- 
phyll-grains; stomata occur in the epidermis. The endo- 
dermis is only demonstrable by faint Casparian dots. 


396 


Fr. J. MATHIESEN. 


GUT 


LA 
DØR 
y ro) 


SD 


[ 
Sy 
986, 


8, 
NÅ 
cq y 





Fig. 13. Castilleita pallida. 
A and B, Transverse section of leaves (5/1). AÅ, var. septentrionale 
(Churchill); B, the principal form (Kola). C, Transverse section of the 
midrib of the leaf (Kola). D and E, The uppermost and lowermost 
layers of the mesophyll, seen in surface view. F, Transverse section 
of leaf. G, A glandular hair, more highly magnified. H and I, Epi- 
dermis from the upper and lower surface of the leaf respectively. 
(DFE EH and I King Pont: GK OlA)(CÆDFERFESE and abont 
EEG about, 3) 


Scrophulariaceae. 397 


In the wood, medullary rays are entirely wanting (the 
radiating lines in Fig. 12, A, B and C indicate only the radial 
arrangement of the elements); exteriorly, the ring of wood 
consists chiefly of wood-fibres; interiorly, vessels become more 
frequent, and are accompanied by partly lignified paren- 
chyma. In the basal portion of the stem the vessels are 
relatively more numerous than shown in Fig. 12, D, and the 
cells of the stereom are thinner-walled. The pith consists 
exteriorly of lignified, rather thick-walled, distinctly porose, 
axially elongated parenchymatous cells, towards the centre 
the cells of the pith are thinner-walled, non-lignified and 
die away, so that the stem becomes hollow. 

The Leaf: Any difference as regards anatomy could 
scarcely be demonstrated in the leaf-types of the different 
forms. Transverse sections, slightly magnified, of the leaf of 
var. septentrionale and of the principal form are shown in 
Fig. 13, A and B, the three main veins and the numerous 
fine anastomoses are seen. 

The vascular bundle of the main veins has on its under- 
side a covering of somewhat collenchymatously thickened 
elements; on the upper side the surface of the lamina dips 
down towards the vascular bundle. The structure of the 
mesophyll is fairly homogeneous, thus in Fig. 13, F it is 
hardly possible to demonstrate a palisade-layer; in specimens 
from Kola the uppermost layer of the mesophyll was how- 
ever elongated in a somewhat palisade-like manner. Chloro- 
phyll-grains occurred abundantly in the entire mesophyll, 
and besides this they were found in the epidermal cells of 
both surfaces. 

The epidermal cells of the upper surface have from 
straight to slightly undulating lateral walls, those of the lower 
surface can be more strongly undulating. Both the outer 
and lateral walls of the epidermal cells are thin; the cuticle 


XXX VIL 26 


398 Fr. J. MATHIESEN. 


of the outer walls is but slightly developed. Stomata occur 
in almost equal number on both sides of the leaf; the guard- 
cells are surrounded by 4—5 cells, and are on a level with, 
or slightly raised above, the surface of the lamina — on 
the large subtending leaves of the flowers I found them 
sometimes raised high above the surface. 

The leaves are densely covered with hairs; the following 
types of hair occur: (1) Non-glandular, thin-walled, pointed 
and with fine cuticular striations; they may either be short 
(1—2 celled) or long (multicellular) and even elongated like 
a whip, the last-mentioned are numerous especially in var. 
unalaschkensis, and particularly in the top of the shoots. 
(2) Glandular hairs with multicellular stalk, and with 1—2 
celled head; the basal cell has sometimes cuticular striations. 
(3) Glandular hairs with unicellular, quite short stalk and 
2-celled head (in Fig. 13, G such a hair is seen in lateral view, 
highly magnified; in H and I they are seen from above), 
in the cells of the head and the stalk comparatively large 
nuclei occur, and a highly granular protoplasm. These various 
forms of hair occur intermixed (ef. Fig. 13, F, H and M/); 
they are also found on the surface of the stem and, as already 


mentioned, in the floral region. 


Euphrasia arctica Lange. 
(Euphrasia latifolia (Pursh) Wettst.) 

Alcohol material from West and East Greenland (Fre- 
drikshaab, leg. ROSENVINGE, 15. 8. 1886; Sydproven; Hekla 
Havn, leg. N. Hartz, 13. 8; Unartok and Tasiusak). 

Herbarium material from West and East Greenland, the 
shore of Hudson Bay, Northern Norway, Lapponia murman. 


Lit.:" LANGE, 1880,-p:79:1887,"p- 264; "WARMING, 1886, pp. VII 
and 43; 1888, pp. 34 and 59; 1890, p. 226; RosENVINGE, 1892, p. 687; 
1895 (1), pi 68; TS95T (IT) ED PENGENE SEPT and 245 EAR DERS 94 


Scrophulariaceae. BØ 


pp. 9, 15, 20, 46 and 57; 1895 (I), pp. 146, 170, 179 and 289; 1895 (11), 
pp. 335, 359, 372 and 377; Hartz and Kruuse, 1911, pp. 359, 364, 
409, 416, 423 and 428; WETTSTEIN, 1896, p. 136; ÅBROMEIT, 1899, 
p: 46; Dusen, 1901, p. 40; Porsizn, 1902, p: 197; 1912, pp. 382 and 
387; 1920, p. 141; Kruuse, 1905, p. 176; 1906, p.'250; 1911: in part 
IV, pp. 196, 229, 242, 247 and 261, and besides these many notes in 
the preceding parts; JØRGENSEN, 1919, p. 99. 


Ås regards the Morphology, Biology and Anatomy of 
the Euphrasia spp. see: MULLER, 1881, p. 279; HOVELACQUE, 
1888, pp. 400, 454 and 477; Linnman, 1887, p. 81; KocxH, 
1895, pp. 140—144; WETTSTEIN, 1896; HEI1INRICHER, 1898 
and 1902; KnurtHxH, 1899, pp. 202—206; and KERNER, 1900. 

Nanophyllous therophyte; the seed germinates during 
early summer. 

The main root is rather scantily branched; above the 
cotyledons there are 1—3 pairs of leaves separated by rather 
long internodes; "the first flowers generally occur in the axils 
of the ård or 4th pair of leaves, frequently in that of the 
2nd pair, more rarely not until in that of the dth, and as 
an exception, even in that of the Ist pair” (JORGENSEN). To- 
wards the apex of the stem, the distance between the leaf- 
pairs becomes very short; the leaves become broader, and 
their teeth longer and more pointed than are those on the 
lower leaves of the stem; they all subtend flowers. The 
branching is, as a rule, scanty; sometimes a few weakly- 
developed branches åre seen to proceed from the uppermost 
pair of leaves below the inflorescence; occasionally, however, 
branches, almost as vigorous as the main axis, may be devel- 
oped from the axils of the two lowermost pairs of leaves — 
in this case it is evidently a matter of rather late-flowering 
individuals (August—September). Stem, leaves and calices 
are more or less densely covered with hairs. 

"Die Pflanze variiert sehr viel, von sehr klein und ein- 
fach, mit wenigen Blithen — so håufig an der Witterung 


26% 


400 Fr. J. MATHIESEN. 


ausgesetzten alpinen Standorten — bis sehr gross, grob und 
veråstelt mit sehr grossen und breiten, grobgezåhnten Blåt- 
tern, die bis uber 2 cm lang und ebenso breit werden” (Jor- 
GENSEN). Similar variations were found in the Greenland 
material. From the above-said it follows that the individuals 
generally agree in habit with the growth-type indicated by 
WETTSTEIN as "fruhblutige Form” (1. c. Fig. 1, p. 44); the 
comparatively short period of vegetation with which the 
individual may be compelled to be satisfied, in consequence 
of the geographical distribution of the species, makes, as 
JØRGENSEN (I. c. p. 104) remarks, this growth-type necessary 
in Arctic regions. 

The flower-morphology and -biology of the Euphrasia 
spp. have been described so often that I can here confine 
myself to the following remarks: E. arctica belongs decidedly 
to the small-flowered forms; in no case did I find the corolla 
to be more than 7 mm in length, more frequently it was 
shorter. The flowers are protogynous, and in the recently 
expanded flower, the stigma generally stands somewhat in 
front of the anthers, afterwards the style curves downwards 
and backwards, so that the stigma is brought into close 
contact with the anthers which åre now quite open (WAR- 
MING, 1890). Sometimes, however, the stigma, even at the 
time when the flower is expanding, is so near to the anthers, 
that self-pollination must be able to take place easily; I even 
found pollen on the stigma of a not yet expanded flower. 

WARMING (1890, p. 227) mentions that a length-incre- 
ment of the corolla-tube can take place — as in larger- 
flowered species —, by which the anthers are carried forward 
and approach the stigma; in some few of the flowers I did 
find the stigma protruding so far that a movement of this 
kind may very probably be of importance. 

Geographical Distribution according to JORGENSEN : 


Scrophulariaceae. 401 


Greenland, Cumberland, Labrador, Iceland, the Færåes, Scot- 
land, Sweden southwards to Herjedalen and Jåmtland, Nor- 
way (common towards the north, rare south of the Trond- 
hjem Fjord), Northern Finland and Russia, in Siberia at 
least as far towards the east as Jenisei. Porsirn (1912, 
p-. 382) records the northern limit of the species in West 
Greenland to be Tartusaq (71725' N. lat.). In East Green- 
land E. arctica extends almost to 74” N. lat. (Lille-Pendulum 
Island: Dusen); from the district of Angmagsalik north- 
wards it occurs most frequently, and grows more luxuriantly 
at the head of the fjords, whilst it is rarer and stunted in 
growth along the coast outside the fjords and on the islands 
off that coast. 

The Habitat is given by the various authors in unison 
as being willow copses, heaths, grassy slopes and "herb- 
slopes”; RosENVINGE (1896 (II), pp. 219 and 245) also found 
the species growing on gravelly flats (Igaliko) and on knolls 
in mossy bogs; HarRTzZ found it in the district of Scoresby 
Sound flowering as late as the middle of September. The 
fruit-setting appeared everywhere to be abundant and good. 

The Anatomy of theroot and the stem has been very 
exhaustively described by HOVELACQUE as regards "E. offi- 
cinalis”; in E. arctica I found nothing which differed from 
his description. 

The Root has only a few root-hairs; the cortex is thin 
and few-layered (3—4 cell-layers in thickness), its elements 
are greatly elongated in a tangential direction; in the endo- 
dermis the Casparian dots are distinct; in proportion to its 
diameter, the xylem part in the full-grown root forms a 
very thick bundle, composed of rather thin-walled vessels 
and wood-fibres. 

The Stem has a rather thick-walled epidermis, upon 
which occur 2—3 celled, non-glandular hairs with fine cuti- 


402 Fr. J. MATHIESEN. 


cular striations, and small glandular hairs with one-celled 
stalk and two-celled head. The cortex is few-layered (3—4 
cell-layers in thickness) as in the root; in the endodermis 
the Casparian dots are faint, but nevertheless demonstrable 


G 





Fig. 14. Euphrasia arctica. 
A,. Epidermis of the upper, and B of the lower surface of the leaf. 
C, Transverse section of the leaf. D and E, Glandular hairs from the 
lower surface of the leaf. F, A glandular hair from the margin of the 
leaf. G, A 2-celled, non-glandular hair. (4, B, C, D, E, F and G about 
ON NE (Greenlande) 


with Sudan III.. The pericycle is 1—2 layered. The outer 
part of the rather broad ring of xylem is composed exclusively 
of wood-fibres. The peripheral cells of the pith have some- 
what thickened walls. Medullary rays are absent. 

The Leaf. A transverse section of one of the lower 


foliage-leaves is shown in Fig. 14, C; there occur 1—2 layers 


Scrophulariaceae. 403 


of short palisade-cells, with rather large intercellular spaces; 
the spongy parenchyma consists of only slightly branched 
cells. The mesophyll of the upper leaves of the stem (i. e. 
the leaves subtending the flowers) appears, on the whole, 
to be more lacunose in structure than that illustrated in 
ITS Er RØR 

The margins of the teeth of the leaves are somewhat 
bent over, whereby a concavity is produced on their under 
side. In this concavity the leaf-surface is densely covered 
with glandular hairs, and the cells of the epidermis differ in 
character from the other epidermal cells of the leaf. For 
whilst these, both on the upper and lower surface — and 
especially on the upper side of the teeth of the leaves — 
have strongly undulating lateral walls (Fig. 14, A and B), 
which only here and there show a small flanged thickening, 
the epidermal cells in the concavities, and especially those 
which bear the glandular hairs, have straight or only slightly 
wavy lateral walls (Fig. 14, D and E), which are often (not, 
however, in the places illustrated in Fig. 14, D) furnished 
with flanged thickenings, the one by the side of the other. 
Stomata are almost equally abundant on both leaf-surfaces, 
in the concavities of the lower side of the teeth, however, 
they are less frequent; the guard-cells are on a level with 
the surface. In the teeth the veins break up into a fine net- 
work of tracheids. 

The glandular hairs which occur on the lower side of 
the teeth are of two types: (1) a smaller one with one-celled 
stalk and two-celled, globular head, and (2) å larger one with 
two or more frequently four-celled cupola-shaped head, seated 
upon a short and broad stalk-cell which partly sinks below 
the leaf surface, the inner-wall of the stalk-cell, during the 
development of the hair, becoming rounded inwards, thus 
pressing the layer of epidermal cells, situated under the stalk- 


404 Fr. J. MATHIESEN. 


cell, in the same direction (Fig. 14, C). The stalk-cell of the 
latter type of hair rests upon the connecting parts of 4—8 
epidermal cells, which åre produced by the division of a 
single cell, viz., the epidermal cell which has been the mother- 
cell of the glandular hair. Below the stalk-cell small, circular 
intercellular-spaces are formed between the lateral walls of the 
cells of the epidermal layer (cf. Fig. 14, C and D; in D that 
part of the lateral walls of the epidermal cells which lie under 
the stalk-cells, is indicated by dotted lines — the dotted circle 
inside the head of the small glandular hairs indicates the 
outline of the stalk-cell). 

These large, peltate glands appear, therefore, to belong 
to the same type as the ”Schilddrusen” of Lathraea (and 
Bartschia), the development of which has been so thoroughly 


described by SCHERFFEL; the intercellular spaces in the layer 


under the stalk-cell have not been perceived by HoVELACQUE 
or WETTSTEIN. 

The small glandular hairs belonging to type 1 are also 
found in other places on the leaf, but nowhere in such abun- 
dance, as in the concavities on the under surface of the teeth. 

Glandular hairs with 1—2 celled head and multicellular 
stalk (Fig. 14, F), such as those required for the diagnosis 
of species, are found both along the margin of the leaf and 
also — hut scantily — upon the lower surface. 

Non-glandular hairs occur upon both leaf-surfaces, and 
are especially numerous towards the margin and on the 
veins; they are most frequently one-celled, but can also be 
2—3 celled, pointed, and have fine cuticular striations (Fig. 
14, 4, C and G; C and G are seen in optical section; the 
thickness of the walls is evident from the figures). The epi- 
dermal cell or cells (2—4) which bear them are distinguished 
from the surrounding cells by having less strongly undulating 
lateral walls, and outer-walls which are more or less strongly 


arched outwards. 


Scrophulariaceae. 405 


In the concavities on the under surface of the teeth I found 
the epidermis covered with crystal aggregates, which proved 
to consist of calcium-carbonate. — The same observation 
has been made by WETrTSTEIN (l. c. p. 19). I have sometimes 
also found crystals of the said substance in the cells of the 
large glandular hairs, as shown in Fig. 14, E. 

The palisade-tissue and spongy parenchyma were rich 
in chlorophyll-grains; and such were also found in the epi- 
dermis of the lower surface. 


Bartschia alpina L. 


Alcohol material from Norway (Aursundséen, leg. TH. 
ResvoLL, 31. 8. 1918; Bosekop, Kaafjord, Alten, .leg. E. 
WARMING, 11, 17. 8. 1885), Greenland (Sukkertoppen, 5. 7. 
1884; Lyngmarken, leg. L. K. RoOSENVINGE, 25. 7. 1886; Kan- 
gerdluarsuk, 5. 8. 1884 and 30. 7. 1885; Holsteinsborg, 15. 7. 
1884; Sarfanguak, 15. 7), and Iceland (Gullfos and Tungafos, 
leg. A. FEDDERSEN, 25. 6. and 30. 6. 1886). 

Herbarium material from Fennoscandia, Island of Kol- 
gujew, the Færåes, Iceland, Greenland, the coast of Hud- 
son Bay. 

Free PASSEDE LS GO OS BANGE ET 88 0 Ep SEEST SEP 263; 
MULLER, 1881, p. 283; WARMING, 1886, pp. II, VI, 7 and 43; 1888, 
pp. 34, 39, 67, 75, 93 and 188; 1890, -p. 226; Lunnman, 1887," p. 82; 
HoVvELACQUE, 1888, pp. 403, 451 and 478; RosENVINGE, 1892, p. 687; 
1189 GE pp SEAL 6 ISEN GS Handl 6 SS EAR ere SOLE pp OND BENE 
43; 57; 1895 (I), p. 302: 1895 (II), pp. 359, 372 and 392; JUNGNER, 
1895, pp. 224 and 238; Norman, 1895, p. 455; ABROMEIT, 1899, p. 45; 
Knurtx, 1899, p. 197; KERNER, 1900, Bd. I, pp. 107, 129, 131 and 32, 
i 3an de 60 GB Lp pA ES OP sande ON Greve ORE Pp: 
89 RSS NAK 1076 and 89: "Hejnricher IT 190" Porsrep, 1902; 
pp- 181 and 194; 1912, p. 382, 387; 1920, p. 141; Kruuse, 1905, p. 177; 
1906;p: 249: 1911" im part IV, pp. 202, 229; 242, 244 and 254, and 
besides these numerous places in the preceding sections; SYLVEN, 1906, 
p. 86; Horstern, 1907, p. 126; ScaRrRårteErR, 1908, pp. 465—68 and 772. 

A nanophyllous to microphyllous proto-hemicryptophyte 
with subterranean runners; the shoot-development usually 


extends over 2 years. 


406 Fr. J. MATHIESEN. 


According to HEINRICHER and SYLVEN the seed germin- 
ates in early summer. The first-named author, who has 
studied Bartschia very thoroughly, has pointed out that it 
belongs to the comparatively few plants which produce buds 
in the axils of the cotyledons. Whilst the uppermost part of 
the primary shoot dies away, the following year one of these 
buds develops into a foliage-leaf-bearing shoot, in the basal 
portion of which, small innovation-shoots arise, which, like 
their parent-shoot, will continue their growth in their second 
year, and become foliage-leaf-bearing shoots. As was the 
case with the primary shoot, the uppermost part of the axes 
of the succeeding shoot-generations dies down to above the 
innovation-shoots. Even after the 4th vegetative period, 
HEINRICHER'S cultures did not yet bear flowers; the young 
plants had formed small sympodial, subterranean rhizomes, 
composed of the persistent, basal portions of the innovation- 
shoots. 

HEINRICHER is of opinion that, in Nature, the first vege- 
tative-stage is of 4—5 years duration. 

With regard to the morphology of the full-grown plant, 
in the following description the statements given in the cited 
work of HEINRICHER will be found verified; for criticism 
of preceding investigations (HOVELACQUE, KERNER), reference 
may be made to the same author. 

Fig. 15, A and D show the structure of the shoot, and 
the mode of branching. In Fig. 15, A, each of the two shoots, 
II? and II?, had terminated in an inflorescence. The shoots 
aré in their 2nd year of development; during the first year 
each of them had developed a subterranean portion, bearing 
decussated scale-leaves which have now withered; the upper- 
most pairs of these scale-leaves has served as bud-scales for 
the winter-buds. The first year's growth-increment especially 


of II” is seen to have the character of a runner, the growth- 


Scrophulariaceae. 407 


direction has been horizontal, and the distance between the 
nodes is comparatively considerable. The apex of such a 
runner is shown int Fries ION CS In Eje SIS DIT Mast not 
borne flowers; the sister-shoot 7/77 has been arrested in its 





Fig. 15. Bartschia alpina. 
A, The lower portion of a flowering plant; both 7/1" and II? have borne 
inflorescences (Holsteinsborg, 5. 7. 1884). B, A foliage-leaf. C, The 
apex of a runner. D, Portion of a sympodium in which the runner- 
like portion of shoot // is particularly long; r is an adventitious root. 
(C and D, Aursundséen, 31. 8. 1918.) (About ”/,.) 


development, the two small basal pairs of leaves are from 
the previous year, and withered, but the bud is in a fresh 
condition; like shoot 77 shoot / has had a long runner-like 
portion. The innovation-shoots are always developed from 
the portion of the shoot of the first year; in case the latter 


408 Fr. J. MATHIESEN. 


shoot is elongated in a runner-like manner, the new shoots 
generally appear near the spot where it begins to bend up- 
wards, but if it is short and vertical, the new shoots most 
frequently appear at the base — sometimes, however, also 
higher up; as an exception, shoots ("enriching-shoots”) may 
appear in the axils of the lowermost foliage-leaves: such 
shoots, however, will probably be always purely vegetative 
and terminate their development in the same year as the 
parent-shoot. After flowering and fruit-setting, the latter 
shoot dies down to immediately above the innovation-shoots. 

Innovation-shoots are generally developed to the number 
of two on the parent-shoots; //7 and II?” in A and III? and 
III? im D are seen to have arisen in the axils of two opposite 
scale-leaves; this is most frequently the case, and explains 
the fact why two flower-bearing axes are frequently found 
to stand together. In Fig. 15, D å non-expanded bud is seen 
in the axil of the first scale-leaf to the right of the subtending 
leaves of the innovation-shoots. 

The innovation-shoots are developed rather late in the 
Arctic summer; in the abundant herbarium-material from 
Greenland which I have at my disposal I never found the 
young shoots to be visible in individuals collected in June— 
July, whereas in those collected in August they were about 
one centimetre long. 

This does not quite agree with the statement of HE1n- 
RICHER, according to which they already expand simultane- 
ously with the foliage-leaves of the mother-shoot. The dif- 
ference may perhaps be explained by the more unfavourable 
conditions of growth, presented by the Arctic summer, in 
which the building-up of new organs can be compelled to 
extend over a longer space of time. 

The same author states, that small buds for the inno- 
vation-shoots are already to be found the year previous to 


Scrophulariaceae. 409 


that, in which they expand; the same is the case with the 
buds for the flowers. 

Ås already mentioned, the portion of the shoot devel- 
oped during the first year of growth, may either have elong- 
ated internodes and be runner-like, or — as e. g. figured by 
KERNER on p. 129 (1. c.) — it may have quite short inter- 
nodes with close-set scale-leaves; the first 2—3 internodes 
of the stem are, however, generally somewhat longer than 
those following; and here the scale-leaves are smaller. If by 
means of the latter form of shoots the ramification should 
be continued through several shoot-generations, the indivi- 
dual becomes gradually cæspitose in habit. A tuft of this 
kind collected by M. Porsirn in West Greenland ("heathy 
slopes”) had about 50 older and withered shoots, and some 
20 fresh ones — both vegetative and flowering. The indivi- 
dual had sometimes formed quite short runners, adventitious 
roots were scantily developed, and the entire system was 
borne by a single root (whether main root or adventitious 
root was not clear) about 5 mm thick. 

The adventitious roots spring — often abundantly — 
from the nodes. 

The rhizome, probably as a rule, dies away rather quickly 
from behind; however, in a specimen from East Greenland 
with cæspitose habit, a part of it had attained a thickness 
of fully 5 mm, and the whole system was evidently old. 

The plants are densely covered with hairs especially at 
the top; hairs (non-glandular) also occur on the subterranean 
portions of the shoots, KERNER ascribes importance to these 
hairs as organs of absorption; according to HEINRICHER, 
however, there is no reason to assume that they have such a 
function. 

The Flower-biology has been dealt with by H. MiLLERr, 
E. WARMING, LINDMAN, KERNER and others. The accompa- 


410 Fr. J. MATHIESEN. 


nying figures are borrowed from WARMING, 1886, and the 
following is cited from his description: "The Scandinavian 
and Greenland individuals of this species appear to me to be 
exactly alike in every way, but differ from those of the Alps 
in at least one respect, that is, provided that H. MiLLER has 
informed us of all the forms occurring there. Fig. 16 shows 





Fig. 16. Bartschia alpina. 
A and B from East Greenland (EBEeRrLin, 30. 7. 1885); C from Kaa- 
fjord near Alten in West Finmark; D from Godthaab in West Green- 
land (28. 6. 1884); all the anthers were open, and pollen had already 
germinated upon the stigma. E and F are from Holsteinsborg in West 





Greenland (15. 7. 1884). A—F are twice the natural size; the others, 
which are all drawn from Greenland material, are in various magni- 
fications. For further particulars see text (WAarRmInG, 1886). 


the structure of this flower, partly from Norwegian and partly 
from Greenland material (in the majority of cases the hair- 
covering has been omitted or only indicated along the outlines 
of the figures). The conspicuousness of the dull purple-violet 


Scrophulariaceae. 411 


corolla, which like the calyx, is densely covered with glandular 
hairs, is increased by the fact that the subtending bracts 
are also of a dull violet. I have not noticed any scent, but 
honey is secreted by the greenish nectary on the front of the 
ovary (Fig. 16, G). MuLLer has described how humble-bees 
and other insects, by thrusting their proboscis and also their 
heads into the flower (see Fig. /, the corolla seen from the 
front) strike against the pointed lower ends of the anthers 
(Fig. 16, K and L), whereby the anthers which adhere by 
their hairs, are violently torn from one another, by which 
means the dry and light pollen-grains are scattered and fall 
upon the insect, which then easily conveys them to another 
flower and deposits them upon the stigma (Fig. 16, A—E, M) 
which protrudes even in the bud (Fig. 16, B). Specimens with 
flowers, the structure of which is exactly like that of the 
flowers of the Alps, have also been found to occur both in 
Norway and in Greenland?, these specimens have evidently 
marked insect-visited flowers which either cannot pollinate 
themselves, or can only do so with great difficulty. My figures 
(which have all been drawn from full-grown flowers) show 
that both the length of the style and the size of the whole 
flower vary considerably, apart from the difference of age. 
Thus specimens are found which have so short a style through- 
out their whole life, that the stigma does not become visible 
outside the corolla; in some, however, the stigma is situated 
immediately inside the throat of the flower (Fig. 16, F, K), 
but in others it lies even as far inside as above the hind- 
most anthers (Fig. 16, L). In these cases the stigma lies 
against the anthers, and self-pollination, as far as I can see, 


is inevitable; and in such flowers I have also distinctly seen 


1 MULLER does not mention that the style is beset with stiff 
hairs directed forwards, except at its uppermost end (Fig. 16, H); but 
it is probable that it has this feature also in the Alps. 


2 Fr. J. MATHIESEN. 


pollen-grains deposited especially upon the lower part of the 
stigma — the part which is in contact with the anthers 
(Fig. 16, M). For the rest, the pollen-grains fall out very 
easily, and it is very common for masses of pollen-grains to 
be found scattered everywhere in the flower: on the corolla, 
the filaments, the style, etc. Even if the stigma ripens shortly 
before the anthers — which I found to be the case in the 
district of Holsteinsborg — the two organs nevertheless very 
soon become simultaneously functional and this throughout 
the greater part of the life of the plant, so that self-polli- 
nation is possible. That Bartschia, in many places in Green- 
land, produces ripe fruit, I saw in 1884, from the fragments 
of such fruit which had remained over from the preceding 
year.” 

According to KERNER (II, p. 301) wind-pollination takes 
place regularly in Bartschia, in a similar manner as for in- 
stance in Lathræa, at the end of the flowering period: the 
style in the older flower withers, the filaments are elongated 
and carry the anthers outwards, the latter separate and 
pollen, which may have been left in the anthers is scattered 
by the agency of the wind, and may pollinate the younger 
(upper) flowers of the inflorescence. This probably explains 
the fact mentioned by Linnman that the anthers may pro- 
ject beyond the edge of the corolla. This author also men- 
tions the same variations in the length of the style as those 
of which WARMING speaks. 

Also in the material at my disposal there were flowers 
im which the anthers protruded, and it was generally the 
lowermost (oldest) flowers of the inflorescence in which this 
was the case. 

AÅAuG. ScHuLz differs from WARMING in finding that in the 
Riesengebirge (Bibl. bot. No. 10, 1888, p. 74 — here cited 
from Knuth) the length of the style, in relation to the corolla, 


Scrophulariaceae. 413 


differs in older and younger flowers, so that the style of the 
younger flower protrudes farther than that of the older, the 
corolla-tube of the latter having increased in length. The 
elongation may be as much as 5 mm and bring the anthers 
— the filaments follow the corolla in growth — into contact 
with the stigma, so that self-pollination takes place. War- 
MING (1890, p. 226) admits that such a growth can take place, 
and disturb the relations of the organs in question, but main- 
tains that individual differences also occur in the relative 
length of the style and the corolla. I found the same forms 
that have been illustrated by WARMING (1886) in his Fig. 
3, A—F, K and L (reproduced in this paper as Fig. 16), 
and even if there might possibly occur some difference in 
the length of the corolla of older and of younger flowers of 
the same inflorescence, yet I never found the style pro- 
truded far in the young flowers and enclosed in the older 
ones; that, at any rate under more northerly latitudes, it is 
really a matter of individual differences, is supported by a 
statement of Mr. NyHuus of Tromsé, who in a letter to 
E. WARMING in 1885 writes: "On Aug. 24th I found Bartsia 
there (i. e. Dalfjæld in Marknæsdalen) in abundance (towards 
the south) from a height of 2500 feet and down over the 
whole mountain; but from about 2500 to 1500 feet I did 
not find a single flower with protruding style although I can 
truthfully say that I examined several hundred plants. On 
the other hand, lower than that, long-styled flowers became 
constantly more frequent” (E. Warming, 1886, p. 10). 

I found the fruit-setting to be good and abundant every- 
where. "Die geflugelten Samen werden durch den Wind ver- 
breitet.” (SCHROTER). 

Geographical distribution in the Arctics according 
to LANGE and Porsirn: Labrador, Arctic Russia and Siberia, 
(according to F. R. KJELLMAN's lists of the flora, in Siberia 


XXXVII. 97 


414 ' Fr. J. MATHIESEN. 


it does not seem, however, to occur at or near the coast), 
Scandinavia, Iceland, the Færåes and Great Britain, in 
addition, the species extends over "ein mittel-europåisches 
Gebirgsareal von den Pyrenåen bis zum Balkan mit Vor- 
ståssen zu den deutschen Mittelgebirgen und zahlreichen 
Standorten im Vorland” (ScHROTER). In West Greenland its 
northern limit is the Ignerit Fjord (Umanak), in East Green- 
land it does not reach so far as to the region of Scoresby 
Sound. KruusE (1905, p.177) states that the northern 
limit there is at 6828' N. lat. 

Habitat. According to WARMING in West Greenland 
it grows in willow-copses and on "herb-flats”; in the most 
southerly West Greenland RosENVINGE found it in willow 
and birch copses, on "herb-slopes” and grassy slopes. 

Ås regards the conditions pertaining to the growth of 
Bartschia in East Greenland Kruuse (1905, p. 177) writes: 
St is very rare north of 66720' and only 10 cm. high, but 
yet it sets flower everywhere and as far as can be judged 
from the collected material also fruit. It shuns here the 
coast, and all finding places are well sheltered, exposed to 
the south and have the character of herby slopes, while the 
species more to the south prefers heath”, and in the Ang- 
magsalik district it is (1906, p. 249) "commonly distributed 
on herby slopes and in fertile heath”. 

NORMAN records it from Northern Norway as growing 
Sin bogs, on damp cliffs, in birch-woods, on flats at the river- 
banks and on the beach, as an exception below the upper- 
most belt of sea-weeds; it occurs chiefly on the sunny sides, 
somewhat more seldom on the indifferent (eastern and west- 
ern) sides and as an exception on the shady sides”. A. CLEVE 
records that on the mountains in the north of Sweden it is 
"Ein sparsamer, exclusiver Bewohner der trockenen Bliten- 


wiese mit vollen S-Exposition . .. Ein typischer hydrophyt 


Scrophulariaceae. 415 


der Weiden- und Waldregionen Lapplands, kann aber diese 
Art im Hochgebirge sich nur an den viel Wårme darbieten- 
den Stellen behaupten, mågen letztere auch viel trockenen 
sein, als es gewohnlich in dem Tieflande der Fall ist.” 

In the Alps it grows on "alle Formen alp. Wiesen und 
den Quellfluren von den Vorbergen bis gegen die Nivalregion 
(ScHROTER)”. 

In the Arctics Bartschia belongs to the group of plants 
of later summer, which is easily understood when one con- 
siders the considerable vegetative work the shoots must 
carry out, before flowering can take place. 

Anatomy. The Root. An adventitious root, about 
1 mm thick, showed the following structure: The epidermis 
had died away, only here and there a few of its cells remained 
in a collapsed condition. The outermost layer of the primary 
cortex had cuticularized cell-walls; in many of its cells a 
division had taken place during growth, and in these cells 
one very thin, non-cuticularised radial wall was found. The 
cells in the inner layers of the cortex were distinctly tan- 
gentially elongated and were likewise divided secondarily by 
thin radial walls — in the cells of the endodermis ås many 
as four such walls were found. The wall of each of the ori- 
ginal endodermal cells is in its whole circumference furnished 
with a cuticularized layer, but as in the exodermis, the thin 
radial walls which had developed later, were not cuticularised 
here either. The root was 3-rayed; in transverse section the 
xylem-part now formed a circular bundle, consisting of ves- 
sels and wood-fibres, since the cambium had commenced its 
activity all the way round. The epidermis evidently dies 
away very early; even in a root hardly 0.5 mm in thickness, 
the epidermal cells were found in a collapsed condition. 

The root of 5 millimetres thickness, with which the indi- 
vidual from West Greenland, mentioned above, was fur- 


27% 


416 Fr. J. MATHIESEN. 


nished, had a xylem-cylinder, 3 mm thick, which consisted 
for the most part of vessels, thin-walled and rather wide; 
in addition thin-walled wood-fibres occurred. 

Parenchyma rays did not occur, neither were there any 
indication of growth-zones. In the secondary cortex sieve- 
tubes were rather 

scantily present. 

Cork was not devel- 
oped. In the roots 
both the endoder- 
mal and the exor- 
dermal cells were 
full of a brown sub- 
stance. 

The Stem. In 
HOVELAQUE's work 





it is particularly the 
FLAPS i j anatomy of the stem 
Fig. 17. Bartschia alpina. å g 
Diagrammatic transverse sections of the stem: which has been ex- 
A and B, from runners; C, from the aerial  haustively treated, 
stem taken between the 3rd and 4th pair of É 
foliage-leaves; D, taken at the point of tran- 
sition between rhizome and the aerial stem on this point, since 
(the black ring in C and D is the hard-bast, 
whilst the wood-ring in all the figures is radi- 
ally shaded). (About 78/,.) ated by HOLLSTEIN. 
To this I have noth- 


ing new to add, and can therefore confine myself to the fol- 


and Bartschia has, 


been again investig- 


lowing brief description. 

Å section taken from a little above the middle of the 
foliage-leaf-bearing part of the stem (a diagrammatic repre- 
sentation is given in Fig. 17, C) shows a thin-walled epider- 
mis, beneath which there is a cortex, the outermost layer of 
which has somewhat thickened cell-walls; several of the cor- 


tical cells have undergone division by means of thin radial 


Scrophulariaceae. 417 


walls. In the endodermis — contrary to HoVELAQUE, who 
mentions its cells as being "sans cadres” — I found distinct, 
although rather faint, Casparian dots. The outermost layers 
of the stele consist here of rather thin-walled lignified prosen- 
chymatous cells; the sieve-tissue forms a continuous ring 
outside the xylem-part, which in the whole extent of its 
periphery consists almost exclusively of wood-fibres. The 
pith consists of thin-walled cells which collapse early in the 
centre of the stem. 

A section taken at about the middle of the subterranean 
part of the axis, is distinguished from the above by having 
a more distinct endodermis, the cells of which have, as in 
the root, a suberised lamella in the whole circumference of 
the wall — the contents of the endodermal cells have a 
brown colour —, by entirely wanting hard bast, or at any 
rate, by having it only in a far more slightly developed 
degree (in the two diagrammatic figures, Fig. 17, A and B, 
the line between the outer edge of the xylem-ring and the 
periphery is the endodermis, the black dots within this, in 
Fig. B, are small groups of hard-bast cells — in A these 
are totally wanting), and lastly, by a comparatively nar- 
rower pith and broader xylem-ring, in which wood-fibres 
are less numerous and thin-walled. 

At the point of transition between rhizome and the 
aérial stem, the hard-bast appears successively (the cells of 
the hard-bast are here somewhat more thick-walled than at 
the top of the shoot), the pith widens, the xylem-ring becomes 
thinner and richer in wood-fibres throughout its periphery. 
The sieve-tissue is divided into four larger groups, since the 
hard bast and the wood-fibres of the xylem-ring at four 
points which decussate, tend to approach one another, and 
may practically coalesce at these points (this phenomenon 
is connected with the passing-out of the leaf-trace-bundles 


418 Fr. J. MATHIESEN. 


from the stele, compare Fig. 17, D, which is a diagrammatic 
representation of a part like the one just described — the 
section was taken between the uppermost scale-leaf and the 
first pair of foliage-leaves), small groups of sieve-tissue 


A B 








Fig. 18. Bartschia alpina. 
A and.B, Transverse sections of foliage-leaves. C and D, Portions of 
the epidermis of the upper and of the lower surface respectively. (West 
Greenland) HA Fandt otaku EEC Kande Dab out e) 


elements, however, are always found enclosed in the stereom 
(well-illustrated in HoveLAaQUE's Fig. 334, p. 405). 

Medullary rays do not occur. The epidermis of the rhi- 
zome persists for a long time; cork is not developed (Hor- 
STEIN); in the older rhizomes a successive dying-away of the 
outer layers of the cortex takes place. f 

The Leaf. The epidermal cells of both the upper and 
the lower surface have undulating lateral walls, this being 
somewhat more strongly the case with those of the lower 
surface than with those of the upper (Fig. 18, C and D). 


Scrophulariaceae. 419 


Stomata occur on both leaf-surfaces, but they are more 
numerous on the lower surface; the proportion between the 
number of stomata on both surfaces proved to be about 1 
to 2. On the upper surface the lateral walls of the epidermal 
cells are slightly porose, especially near stomata; the guard- 
cells are surrounded by 3—6 cells, and are on a level with 
the surface, or project slightly (Fig. 18, A and BJ). 

The transverse section (Fig. 18, A and B) shows 1—3 
palisade-layers. Both A and B were drawn from sections 
of leaves taken from specimens from West Greenland, and 
in both cases the leaves were chosen from the middle of the 
stem. Å is seen to be somewhat thicker than B, and to have 
an assimilatory tissue consisting of as many as 3 cell-layers, 
the elements of this tissue are somewhat irregular and 
some of them are slightly branched; B probably represents 
the more common type. The transverse section of the leaves 
of the Norwegian plants resembles, on the whole, the trans- 
verse section shown in B, with the exception that the pali- 
sade-cells are slightly higher. The spongy parenchyma con- 
sists of rather copiously-branched cells. In the palisade- 
tissue and spongy paårenchyma chlorophyll-grains are pre- 
sent in abundance. 

The leaves are rather densely covered with hairs; the 
following types of hair occur: — 

(1) Non-glandular hairs, with one or many cells, thin- 
walled, glabrous and pointed (Fig. 19, C and E); (2) Glan- 
dular hairs with longer, two- or many-celled stalk and a 
2—4 celled head (Fig. 19, A, B and D); (3) Glandular hairs 
with short, one-celled stalk and 2 or commonly 4-celled head 
(Fig. 19, F); (4) Glandular hairs of the kind illustrated in 
Fig. 19, G, H, I and K with a low somewhat sunken stalk- 
cell, and a cupola-like extremely thin-walled 4-celled head. 

By comparision of the figures in question with Fig. 14. 


420 Fr. J. MATHIESEN. 


C, D and E, it becomes evident, that this form of glands 
in Bartschia — as already pointed out by HEINRICHER (cf. 
his addition to the cited paper of ScHERFFEL) — belongs 
to the Lathraea-type like the similar ones in Euphrasia. Fig. 
19, I and K show these glandular hairs seen from above; 





Fig. 19. Bartschia alpina. 
Different types of non-glandular and glandular hairs from the leaf. 
(A—E, about ?%/,; F—K, about. 299/,,) 


below the stalk-cell the lateral walls of the epidermal cells 
are indicated by dots, in K two intercellular spaces are formed 
between them, 2—4 of these are found rather commonly 
under the hairs. A glandular hair, similar to that which is 
seen from above in K, is shown in M in transverse section, 
here also 2 intercellular spaces are distinctly seen. G and I 
have no intercellular spaces of this kind. Glandular hairs of 
type 4 occur only on the lower surface of the leaf, under 
the branches of the veins of higher order; in the same place 
there is found in addition to this a great number of hair 


Scrophulariaceae. 421 


of type 3, but the latter kind of hair is also found on the 
upper surface of the leaf, located at the margin of the leaf 
in the innermost part of its indentations, i. e. above the 
extreme points of the veins (cf. Fig. 15, B). The non-glan- 
dular hairs and the long-stalked glandular hairs occur on 
both leaf-surfaces, and they grow most densely near or upon 
the veins. 

Upon the scale-leaves at the base of the shoots glandular 
hairs of the type shown in Fig. 19, F predominate, they are 
especially numerous on the lower surface, where the majo- 
rity of them are found arranged in two broad stripes, one 
along each of the margins of the leaf; mingled with them 
are found glandular hairs of type 4. 

The stems are covered with multicellular non-glandular 
hairs, and glandular hairs which frequently have multicellular 
stalks. 


Pedicularis lapponica L. 


Alcohol-material from Norway (Aursunden, leg. TH. R. 
REesvoLL, 8. 8. 1918; Skådavara in West Finmark, leg. E. 
WARMING, 7. 7. 1885). 

Herbarium-material from West and East Greenland, 
Lapponia murmanica, Nova Zembla, Labrador, Arctic Sibe- 
ria and the coast of Hudson Bay. 


Lit.: AxELL, 1869, p. 102; LANGE, 1871, pp. 256 and 265, tab. 
III, fig. 16 (the seed); 1880, p. 74; 1887, p. 262; WARMING, 1886, 
ppi 44Fand 47:11888 pp 357 39759and 8751890) pp..207, 208, 210, 
211 and 219; Linnman, 1887, p. 84; RosENVINGE, 1892, p. 686; 1896 
(I), p. 68; 1896 (II), p. 78; HarTz, 1894, pp. 37 and 39; 1895 (I), pp. 137, 
175, 219, 271 and 288; 1895 (II), pp. 335, 359 and 372; HAarTz and 
KRUUSE, 1911, pp. 347, 359, 365, 388, 409, 419, 423 and 428; EKSTAM, 
1897, pp. 119, 167 and 168; Kruuse, 1898, pp. 373, 380 and 394; 
1905, p. 175; ABROMEIT, 1899, p. 42; Knurtx, 1899, p. 190; Dusén, 
1901, p. 39; Norman, 1901, p. 457; CLEvE, 1901, pp. 9, 39, 57, 70, 
71, 76 and 89; SKOTTSBERG, 1901, p. 8; Porsizn, 1902, pp. 114, 167, 
175 and 187; 1910, pp. 267 and 271; 1912, pp. 382 and 387; 1920, 


422 Fr. J. MATHIESEN. 


p. 142; Porrius, 1903, p. 48; SyLven, 1905, p. 88; SILÉN, 1906,.p. 92; 
Simmons, 1906, p. 34. 


Hemicrytophyte, with a primary root which dies away 
rather quickly; it spreads by means of slender, subterranean 
runners. Ås a rule, these are monopodial: their short ver- 
tical portion bears leaf-rosettes, usually through a limited 
number of growth-periods (1—3), the floral shoots arising 
as lateral axes on the rosette-axis. 

As shown in Fig. 20, A and B the horizontal portion 
of the runners has elongated internodes, with small scale- 
leaves; in the axils of these, buds are formed which grow 
out into new runners. New runners arise only in the hori- 
zontal portion of the parent-shoot. The runners are provided 
with adventitious roots, which must be described as ”re- 
stricted to one position only”, inasmuch as they arise in 
connection with the axils of the scale-leaves, as a rule one 
at each. Haustoria were found on the roots. 

When the runner bends upwards, its internodes become 
short, and the scale-leaves pass without intermediate forms, 
into long-stalked foliage-leaves, of which it bears 3—7 in 
the first year (at x in Fig. 20, A and B the dead stalks of 
the rosette-leaves of the previous year are seen); at the end 
of the season of growth a number of scale-leaves are again 
formed, which function as bud-scales for the winter-bud. 
Some runners pass over directly into their winter-rest, with- 
out having succeeded in forming leaf-rosettes in the first year. 

Both the scale-leaves under the rosettes, and the rosette- 
leaves, as well as the scales of the winter-bud, can subtend 
buds (Fig. 20, 4, B and D); the lowermost buds must prob- 
ably be regarded as reserve-buds; any true principal bud 
does not occur. 

At the beginning of the next season of growth, one or 


several — as many as 3 — of the buds can grow out into 


Scrophulariaceae. 423 





Fig. 20. Pedicularis lapponica. 
A, Å part of a flowering plant. On the runner two new runners (a 
and b) have developed as lateral shoots, and an erect, floral shoot, 
of which only the lower half has been figured; », adventitious roots. 
The uppermost of the two buds seen just below the floral shoot is the 
terminal bud of the runner. The runner a is in its second year of growth; 
the withered remains of the rosette-leaves (7) of the previous year 
are seen under the scales of the winter-bud. B, The apex of a runner; 
under the leaf-rosette one sees the scales of the winter-bud, — which 
scales subtend small buds, — and the withered remains of the foliage- 
leaves (7) of the previous year; a floral lateral shoot has been devel- 
oped. C and D, Apices of two runners; C with dead, terminal bud 


494 Fr. J. MATHIESEN. 


and two lateral shoots (floral), D with a floral lateral shoot and four 

reserve-buds. E, One of the leaves of a floral shoot. (Å and E, Green- 

land.) A and B about nat. size; C and D about ”/,; E about Z/,. (B,C 
and D are drawn from sketches by E. WARMING.) 


floral shoots; as shown in the figures, they bear scale-leaves 
(ex parte bud-scales) at their base; these pass gradually into 
foliage-leaves in the middle of the shoot. The foliage-leaves 
of the floral axes differ somewhat in form from the rosette- 
leaves; the midrib is broader, and the incisions of the margin 
are less deep (Fig. 20, E). The inflorescence is a crowded, 
almost capitulate raceme, without a terminal flower; accord- 
ing to WARMING, the flowers in the middle of the inflores- 
cence, are the first to expand. The terminal bud of the run- 
ner, as a rule, continues its growth by forming a second leaf- 
rosette; exceptionally, it may produce a floral shoot, which 
consequently becomes terminal; the latter has only scale- 
leaves at its base. In addition I must remark, that in a 
single case, I found a runner which, without a previous 
vegetative-stage (leaf-rosette), terminated directly in an in- 
florescence. 

Å runner hardly ever forms leaf-rosettes for more than 
3 successive growth-periods. Also in the axils of the scale- 
leaves of the floral shoots buds occur; in cases in which these 
develop further, they frequently form, in the following year, 
shoots of similar structure to the parent-shoot. WARMING 
mentions that at the base of the floral shoots also, rosettes 
of foliage-leaves may be produced; in some cases I suc- 
ceeded in demonstrating such leaf-rosettes, but they are 
evidently of rare occurrence. 

The structure and biology of the flower are explained 
by LinnmMaNn and E. WarmiInG, of whose descriptions the 
following is an extract: The lower lip of the almost horizont- 


ally projecting flower incline very decidedly to the right 


- 


Scrophulariaceae. 425 


(seen from the front); besides this, the whole flower is subject 
to torsion around its axis to the right; this causes the upper 
lip to slope even more to the right, while its helmet-shaped 
apex, and the part of the style turned downwards, point 
to the left; the lower lip stands almost in the vertical plane 
(Lindnman, p. 84, Pl. IV, fig. 47, Å and DB). 

"The calyx is dark-red, the corolla pale yellow or brim- 
stone-coloured. On the lower lip there åre two conspicuous 
convexities covered with 
hairs, the corolla-tube is 
6—8 mm long” ... "The 
slit of the upper lip is 
about 1—17/, mm wide; 
the front stamens are 





slightly hairy or glabrous. 


Fig. 21. Pedicularis lapponica. 
The style may be enclosed  A flower with a far-protruding style. 


(and this is the case at the (Greenland, about 67? N. lat.) 
(E. WARMING, 1890.) 


same time as the anthers 
are open), but usually it protrudes more or less, from 
1.5 to 2.5 mm or even more (without being moved away 
from its natural position up to the dorsal side of the upper 
lip). Even in the bud it protrudes somewhat, so that the 
stigma comes into contact with the lower lip; the former 
is even at that time papillose, and appears ripe, consequently, 
slight protogyny appears to prevail. Afterwards the anthers 
are seen to have dehisced, without the length of the style 
having altered; consequently, self-pollination appears to be 
able to take place, but not easily, because although it is 
true that the anthers come to stand vertically above the 
stigma, yet the distance between them is comparatively 
great”. (WARMING, p. 219, fig. 33, reproduced here as Fig. 21). 

All authors join in attributing perfume to the flowers 
of P. lapponica; LIiNDMAN mentions its "strong perfume”, 


426 Fr. J. MATHIESEN. 


and Hartz (1895 (I), p. 271) writes, that P. lapponica and 
Viscaria filled the air with perfume (East Greenland, 7. 7.92). 

Pollination takes place by the agency of humble-bees, 
which, according to LINDMAN, alight on the upwardly turned, 
left side of the lower lip; the stigma is consequently, by 
the torsion described above, turned towards the visitor. ÅAc- 
cording to the same author, the humble-bee thrusts its pro- 
boscis into the flower, at the widest part of the slit of the 
upper lip, at about the middle of it, just above the revolute 
portion of the edge of the upper lip, but not into the narrow 
groove between the two convexities of the lower lip. 

Several of the specimens from West Greenland had one- 
year-old fruits, from East Greenland I saw fruit-bearing 
specimens from Danmarks Ø (N. Hartz leg.). 

According to PorsiLn (in litt.) visits of humble-bees are 
rare, and fruit-setting (in contradistinction to the other Pedi- 
cularis spp.) is generally not good; many of the capsules are 
empty, and in others only a few seeds are found. EKSTAM's 
observations from Nova Zembla agree with those of PoRrsiILD. 

A description and beautiful illustration (Pl. III, Fig. 16) 
of the seed is found in LANGE, 1870. 

P. Lapponica is a decidedly middle-summer-flowering 
species. 

Geographical Distribution: Greenland, Arctic Ame- 
rica, Labrador, Kamschatka, Arctic Siberia, Nova Zembla, 
Arctic Russia, Scandinavia and Lapland. RosENVINGE (1892) 
records, that in West Greenland it is rather common north 
of 64" N. lat., but very rare south of this. "The northern 
limit im West Greenland is still unknown, in the fjords at 
72723" it was so commonly distributed that the northern 
limit hardly can be here” (Porsirn, 1920). In East Green- 
land the species is only found between 69725" and 73710' 
Naar m(Kru use 9 0S pE] 


Scrophulariaceae. 427 


Habitat: In Greenland it grows in mossy bogs, and 
-m damp places on heaths; the runners creep in the moss, 
or in slightly decomposed vegetable matter (M. PorsiLD, in 
litt.); it is "covered by thick layers of snow during winter” 
(Porsirn, 1920). WARMING, 1888, mentions it as inhabiting 
willow copses, herb-slopes and heaths. On Nova Zembla Ex- 
STAM found it growing in dry localities, which were exposed 
to strong insolation. A. CLEVE finds it on mountain heights 
im Lule Lappmark, avoiding wet ground and the poorest 
heath; she writes: "Sie 
gedeiht am besten in 
den Waldregionen und 







sucht im Hochgebirge 
relativ trockene Stand- 


GES 


orte auf, was schon 





Wahlenberg aufgefallen 






me 
DS ADED: 








war.” 







SS 


CE 


Anatomy. The 
Root: The structure is 
shown in the transverse 
section figured in Fig. 22. 


The secondary growth Fig. 22. Pedicularis lapponica. 
Transverse section of an adventitious 


is not considerable, and MOE bore) 


does not greatly exceed 

what is shown in the figure in question. The endodermis 
is a very beautifully developed Casparian sheath. The epi- 
dermis dies away quickly; the outer walls of the exodermis 
were found to be cuticularised. On the surface of the roots, 
dark-coloured, fungal hyphæ were frequently found. 

The Stem: (1) Runners: The epidermal cells have 
farly thick outer walls; they are filled with a homogeneous 
brown mass, which gave the reaction of tannin. The outer- 
most cortical layer adjoins the epidermis without inter- 


428 Fr. J. MATHIESEN. 


cellular spaces; the cells of the cortex show radial divisions 
by thin walls of secondary formation. The endodermis is 
exactly as in the root. The pericycle is 4—7 layered, its 
cells, like those of the cortex, have somewhat thickened 
walls. The wood-cylinder is continuous, without parenchyma- 
rays, but is often somewhat excentrically developed, so that 
the wood on that side of the shoots which is turned down- 


LED 
RER 
DE 


mm! 
il 
ca) 
QYY 
HI 
07 
CT 





Fig. 23. Pedicularis lapponica. 
A, Portion of transverse section of runner. B, Portion of transverse 
section of flower-bearing stem. (Greenland.) (About 119/,,) 


wards, is thinner than on the upturned side. As shown in 
Fig. 23, Å, annual rings can be formed in the runners. The 
two annual rings are separated by thin-walled parenchyma, 
mixed with small vessels, and the vessels are much larger 
in the second than in the first annual ring. Some stereom 
(wood-fibres) is seen, especially in the inner annual ring. 
(2) The Flower-bearing stem (Fig. 23, Bb) differs from 
the runners in that the cells of its cortex and of its only 
1—2 layered pericycle, are much thinner-walled than in the 
runners, moreover, by the Casparian spots in the endodermis 
being much fainter, by the narrower vessels of the xylem- 


Scrophulariaceae. 429 


ring, and the almost continuous ring of wood-fibres on the 
outer side of the latter, and by the outér layers of cells of 
the pith being thickened and lignified. The central part of 
the pith has died away, and the stem is consequently hol- 
low. Stomata occur in the thick-walled epidermis, and chloro- 





Fig. 24. Pedicularis lapponica. 
A, Transverse section of the leaf. B, Epidermis of the upper, and C 
of the lower surface of the leaf. (Greenland.) (A. about ""%/,; B and C 
abon 5R) 


phyll in the cortical cells. In the cases in which the basal 
portions of the flowering-shoots persist and bear lateral shoots, 
a second growth-ring —often excentric— may be formed here. 

The Leaf: As regards the anatomy, there appears to 
be very little difference between the rosette-leaves and the 
leaves of the floral shoots. The epidermis of the upper sur- 
face (Fig. 24, B) has straight to slightly undulating lateral 


XX VIL: 28 


430 Fr. J. MATHIESEN. 


walls; those of the epidermis of the lower surface (Fig. 24, C) 
are more strongly undulating. The lateral walls of the epi- 
dermal cells of both the upper and lower surface, although 
thin, are yet very distinctly porose, those of the upper sur- 
face, more strongly so. Stomata occur only upon the lower 
surface, the guard-cells are surrounded by 4—f6 cells. In trans- 
verse section (Fig. 24, A) 1—2 
layers of palisade-cells are seen ; 
the spongy parenchyma is 
composed of highly-branched 


ef lt UABRADI 


OBS GAM 


cells (the small circles inside 
the cells of the spongy paren- 
chyma indicate the branches 


1, 


which have been cut through 
on preparing the section); 
chlorophyll-grains are present 





OMA IMAM) DU, 


abundantly in the whole of 
Fig. 25.  Pedicularis lapponica. 
A portion of a leaf-section seen 
from the lower surface: the  dermis of the lower surface; 


glandular hairs are seen to be +fhe stomata occur on a level 
crowded under the veins (about É Å j 
35/,). with, or are raised slightly 


the mesophyll, also in the epi- 


” above, the leaf-surface. The 
final branchings of the veins end in an epithema-like tissue 
of somewhat inflated, thin-walled tracheids. 

Very characteristic of the leaves of the Pedicularis spp. 
are the numerous glandular hairs of the type shown in 
Fig. 24, A and C. Fig. 25 shows a portion of a leaf-section 
seen from the lower surface,"— the glandular hairs are found 
here only —, the veins are drawn as if visible through the 
mesophyll, and the glandular hairs are seen to be crowded 
under the veins, the same feature is recorded by HOVELACQUE 
(1. c.) in the case of other species of Pedicularis. In Fig. 24, C 
three glandular hairs are seen from above; the epidermal 


Scrophulariaceae. 431 


cells from which they have developed, have far thicker and 
more highly porose lateral walls than the surrounding cells; 
the finely-drawn circle which surrounds the thickly-drawn 
glandular-head, indicates the place of contact between the 
basal cell of the hair, and the outer wall of the epidermal 
cell; the smaller circle at the top of the head indicates that 
the cuticle at this place is lacking, thus forming a pore 
only spanned by the cellulose wall, as described by PERCY 
Groom for P. palustris. 


Pedicularis sudetica Willd. 


Herbarium-material from the shore of Hudson Bay 
(Churchill), King Point, King William's Land, Southhampton 
Island, Port Clarence, Taimyr Peninsula, Chabarowa, Nova 
Zembla and Lapland. 


Lit.: LANGE, 1880, p. 75; KJELLMAN, 1882 (I), p. 256; 1882 (II), 
p. 325; 1882 (III), pp. 361, 363 and 366; 1882 (IV), p. 509; KJELLMAN 
and LuNDSTROM, 1882, p. 303; HorLm, 1885, p. 43, tab. VI, fig. 10, 
tab. VII, figs. 1—7; WARMING, 1890, pp. 207 and 215; Egxstam, 1897, 
pp. 118, 166, 168, 176 and 180; Knurtx, 1899, p. 191; Simmons, 1913, 
pp. 123 and 138. 


Sympodial hemicryptophyte of semi or entirely rosette 
type, the above-ground part of the shoot bearing the in- 
florescence being either furnished with a few foliage-leaves, 
or being totally devoid of them. 

The species may form tufts, (a specimen from Herschell 
Island belonging to forma /anata Walpers, had numerous 
rosettes of foliage-leaves, even seven flowering shoots and 
10 withered peduncles from previous years) and in this case 
it has a short mesocorme (”rThizoma multiceps”), a fairly 
thick primary root, sometimes probably of long duration, 
adventitious roots being only few in number, but rather 
vigorous; more frequently it appears, however, that the 
basal portion of the main shoot dies away, perhaps even 


28% 


432 Fr. J. MATHIESEN. 


after the first flowering period; the main root is then less 
vigorously developed, and slender adventitious roots, also from 
the rhizome-portions of the innovation shoots, are abundantly 
developed, a circumstance which permits those shoots to 
continue their existence as independent individuals, after iso- 
lation from the main shoot — consequently, the species can 
be propagated vegetatively (cf. Ekstam). 

This difference in the growth is probably conditioned 
by the nature of the habitat; the tufted individuals must 
be assumed to have grown on drier soil, while the other 
form must have lived in damper localities; the fact of my 
having found Sphagnum in connection with the rhizomes of 
many specimens of the latter type, indicates moist moss- 
tufts as their habitats. 

As regards the duration of the first vegetative stage, it 
is not possible for me to state anything with certainty, owing 
to want of material; in a comparatively weakly-developed 
individual, I estimated its extent as three years. 

Fig. 26, 1 shows an individual produced from an iso- 
lated innovation shoot; (the figure has been drawn from 
soaked herbarium-material, and is somewhat diagrammatic). 
At the base of the relatively-main axis, we see the separation 
surface (scar) along which it has loosened itself from the 
parent-shoot; during its first year, the shoot has formed a 
few-leaved rosette of foliage-leaves, and completed the year's 
growth by the formation of scale-leaves, for the protection 
of the winter-bud; (at 7 some fragments of the first year's 
leaves are seen; the bud A is dead). Next year a piece of 
stem, about 1 cm long and only bearing 2 leaves, was first 
produced, next a rosette of foliage-leaves, followed in turn 
by scale-leaves, of which three still remain (two only can 
be seen). The process is repeated in the following (3rd) year, 


and simultaneously the buds C and D in the axils of the 


Scrophulariaceae. 433 


scale-leaves of the 2nd year, expand and produce a few foliage- 
leaves, some of which still remain in a withered condition 
(ce; and d; respectively). In the fourth year the individual 
at last flowered; /V indicates the lower portion of the pedun- 
cle. C and D which, like the terminal bud of the shoot, 





Fig. 26. Pedicularis sudetica. 
1, Basal portion of a plant, somewhat diagrammatic; as regards the 
explanation of letters and numbers cf. text on pp. 432—434. 2, A flower 
seen in side view. 3, The upper lip seen in side view; the reciprocal 
position of the anthers and pistil is indicated. 4, Stigma, more highly 
magnified. (Nova Zembla.) (1 about nat. size; 2 and 3 about Z/; 
4 about 59/,.) 


have been protected during the winter-rest by scale-leaves, 
are forming their second leaf-rosette, the leaves of which are 
indicated by c, and d, respectively; and B, which rested during 
the previous summer, has now developed a foliage-leaf, bs; 


434 Fr. J. MATHIESEN. 


the now dead leaves x and y are from the leaf-rosette formed 
the year before the main shoot flowered; from the axil of one 
of the uppermost bud-scales, an innovation shoot Æ has also 
been developed, and is expanding its first two foliage-leaves. 

Here, consequently, the vegetative-stage has lasted 3 
years, and this appears to be the rule in individuals of this 
type; in the tuft-forming type in which, in dried material, 
this question is far more difficult to decide, it appears to me 
that the vegetative stage can be restricted to two years. 

It still remains to be noted that, although the majority 
of the individuals which result from isolated innovation 
shoots, had elongated internodes inserted between the ro- 
settes, yet only rarely were they as long as is shown in Fig. 
26, 1; the phenomenon is probably due toY"the endeavour of 
the plant to keep its terminal bud in a certain position rela- 
tive to the surface of the substratum. In Fig. 26, 1 adventi- 
tious roots are developed only at the base of the shoot, but 
these may also be developed below the other rosettes, often 
in great numbers. They are 1—2 mm thick. I never found 
haustoria on such adventitious roots, but they occurred on 
the root-branches in the specimen from Herschell Island, 
mentioned above. 

From Nova Zembla plants were to hand of which the 
peduncle was only 5 cm high during flowering; in specimens 
from Chabarowa it was 17 cm, and in one from Churchill, 
even 25 cm high; after flowering an elongation of the axis 
of the inflorescence takes place. 

Fig. 26, 2 shows a flower seen from the side. The posi- 
tion of the stigma should be noted, and the two pointed 
lobes in which the front part of the helmet terminates, and 
the ridge, with the veins which proceed from it into the 
helmet, also the slightly warty revolute portion of the edge 
of the upper lip, and the two convexities on the lower lip; 


Scrophulariaceae. 435 


as stated by E. WARMING, the latter is somewhat oblique. 
The calyx is drawn as densely hairy, this is the case in forma 
lanata Walpers. In Fig. 26, 3 the position of the stamens 
inside the helmet is shown. It must be borne in mind that the 
natural position of the flower is almost horizontal, whence it 
follows that self-pollination must be able to take place with 
comparative ease, for the pollen will be sprinkled out of the 
front opening of the helmet by accidental movements of the 
plant, caused for instance by the wind, and will fall upon 
the hinder side of the stigma, which, as shown in Fig. 26, 4 
is almost globular. 

Ekstam has studied the species in Nova Zembla, and 
he writes concerning this: "Ziemlich håufig — håufig an beson- 
ders nassen Orten. Besitzt einen ziemlich starken jasmin- 
åhnlichen Wohlgeruch und hell- bis dunkelrote Bliten. 
Schon in sehr jungen Knospen ragt die Narbe aus der Ober- 
lippe hervor — in einer ganz aufgebluihten Blume etwa 
21/,—3 mm — und wird bald papillås, gewohnlich bevor die 
Blume ganz aufgebluht ist, und die Antheren sich gedffnet 
haben. Selbstbeståubung durfte dadurch ermåglicht werden, 
dass die kugeliche Narbe rund umher papillås ist, und in 
der Falllinie der Pollenkårner gelegen ist. Nachdem die An- 
theren sich entstaubt haben, bleibt die Narbe noch eine Zeit 
lang glånzend, papillås. Bluth im Hochsommer. Mit reifen 
Fruchten beobachtet. Anemophile Samenverbreitung. .... 
Wurde am 3/, von einer kleinen Fliege, am &/, von einem 
grossen Bombus hyperboreus Schonh., am 72/, ebenfalls von 
einem grossen Bombus hyperboreus Schonh. besucht. Im 
Sommer 1891 trotz fleissiger Beobachtung kein einziger In- 
sektenbesuch bemerkt.” 

In somewhat vigorous specimens, the exceedingly dense 
inflorescence, with the deep-red corollas, cannot fail to be 
very Cconspicuous. 


436 Fr. J. MATHIESEN. 


The Geographical Distribution, according to SIM- 
MONS and LANGE: Western Arctic and Subarcetie North Ame- 
rica, (both the continent and the archipelago), Arctic Siberia, 
Nova Zembla, Arctic Russia and besides on mountains 









VI33 
== 
1 
















sæ 
= TE 





Fig. 27. Pedicularis sudetica. 
A and B, Portions of transverse sections of the rhizome of the individ- 
ual illustrated in fig 26, 7 (cf. text). C, Epidermis and the outer layers 
of the cortex of the same rhizome. D, Transverse section of the leaf. 
(Nova Zembla,) AA and B'about "97,0 "about. 7"; D about 11979) 


(Sudeten and Riesengebirge) of Central Europe. Does not 
occur in Greenland. 

According to KJELLMAN (1882 (III)) the fu 7ering year's- 
shoots of P. sudetica in a pressed and fernij: …d condition, 
and the roots, rhizomes and buds in fresh condition, are 
used as food by the Chuckes in Eastern Arctic Siberia. 

Anatomy. Ån adventitious root about 2 mm thick 


Scrophulariaceae. 437 


showed the following structure: Of the exceedingly thin- 
walled epidermis only scattered fragments were left, the cells 
were dead and had collapsed. The outermost layer of the 
cortex had cuticularised outer and lateral walls (it must 
certainly be this layer which TH. Horm in his Fig. 7, Tab. TIT, 
designates epidermis); the layer just within the outermost 
layer adjoins the latter without or with small intercellular 
spaces between them; the endodermis has distinct Casparian 
dots. The intervening part of the cortex consists of a very 
lacunose tissue, which corresponds exactly with that in the 
rhizome (shown in Fig. 27, C). Such slender adventitious 
roots complete their development during one season of growth; 
their structure is 2-rayed, and by the secondary growth 
there is formed a bundle of xylem-tissue, about 0.5 mm thick 
consisting of vessels with intervening non-lignified tissue. 

Of the individual figured in Fig. 26, 1 a transverse 
section of the axis was taken from the part between I 
and 77, and- also one between IM and III; the former 
part must be assumed to be 3 years old, and the latter 
2 years. In the part between / and II, there was found 
in the stele a circle of vascular bundles, separated by 
broad, non-lignified, parenchyma-rays (cf. Fig. 27, A, which 
shows the xylem part of a vascular bundle). In the second 
year of growth an increase of the vascular elements has 
evidently taken place, the lowermost (i. e. in the stem inner- 
most) group of wide vessels probably indicates the limit of 
the first year's growth; what is found outside that belongs 
then to a later (probably the second) year. Wood-fibres are 
quite absent:  ”e the pith does not die away. Fig. 27, Bis 
drawn from a. on taken from the part between // and III, 
at about the middle of it. Here the axis is hollow, because 
the central part of the pith, as in the peduncle, has died 
away; the walls of the outermost layers of the pith have 


438 Fr. J. MATHIESEN. 


become thickened, lignified and porose, the same is seen to 
be the case as regards the inner part of the parenchyma- 
rays, which separate the vascular bundles. Around the ves- 
sels non-lignified elements occur, but the limit of the first- 
year's growth is shown in the vascular bundles by a layer 
of wood-fibres, 2—3 cells thick, which adjoins the lignified 
parenchyma rays, so that a continuous stereom results. The 
few scattered vessels outside this I regard as the growth of 
the second year; such small groups of vessels may also occur 
in the space between any two of the original vascular bundles. 
A second individual exhibited a similar structure, but in a 
third I found no trace of a second-year's growth; unfortu- 
nately the scantiness of the material prevented my studying 
this point more thoroughly. Å portion of the cortex is shown 
in Fig. 27, C; with the exception of the two outermost layers 
it is very lacunose in structure. The epidermis is small- 
celled, and furnished with a rather thick cuticle. 

The Leaf. The epidermis is of the usual Pedicularis- 
type, the lateral walls of the cells are thin and only slightly 
porose, very similar to those in P. hirsuta and lanata. Sto- 
mata occur only on the lower surface of the leaf; the guard- 
cells are on a level with the other epidermal cells. Å trans- 
verse section is shown in Fig. 27, D; the part which has 
been illustrated is typical and also agrees well with TH. 
Horm's Fig. 4, Tab. VII. The upper half of the mesophyll 
consists of 2—3 layers of short, thick palisade-cells, and the 
lower half of spongy parenchyma composed of rather deci- 
dedly branched cells. Chlorophyll occurs abundantly in the 
whole of the mesophyll. Glandular hairs of the same form 
and size as in the other species of Pedicularis, occur also 
in P. sudetica on the lower surface of the leaf — not as TH. 
Horm (l. c. p. 44) writes "here and there,” but in abun- 
dance below the branches of the veins of tertiary and higher 


Scrophulariaceae. 439 


order. On the lower surface of the leaf, and especially in 
the middle of the sections, thin-walled 1- to 3-celled non- 


glandular hairs occur. 


Pedicularis euphrasioides Steph. 


Alcohol-material from Greenland (Itivnek near Hol- 
steinsborg, leg. EuG. WARMING, 13. 7. 1884). 

Herbarium-material from Greenland, the shore of Hud- 
son Bay (Churchill), Port Clarence near Behrings Strait, and 
Siberia (by the Boganida River). 

Lit.: LANGE, 1871, pp. 254 et seq, 264, tab. II, fig. 10 (the seed); 
1880, p. 74; 1887, p. 262; WARMING, 1886, pp. 19, 44 and 58; 1890, 
pp. 206, 210, 211 and 218; RosENVvINGE, 1892, p. 686; 1896 (I), p. 68; 
TEGEL Sp 78 ET ARTE, 1694 "p: 3977 KNUTE, 1899, "p-190FPORSIED, 
1920 p. 142. 

Spot-bound hemicryptophyte of the semi-rosette type, 
biennial to pluriennial hapaxanth, or — what is certainly 
the most common — a pollacanth, with scantily-branched 
main root, the longevity of which corresponds with that of 
the individual. Should the species appear as a pollacanth, 
innovation shoots are developed at the base of the flowering 
shoot. | 

The length of the first vegetative-stage varies from one 
to (probably at most) two years. During the vegetative 
stage a leaf-rosette is developed every summer and — towards 
the end of the growth-period — bud-scales for the protection 
of the young organs during the winter-rest; in the summer 
in which the plant flowers, the shoot forms no rosette of 
foliage-leaves. The shorter vegetative-stage (of one-year's 
duration) appears especially to result in weaker individuals, 
which probably often end their lives by fruit-setting in the 
second year. 

This species differs very much in habit from the other 
species described here, owing to the formation of numerous 


440 Fr. J. MATHIESEN. 


branches which, during the flowering year, arise in the axils 
of the foliage-leaves on that part of the stem which has elong- 
ated internodes. These lateral shoots increase in length down- 
wards to some way below the middle of the parent-shoot, 
and then again become shorter towards its base; here they 
are often purely vegetative, while the upper ones bear, in 
addition to 2—3 decussated pairs of foliage-leaves, a small 
inflorescence corresponding with that of the parent-shoot. 

In the pollacanthous individuals the innovation shoots 
are produced in the axils of the uppermost bud-scales; they 
usually develop only a few pairs of opposite foliage-leaves, 
then bud-scales, and then the following year, they most 
often terminate their growth by the formation of an in- 
florescence. I cannot deny that there is a possibility of the 
innovation shoots also flowering without any preceding 
rosette-formation, and therefore, in their first year, develop- 
ing only bud-scales; in this case the species would be a 
transitional form between the semi-rosette type and the 
protohemicryptophyte-type. 

In individuals in which a ”rhizoma multiceps” is devel- 
oped, the uppermost part of the main root may become as 
much as 2 cm thick. 

The flower-bearing stem may become as much as 25 cm 
high; after flowering, the axis of the inflorescence is some- 
what elongated. 

On the basis of material from Greenland (about 67” 
N. lat.) E. WARMING gives the following description of the 
structure of the flower (cf. Fig. 28): "This species, which is 
in no wise common in Greenland, is very conspicuous owing 
to its numerous inflorescences with yellow, fragant flowers; 
the tip and the helmet of the yellow corolla is slightly brown- 
ish. The flowers are very oblique, the lower lip being highly 
oblique (Fig. 28, A, B, D and H); but the upper lip also is 


Scrophulariaceae. 441 


twisted in such a manner that its two-toothed tip turns 
somewhat to the right (E). The ridge reaches the edge slightly 
above the beginning of its revolute portion (ÅA and G). The 
slit between the two edges of the upper lip is usually rather 
narrow. The front filaments are sparingly hairy. The stigma 
protrudes more or less above the two teeth (B, D, E and H). 





Fig. 28. Pedicularis euphrasioides. 

(From Greenland; about 67? N. lat.) 
A, B, A flower seen from the left and the right side; in A the position 
of the style and the stamen is indicated. C, Stigmatic papillæ. D, A 
flower in front view. E, The upper lip seen in front view. F, The 
ovary. G, The upper lip seen in side view; the venation is indicated. 
H shows the same as A. I, Transverse section of ovary. (Drawn by 

E. WARMING, 1890.) 


Judging from the structure of this almost horizontally- 
placed flower (A and B), I must assume that it is only with 
difficulty that it pollinates itself. It produces ripe fruit in 
Greenland at 67? N. lat.” 

Geographical Distribution. West Greenland be- 
tween 62”? (but only in a few localities south of 64 N. lat.) 


442 Fr. J. MATHIESEN. 


and 69730" N. lat. (ROSENVINGE, 1892), on the whole rather 
rare there; Labrador, the Rocky Mountains, the continent of 
Arctic America, (but not mentioned by Simmons from the 
Archipelago), Siberia and Dahuria. 

In Greenland it grows on luxuriant, damp heaths; the 
species requires a snow-covering during its winter-rest (M. 
Porsirn, 1920). 

Anatomy. The root is of the type described later on 
in P. hirsuta and lanata and, as in these, the secondary 
cortex is traversed by numerous radial clefts. There is, how- 
ever, a difference between P. euphrasioides and the two spe- 
cies in question as regards the nature of the xylem-part, 
because, in the two former it is composed of a large quantity 
of non-lignified tissue and of relatively few vessels only, 
whilst in P. euphrasioides one always finds in the root several 
concentric rings, each consisting of numerous vessels with 
intervening  lignified stereom  (thin-walled wood-fibres). 
These continuous rings of lignified elements, the outlines of 
which are however somewhat undulating, are separated by 
portions in which the vessels are not so wide or numerous, 
and are surrounded by non-lignified tissue; here clefts occur 
between the radiating lines of vessels. 

That these rings of vessels and wood-fibres really in- 
dicate zones of annual growths, is evident from the investiga- 
tion of some specimens chosen from among the individuals 
in my material, which had flowered for the first time: such 
as — judging from the leaf-scars — had had only a vegeta- 
tive-stage of one-year's duration (that is to say, flowered 
in the second year) showed two rings; others which, as far 
as I could see, had had a vegetative-stage of two-year's 
duration, showed three rings. The greatest number I suc- 
ceeded in demonstrating amounted to three. 

The cavity of the stem is comparatively large. The 


Scrophulariaceae. 443 


cortex is thin; in the xylem-ring the wood-fibres dominate 
in the 3—4 outermost layers; the cell-layers in the periphery 
of the pith are lignified, thickened and porose. On the epi- 
dermis there are found non-glandular hairs of the same type 
as on the leaves. 

The Leaf. A transverse section of one of the leaves 





Fig. 29. Pedicularis euphrasioides. 
A, Epidermis of the upper, and B of the lower surface of the leaf. 
C, Transverse section of the leaf. D and E, Non-glandular hairs from 
the surface of the leaf. (Greenland.) (A and B about ?2/,; C about 
Ho SEDFandeE abort 0/78) 


from the middle of the stem, is shown in Fig. 29, C. There 
is generally found only one layer of rather elongated pali- 
sade-cells, below these there is usually a layer of cells which, 
although elongated, are nevertheless branched (this is di- 
stinctly seen to the left in Fig. 29, C), and which must there- 
fore most naturally be included in the spongy parenchyma; 
the lower layers of the latter consists of richly-branched cells. 

The epidermal cells of the upper surface has thin, slightly 


444 Fr. J. MATHIESEN. 


undulating lateral walls, those of the lower surface are still 
more thimwalled, their lateral walls more highly undulating 
and, as in the epidermis of the upper surface, furnished with 
flange-like thickenings, which I found to be particularly well- 
developed in the bracts of the flowers, and, on the whole, 
in the upper leaves on the axis. Stomata occur only on the 
lower surface; the guard-cells are on a level with the other 
cells of the epidermis; they are surrounded by 3—6 cells. 
Chlorophyll is abundantly present in the whole of the meso- 
phyll and is also found in the epidermis of the lower surface. 
On the lower surface there are glandular hairs of the usual 
kind, and with the typical distribution; and besides there 
are non-glandular hairs both on the upper and the lower 
surface (Fig. 29, D and E); they are multicellular and thick- 
walled, often however with one or two cells unthickened and 
therefore collapsed. 


Pedicularis Sceptrum carolinum L. 


Alcohol material from Aursundséen, Norway (leg. TH. 
RESVOLL, 11. 8.1918).— Herbarium material from the district 
near the river Boganida, Alten, Mortensnæs in East Finmark, 
and the Murman Coast. 

Fit: LANGE, 1871 pp 254 et" seg: 2677 tab; TITS 25 (the 
seed); WARMING, 1890, p. 215; Norman, 1895, p. 456; KERNER, 1898, 
BASIE PS 655 KEN res VIE EO IE STEN ELO OS EPO 

Å sympodial hemicryptophyte of rosette- or semi- 
rosette type, with vertical or obliquely-ascending mesocorme, 
which dies away behind, and from which adventitious roots, 
1—2 mm thick, are abundantly developed. 

2-year-old specimens, from Aursundséen, showed that 
the plant in its first year forms a few-leaved rosette; the 
winter-bud is protected by scale-leaves, and the following 


year the growth is continued by the formation of a more 


Scrophulariaceae. 445 


luxuriant rosette. I can state nothing as regards the duration 
of the first vegetative-stage, the 2-year-old plants in question 
were still rather frail. The hypocotyl was somewhat elon- 
gated, the adventitious roots had already begun to develop, 
and in one of the individuals, a lateral shoot had been pro- 
duced in the axil of a rosette-leaf of the first year. 

On therhizomes,innovation-shoots, from 1 to 3 in number, 
occur; their vegetative stage, judging from the material at my 
disposal, which is however but scanty, extends over 2 years: 
During the first year 2—4 foliage-leaves are developed, and 
then scale-leaves, which serve as bud-scales for the winter- 
bud; during the next year a somewhat more dense rosette 
is developed, again succeeded by scale-leaves; and finally 
the shoot, during the 3Ird year, after having formed a lux- 
uriant rosette, terminates in an aerial, flower-bearing portion, 
which is often quite destitute of foliage-leaves, and may 
attain a height of even 60 cm. 

The fact that the rhizome dies away behind, in con- 
junction with the abundant development of adventitious 
roots, must allow of vegetative propagation, since the lateral 
shoots gradually become free and independent by the dying 
away of the parent rhizome. 

E. WARMING (I. c. p. 215 et seg, figs. 30 and 31) gives 
the following description of the structure and biology of the 
flower, on the basis of material from Kola, and from Øster- 
dalen in Norway; his two groups of figures are reproduced 
here as Fig. 30 and Fig. 31. 

The flowers are . . . . as much as 32 mm long; they 
stand erect, the subtending bracts are closely adpressed, and 
in addition each flower appears to be completely closed. The 
lower lip (Fig. E) is erect, 14—15 mm long, pressed closely 
to the upper lip, which is a little longer (16 mm), and closes 
the throat completely, but is easily pressed down. The visit- 


SERVE 29 


446 Fr. J. MATHIESEN. 


ing insects must evidently be large and vigorous, such as 
humble bees and moths, and they must, in a way similar 
to that in Antirrhinum and Linaria, press their body or head 
down between the two lips, by which means they primarily 
come into contact with the stigma; Fig. 30, B shows (the 
lower lip having been removed) that the stigma can be so 
protruding that contact with it must be regarded as inevit- 





Fig. 30. Pedicularis Sceptrum carolinum. 
From Kola near the White Sea (collected by Dr. BROTHERUS). 
A, Flower, natural size and position. B, The same after the lower lip 
has been removed (D belongs to this). C, Part of a 25 mm long flower, 
seen in front view; the stigma protrudes less than it does for instance 
in B; the anthers have not yet opened (see F); to this belongs the 
tip of the style shown in H. E, The lower lip of B. G, Ovary. v indi- 
cates the revolute portion of the edge of the upper lip. (E. WARMING, 
1890.) 


able; that cross-pollination must take place by the agency 
of large insects, is also evident from the fact that the distance 
from the stigma to the nectary may be as much as 2.5 cm. 
I have also several times seen the stamens torn off the upper 
lip, evidently by the violence of a large insect. In one of 
the flowers (C) which has been illustrated, the anthers had 


Scrophulariaceae. 


447 


not yet opened (F), and as the stigma appears to be ripe 


(H), protogyny must consequently prevail. When the anthers 


have afterwards opened (D) it must evidently be the insect 


knocking against the filaments, which shakes the pollen down 


over it; strange to say, both the fila- 
ments, anthers, style as well as the 
revolute portions of the edge of the 
upper lip, are quite glabrous (see B, C 
and D) so that there is no special contri- 
vance which by its resistance increases 
the shock. On the other hand, hairs 
occur — which are probably of im- 
portance in preventing the pollen-grains 
from falling out at the sides — viz., 
along the edges of the upper part of 
the upper lip (B, C, D), consequently, 
in the same place as ordinarily, but 
on another organ. 

For the rest, the width of the slit 
of the upper lip, and the degree to which 
the stigma protrudes, differ also. While 
the stigma protrudes far out in the 
quite young flower B, it does not pro- 
trude so far in the young C, and still 
less in Fig. 31 (also in this paper Fig. 
31) of which the anthers are already 
open . . .. Self-pollination appears to 
be able to take place only with diffi- 
culty. As the species sets fruit abun- 














Fig. 31. Pedicularis 
Sceptrum carolinum. 
From Koola. 
The upper lip in front 
and side view. The 
anthers are open. In 
the front there is a 
flat or incurved part, 
and the slit is tightly 
closed above. In this 
case the stigma does 
not protrude, but 
stands just within 
the uppermost end of 
the slit. (E.WARMING, 
1890.) 


dantly in so northerly a region as near Alten (about 70? N. 
lat.) it must undoubtedly be humble-bees that effect pollina- 


tion.” 


Conditions pertaining to the pollination of the flower 


29% 


448 Fr. J. MATHIESEN. 


have afterwards been studied by SILEN, who writes: "Were 
much-visited by humble-bees, which forced themselves into 
the ciosed flowers, which were thereby torn open on the one 
side, seen from the front, the right side, whereby the lobes 
of the lower lip are bent outwards. After the visit of a 
humble-bee, the flower cannot close again. The smaller 
humble-bees penetrate entirely into the flowers, whilst the 
stomach of the larger ones remains visible outside the flower.” 

In flowers from Aursundséen I found the lower lip to 
be relatively somewhat longer than is figured by WARMING; 
the length of the style was, on the whole, the same as that 
shown in Fig. 30, B. Many of the older flowers in the in- 
florescences had been opened by the visits of insects. 

Geographical Distribution: Fennoscandia, Lapland, 
Western and Northern Russia, Siberia as far eastward as 
to the river Kolyma, Mantchooria and Japan. Besides this, 
the species has an area of distribution in Central Europe 
(Germany, with a southern limit in the Bavarian Alps); 
here it is possibly å glacial relict (KERNER). 

According to NoRrRMAN, in Northern Norway the species 


grows in bogs, on boggy plains, on the coast and in willow 


7 
copses. 
Anatomy. The Root. The epidermis remains long 
intact. The cells in the outermost layer of the cortex are 
provided with a cuticularised lamella along the whole of 
their circumference; between this exodermis and epidermis 
there are no intercellular spaces, and such are wanting also 
between the exodermis and the outermost layer but one of 
the cortex; further inwards, the cortex becomes rather lacun- 
ose. Starch occurs in the cortical cells. The endodermis is 
a typical Casparian sheath. When the secondary growth in 
the stele is completed, a small central group of vessels, cir- 


cular in outline, is found to be developed. 


Scrophulariaceae. 449 


The Peduncle. Å portion of a transverse section is 
shown in Fig. 32, A. The layer of complete cells seen upper- 
most in the figure, is the endodermis. The pericycle is several- 
ayered; sieve-tissue occurs rather sparsely. The vigorous 
development of the stereom of the peduncle, is characteristic 


MAOS 





Fig. 32. Pedicularis Sceptrum carolinum. 
A, Portion of a transverse section of peduncle. B, Transverse section, 
and C longitudinal section of the upper part of a rhizome; B and 
C diagrammatic, cf. text. (Aursundséen, Norway.) (A about 220/, ; 
B and C about &/,.) 
of the species. In the mass of wood-fibres only here and 
there a radial row of vessels is inserted — in the figure the 
4åth row from the right hand is such a radial row. In between 
the groups of the primarily formed vessels, of which one is 
seen in the middle of the figure, the wood-fibres internally 
adjoins the peripheral portion of the pith, which also consists 
of strongly thickened, lignified and porose cells, so that in 
the fully developed stem the original groups of vessels are 


450 Fr. J. MATHIESEN. 


found completely surrounded by a compact mass of stereom ; 
only the tissue immediately surrounding the protohadrome 
remains unthickened and unlignified. 

The anatomical structure in that portion of the stem 
which bears the rosette-leaves deserves a more special de- 
scription. 

Å transverse section of an axis of an innovation-shoot 
in its first growth-period, about 3 mm thick, exhibited a 
distinct Casparian sheath, and in the stele a circle of small 
vascular bundles, separated by comparatively broad rays of 
intervening parenchyma; at a depth of about 3—5 layers of 
cells within the cortex there had been formed, even at that 
point of time, a cork-cambium which had produced a 3—6 
layered cork (Sudan III). 

The transverse section of a 2-year-old rhizome, with its 
richer 2nd year's leaf-rosette developed, exhibited in its 
uppermost part a structure similar to that described in the 
case of the one-year-old shoot, only, the diameter of the 
organ was larger (about 5 mm), the vascular bundles were 
more numerous and also individually broader, and they had 
on the outer side of the xylem-portions a thick cap of wood- 
fibres. In the basal portion of this shoot, which had been 
developed the year before, and which was consequently 2 
years old, there occurred, however, a distinct growth-zone 
in the vascular bundles, since, either as a direct continuation 
of the xylem-portion developed during the first year of 
growth, and furnished on the outer side with a cap of wood- 
fibres, or else separated from that xylem-portion by 1—3 
layers of thin, unlignified elements, one more xylem-portion 
was found, which also had a cap of wood-fibres along its 
peripheral portion, and evidently represented the growth of 
the 2nd year. Frequently, such xylem-groups of the 2nd year 
are also developed without being in connection with the 


Scrophulariaceae. 451 


xylem-groups of the Ist year, viz., opposite the parenchyma- 
rays which separate the original vascular bundles. The cork 
in the primary cortex is constantly few-layered only. 

Lastly, a portion of a rhizome of a 3-year-old shoot with 
a terminal peduncle, was investigated. The basal portion 
was found to be unaltered, as described above in the case 
of the 2-year-old shoot; in the part developed during the 
ånd year of growth (illustrated diagrammatically in Fig. 
32, B), fundamentally, the same development has taken place 
as that which, in the second year, took place in the basal 
portion, the only differences being that here the vascular 
groups are larger and more numerous, and the stereom-caps 
much thicker, and also that the two zones of growth are 
separated by a ring of non-lignified tissue (parenchyma) as 
much as 15 cell-layers thick, in which a layer of cork, con- 
sisting of 3—5 layers of cells, is developed; this layer of cork 
surrounds, in a tube-like manner, the xylem developed 
during the first year of growth. In the figure the cambium 
is consequently situated immediately outside the outermost 
circle of xylem-groups; the endodermis is in part distinct, 
and in the figure, is indicated by a fine line; the two thickly- 
drawn circles indicate the outer and inner cork-layer; within 
the pith there is another one like it, also indicated by a 
thickly-drawn line; in the outer circle of xylem-groups, the 
inner, dotted part indicates that this part consists especially 
of vessels, the outer radiately-shaded part, that there wood- 
fibres predominate, in the inner circle where the situation 
of the vascular- and stereom-elements is less sharply distin- 
guishable, the whole of the xylem is shaded. 

Where the rhizome bears the rosette of the 3rd year, 
it shows a transition to the structure of the peduncle, and 
gradually merges evenly into the latter — the wood-fibres 
form an almost continuous ring, and cork-development ceases. 


452 Fr. J. MATHIESEN. 


Fig. 32, C shows a diagrammatic, longitudinal section of 
that portion of the 3-year-old rhizome described above, in 
which the boundary line between 2nd and ård year's growth 
occurs — here, also, the thickly-drawn lines indicate the 
cork-layers, and the xylem is shaded. Ås is evident” from 


Å 


lg 











NDS 


Fig. 33. Pedicularis Sceptrum carolinum. 
A, Transverse section of leaf. B, Palisade cells, and C, spongy paren- 
chyma in surface view. D, Epidermal cells from the upper, and E 
from the lower surface of the leaf. (AA, B and C about 1%/,; D and 
E about ?20/,) (Aursundsåen, Norway.) 


the figure the inner layer of cork is continued horizontally 
in an inward direction, and forms a plate right across the 
pith; the outer layer of cork shows various irregularities. 
Å closer investigation of the process of development was 
not possible, owing to the very scanty material; as regards 
the development of a continuous layer of cork between the 
growth-zones of the xylem, it must be noted, that a similar 


Scrophulariaceae. 453 


structure, according to the investigations of L. Kocm! and 
E. STRASBURGER, is found in the aereal stems of Sedum 
populifolium L. 

The Leaf. The epidermis of the upper surface has 
lateral walls, varying from straight to slightly undulating; 
only along the edges of the sections are the undulations more 
marked, very much as on the lower surface (Fig. 33, D and 
E). The lateral walls are thin, and faintly and distantly 
porose; only those cells on the lower surface which bear 
glandular hairs, have, as is also the case in the other species, 
thicker and more highly porose walls. The transverse section 
shown in Fig. 33, A, exhibits the lacunose structure of the 
mesophyll: the 1—2 layers of short palisade-cells, which tend 
to be almost isodiametric, and the usually abundantly- 
branched cells of the spongy parenchyma; a surface-view of 
the latter cells is shown in Fig. 33, C. Stomata occur only 
on the lower surface; the guard-cells are on a level with, 
or project only slightly above, the epidermal cells, which 
surround them to the number of 4—6. Chlorophyll occurs 
abundantly in the whole of the mesophyll, and also in the 
epidermis of the lower surface. Non-glandular hairs are totally 
absent; glandular hairs of the usual type occur in great 
numbers under the secondary veins, and the veins of higher 


order of the leaf-sections. 


Pedicularis capitata Adams. 


Herbarium-material from St. Lawrence Bay, King 
William's Land (Gjoa Harbour), Ellesmere Land, Kotzebue 
Bay, Island of Iglorlik and from near the Taimyr River. 


1 IL. KocHx: Untersuchungen uber die Entwicklung der Crassula- 
ceen. Heidelberg, 1879, and E. SrRASBURGER: Ueber den Bau und die 
Verrichtungen der Leitungsbanen in den Pflanzen. Jena, 1891. pp. 324 
—326. 


454 Fr. J. MATHIESEN. 


Lit.: LANGE, 1880, p. 78; 1887, 262; KJELLMAN, 1882 (I), p. 257; 
1882 (IV), p. 511; RosENVINGE, 1892, p. 687; Easrwoon, 1902, p. 288; 
Simmons, 1906, p. 26; 1913, pp. 125 and 139. 


A sympodial hemicryptophyte of the rosette-type, which 
spreads by means of slender runners. The runners bear small 
scale-leaves and, rather scantily, adventitious roots, which 
are given off at the nodes; on some of the roots haustoria 
were found. 

The length of the vegetative period of the innovation- 
shoot varies from 1 to 4 years; during the first year of 
growth it develops at its apex either a winter-bud only, 
or it first forms a small rosette of 2—4 leaves. In one single 
case I found the shoot already flowering in the following 
year, but as a rule a rosette of foliage-leaves is formed in 
one, two, (which appears to be the most common), or three 
more years. The flower-bearing axis is terminal, as much 
as 9 cm high, most frequently quite leafless, rarely with 
1—2 foliage-leaves; at its apex it bears 2—6 flowers in a 
capitate raceme; some fresh leaves are always found at its 
base, since the shoot, also in the year in which it flowers, 
begins the year's growth by the development of a small rosette 
of foliage-leaves. The winter-buds are protected byscale-leaves. 

In that part of the shoots where the leaf-rosettes are 
developed, branching seldom takes place; the new shoots 
arise most frequently from the portion of the runner imme- 
diately below the rosettes, or from the part of it, where 
it turns upwards. The runners are as much as 2 mm thick; 
how long they may become, I am not prepared to state, 
owing to the scantiness of the material, which would illu- 
strate this point; several runners measured 3—4 cm. Some- 
times they may be quite short, or a runner-like portion is 
entirely lacking at the base of the innovation-shoot, so that 
several flowering shoots may occur close together. 


Scrophulariaceae. 455 


The structure of the flower is shown in Fig. 34, A—D; 
as I had only dried plants at my disposal, these drawings 
are somewhat diagrammatic. 

As in P. Sceptrum carolinum the lower lip is directed 
upwards, on its inner side (morphological upper side) are 
two convexities. The maximum distance from the base of 
the corolla-tube to the tip of the helmet, was 3 cm; the 





Fig. 34. Pedicularis capitata. 
A, A bud and B a fully expanded flower; in B the position of the anthers 
is indicated. C, Corolla-tube and upper lip cut through the median 
plane of the flower, the lower lip has been removed. D, Uppermost 
half of the upper lip, more highly magnified in order to show the vena- 
tion of the helmet; lowermost in this figure one sees the uppermost 
parts of the ridge and of the revolute portion of the edge of the upper 
lip respectively. (Ellesmere Land.) (4, B and C about Z/,; D about 2/4.) 


colour of the corolla is yellow. The position of the ridge is 
shown in Fig. 34, C, and the venation of the helmet in Fig. 
34, D; in the latter figure the uppermost part of the ridge 
is seen, and a little of the narrow revolute portion of the 
edge of the upper lip (entirely black in the figure). The 
revolute portion appears as if it were smooth, but, under 
the microscope, some of its epidermal cells are seen to be 
furnished with quite small, warty protuberances with cuti- 
cular striations; such protuberances are also found along the 


456 Fr. J. MATHIESEN. 


edges of the helmet, above the revolute portion. The corolla- 
tube is somewhat hairy within, in its lower part. 

The length of the style and the position of the stigma 
varied greatly, even in flowers from the same plant. The 
relatively longest-styled flower in my material was the one 
shown in Fig. 34, B; the plant to which it belonged came 
from Ellesmere Land; Fig. 34, A shows a not yet open flower 
from the same plant. From Ellesmere Land I have, how- 
ever, also had flowers in which the style was so short that 
it hardly reached outside the tip of the helmet, whilst in 
others it protruded 1—2 mm outside, but was bent more or 
less decidedly upwards. 

In several specimens from King Point there were flowers 
in which the end of the style was bent so strongly inwards, 
that it came to lie within the edges of the helmet, close up to 
the anthers. Some flowers from the Island of Iglorlik showed 
that, in the older ones, the point of the style may be bent 
more inwards than in the younger ones of the inflorescence; 
anything corresponding to this I have, however, not succeeded 
in confirming in flowers from the other localities. With the 
exception of the cases in which the stigma protrudes slightly, 
self-pollination by falling pollen must easily be able to take 
place, naturally, more especially where the stigma, is bent 
close under the anthers. 

According to Simmons P. capitata flowers on Ellesmere 
Land at the beginning of July, and in most years does not, 
in all probability, succeed in setting fruit there, for which 
reason it will often be under the necessity of reproducing 
itself vegetatively by means of its runners. 

Ås regards its habitat, Simmons remarks, that the spe- 
cies grows chiefly in marshy soil, but may also be found 
in drier localities where, however, the inflorescence becomes 


poorer (1—2 flowered). 


Scrophulariaceae. 457 


According to the same author, the Geographical. Di- 
stribution of P. capitata is as follows: North-western Green- 
land, Arctic American Archipelago, Arctic America, Una- 
laschka, East-Arctic Siberia to Taimyr Peninsula, and Kam- 
schatka. 


Anatomy. The Root: The structure of evidently full- 





Fig. 35. Pedicularis capitata. 
A, Portion of a transverse section of root. B, Transverse section of 
a runner, diagrammatic, cf. text. C, The peripheral layers of aåa runner 
in transverse section, showing the cork-formation. D, Two non-glan- 
dular hairs from the lower surface of the leaf. E, Transverse section 
ofleaf. (4 and C about ?20/, - B-about 1%/,; D about "9/,; E about ""9/7.) 


grown adventitious roots, about 0.5 mm thick, was as fol- 
lows: The epidermis appears to die away rather quickly; 
the cells in the outermost layer of the cortex have cuticular- 
ised outer and lateral walls; the cells under the exodermis 


were frequently divided by a thin tangential wall. The 


458 Fr. J. MATHIESEN. 


cortex is few-layered (35—7 layers), the cells in the 3 outer- 
most laåyers unite closely without any intercellular spaces; 
starch is found in the cortical cells. 

During the secondary growth of the root, the endodermal 
cells are elongated tangentially, as are the other elements 
of the cortex and, are divided ås the exodermal cells, by 
thin radial walls, into 2—4 cells; moreover, in the inner 
walls, and in the primary radial walls, there is formed a 
cuticularised lamella (the thickly drawn line in Fig. 35, 4), 
so that the stele becomes, so to speak, surrounded by a tube 
of cuticularised material, only interrupted by non-cuticula- 
rised passage-cells opposite to the protohadrome-rays; (the 
roots investigated were 2—3 rayed). The secondary growth 
of the root stops when a thin bundle of xylem-tissue, cir- 
cular in outline, and surrounded by a narrow zone of sieve- 
elements, is developed in the stele. 

The Peduncle presents nothing particular in its ana- 
tomy. Chlorophyll-grains are found in the cortical cells, the 
cells of the outer layer of the cortex have somewhat thickened 
outer and inner walls; the endodermis has recognizable Cas- 
parian dots. In the periphery of the xylem-ring the wood- 
fibres dominate; the outer layer of the pith has thickened, 
woody, porose cell-walls; the central cells of the pith are 
thin-walled and die away. On the surface multicellular non- 
glandular hairs occur. 

On the other hand, the anatomical structure of the run- 
ner-like part of the shoots deserves a fuller treatment. Å 
diagrammatic transverse section is shown in Fig. 35, B; the 
thickly drawn line slightly within the periphery indicates a 
cork-layer, a portion of it is shown in Fig. 35, C under higher 
magnification. The outermost (uppermost) layer of cells in 
the latter figure, is the epidermis; this, together with the 
2—3 outermost layers of the cortex, naturally dies away as" 


Scrøphulariaceae, 459 
i 


soon as cork-formation commences, yet how early in the 
development of the runner I am not prepared to state, but 
it probably happens rather early. The amount of cork 
developed is small; møre than five layers of corky cells (the 
thin-walled elements placed in radial rows seen in the figure) 
never occur, in fact, often only 2—5 layers; sometimes the 
cørtical layer situated immediately under the epidermis, 
forms the cork-cambium. This cork-føormation is continued 
from the runner-like, into the røsette-bearing part of the shoot. 

In the stele, which is surrounded by an endodermis of 
a similar kind as that of the root (in Fig. 35, B the inner- 
most circle indicates the endødermis), two groups of xylem are 
found; the vessels are intermingled with non-lignified paren- 
chyma, and the groups are separated by two broad medul- 
lary rays, diametrically opposite to each other, in the outer 
part of which a small group of narrow vessels occurs (leaf- 
traces). The pericycle is several-layered; like the cortex, it 
consists of cells with somewhat thickened walls. In the peri- 
fery of the xylem-groups a few wood-fibres sometimes occur. 

The Leaf. The epidermis of the upper surface has 
lateral walls varying from straight to slightly undulating, 
comparatively thick, but only slightly porøse; on the lower 
surface of the leaf the lateral walls of the epidermal cells 
are found to be slightly undulating; the cells are essentially 
more thin-walled than those of the upper surface. Stomata 
occur only on the lower surface of the leaf; they are sur- 
rounded by 4—9 cells. 

Å transverse section of the leaf is shown in Fig. 35, E; 
as far as I could judge from the material at my disposal, 
the portion which has been selected for illustration is typical: 
2 layers of palisade-cells, and a few-layered spongy paren- 
chyma with cells, which do not, on the whole, branch very 
Copiously, are seen. 


460 Fr. J.' MATHIESEN. 


On the lower side of the leaf-lobes, glandular hairs of 
the usual Pedicularis-type are found, below the branches of 
the veins of higher order; the dense covering of non-glandular 
hairs on the lower surface of the leaf as well as on the leaf- 
stalk, is very characteristic of the species; the hairs (Fig. 
35, D) are 2—5å-celled, rather thick-walled, pointed and 
smooth, they are especially numerous below the branches 
of the veins of lower order, and along the leaf-margin, but 
are also found scattered here and there among the glandular 
hairs, and on the spaces between the ;veins. The epidermal 
cells from which they originate (Fig. 35, D) may be very 
thick-walled; they are often arched cupola-like. 


Pedicularis hirsuta L. 


Alcohol-material from numerous localities both in East 
and West Greenland, collected by E. WARMING (11. 7. 1884), 
FRÆDER (MES 8 brandes ET) S HANSEN (PN TÆNSSSE 
NEEAR TE SOME ID and FAE EN DN EET PIN ONE 
and 20. 6. 1908); also from Spitzbergen (Belsund, leg. NAT- 
HORST, 1. 7. 1888) and Norway (Talvik near Alten, leg. E. 
WARMING, 17. 7. 1885). 

Herbarium-material from Greenland, Ellesmere Land, 
Grant's Land, Spitzbergen, Nova Zembla and Northern Nor- 
way (Alten). 


BE RSESBANGE ENE 23 Resen 260 tab SEER EPE ele 
seed); 1880, p. 76; 1887, p. 262; NATHoRrSsST, 1883, p. 10; WARMING, 
11866 pp VILS EVIT TD 21 4 ande L-ALE ESP PSB TES SED OASE SEE 
97 and 103; 1890, pp. 206, 208 and 213; RosENVINGE, 1892, p. 686; 
VED GIN) Sp 78 HARTE TSA DNS Fan de BODEN) ED pE ERE: 
DRE EA ENS SEE ne SS TESS (HU DET og ONDE DE BID] Bunke Sy as 
HArTtz and Kruuse, 1911, pp. 339, 342, 343, 345, 347, 348, 353, 359, 
364, 370, 375, 376, 378, 385, 402, 405, 409, 416, 418, 419, 423 and 428; 
Norman, 1895, p. 458; Ekstam, ;1897, pp. 118, 168 and 180; 1899, 
pp-—33 3740 SA ST ande 0 KRUUSE RES JEEP NS OSS SSO RISE 
304Fandi398r 190 on de EO GE DRED NOE Spar VED PD ERDGr 
202, 244, 254, 266, 269 and 272, besides notes in the preceding parts; 


Scrophulariaceae. 461 


ÅBROMEIT, 1899, p. 43; KnurtxH, 1899, p. 190; ANDErsson and HEes- 
SELMAN LIONS PSG CLE VE 190 pp TOR SOE SEG TO TA 823 
83 and 88; Dusén, 1901, p. 39; Easrwoon, 1902, p. 290; PoRrsiLD, 
1902, pp. 110, 114, 124 and 216; 1910, pp. 259, 267 and 271; 1912, 
pp. 382 and 385; 1920, p. 143; SyLven, 1905, p. 88; Simmons, 1906, 
PEP NAR tabD IE hes 8 bab SITE he SEN OT SS pp ES ES 7 ande ie 
OSTENFELD and LUNDAGER, 1910, p. 31; OSTENFELD, 1915, p. 381; 
LUNDAGER, 1917, pp. 360, 394, 399 and 402; REesvorLr, 1917, p. 214. 


Spot-bound, sympodial hemicryptophyte of semi-rosette 
type, with primary root of long duration, attaining as much 
as 2 cm in thickness and often furnished with some vigorous 
branches. Ås recorded by Tx. REsvoLL, in older specimens 
adventitious roots can be developed. 

SYLvenN describes only mere seedlings. The seed ger- 
minates in spring; during the summer a rosette consisting 
of a few small foliage-leaves, is formed, and the young plant 
enters upon its winter-rest with its shoot-apex protected by 
4—6 hbud-scales (Tx. ResvoLt): the following summer a 
rosette of more abundant leaves is formed, then again bud- 
scales and so on, until the time of flowering is reached. A 
flower-bearing shoot-portion with elongated internodes ter- 
minates the growth of the main axis. Also as TH. REsSVOLL 
remarks, during the flowering year, no rosette of radical 
leaves is formed. As regards the length of the first vegeta- 
tive stage, the said author has found, that two-year-old 
plants had not yet any floral organs developed in the ter- 
minal bud. 

At the rhizome-portion of the floral shoots, 1—3 innova- 
tion-shoots are developed — most frequently two; the suc- 
cession of shoots with short internodes which are thereby 
formed, gradually brings about the formation of a ”rhizoma 
multiceps” (a mesocorme). 

The innovation-shoots pass through a vegetative-stage 
of two years duration before flowering; every summer, a 
rosette consisting of a few foliage-leaves, is formed and, as 


XXXVII. 30 


462 Fr. J. MATHIESEN. 


in the main shoot, bud-scales, for the protection of the 
winter-buds. In the adventitious shoots, also, no rosette of 
fresh foliage-leaves occurs in the flowering year. 

I have been so fortunate as to have Arctic material at 
my disposal, collected as late as on the 1st of September 
(Danmarks Ø, leg. HArRTZz); it contained both buds which 
would flower next year, and also younger buds. The former 
were of a considerable size (1.5 cm long, 1 cm thick); ex- 


: 


BUE 32: 





Fig. 36. A—E, Pedicularis hirsuta. 

A, A bud-scale. B, Transitional form between bud-scale and foliage- 
leaf. C and D, Leaves from a part of the stem with elongated inter- 
nodes. E, Rosette-leaf. (Greenland). 

F—H, Pedicularis lanata. 

F, A lower and G an upper leaf from the rosette of the same year. 
H, AÅ leaf from the part of the stem with elongated internodes. (Green- 
land). All about nat. size. 


ternally they were protected by the somewhat broadened 
bases of the dead leaves of the rosette, then followed 10—12 
arched, pointed bud-scales, 10—13 mm long, and 5 mm broad, 
and hairy along their edges; the outermost of these bud- 
scales were withered. The young foliar organs were already 
large, and in the axils of the subtending-leaves of the in- 
florescence were floral buds about 0.5 mm in size, all enveloped 


Scrophulariaceae. 463 


in non-glandular hairs. The purely vegetative buds were in 
all cases smaller, and the bud-scales fewer in number; also 
in them the young foliar organs were well-developed. 

In Fig. 36 are shown 2 bud-scales (of which the one, B, 
was uppermost in a bud, and shows a rudimentary leaf- 
blade), a rosette-leaf and 2 leaves from the part of the shoot 
with elongated internodes (both kinds of foliage-leaves vary 
greatly in form). The leaves are hairy, especially at the 
base, and the stems also are abundantly covered with hairs, 
especially in the upper part. 

The height of the flowering plant may vary greatly. 
From Spitzbergen (leg. Tx. Wurrr) I had specimens scarcely 
2 cm high, which nevertheless bore as many as 4 flowers of 
normal size. It appears that during the end of the flowering 
period, a rather considerable elongation of the stem, especially 
of the upper part, takes place; two fruit-bearing specimens 
from Greenland (Amitasiarsak Fjord) were as much as 36 cm 
high. 

The development of a "”rThizoma multiceps” causes a 
somewhat tufted growth of the species. One specimen from 
Hvalrosodde (North-east Greenland, leg. ANDR. LUNDAGER) 
had at the same time as many as eleven flowering shoots, 
numerous leaf-rosettes and ten dead inflorescence-axes from 
previous years. This tuft-forming growth must afford ad- 
ditional protection to the winter-buds, the latter being 
covered by the numerous withered leaves. 

Haustoria are found on the roots; according to ROSEN- 
VINGE this species is parasitic on Vaccinium uliginosum and 
Salix herbacea, for example. 

In the fresh condition the root has a slightly yellowish 
colour, and a somewhat sweetish taste (NarxHorst). The 


radiating clefts, which are formed in the tissue of the root, 
30% 


464 Fr. J. MATHIESEN. 


especially in the cortex (see below), are, during the winter, 
found to be full of ice (M. PorsiLD, in litt.). 

The structure of the flower will be evident from Fig. 37 
(= WARMING, 1886, fig. 12). A shows a flower seen from 


NNEFSES SB 
DYK SM 


WIZ 
| 





Fig. 37. Pedicularis hirsuta. 
A and B (the same flower) as well as C—F and K—N are drawn from 
Greenland material; I from a flower from Northern Norway (leg. Ny- 
HUUS), O and P from flowers from Spitzbergen (leg. NAatHorst). (Cf. 
text.) (E. WARMING, 1886.) 


the side; B the same flower, and K another flower, both in 
front view: in Å the densely hairy calyx should be noted, 
the short corolla-tube, the short, warty revolute portion of 
the edge of the upper lip and the ridge which passes down- 
wards from its uppermost point, into the back of the corolla- 
tube; also the two convexities on the lower lip, and the 


Scrophulariaceae. 465 


furrow between them; in B the slightly oblique lower lip 
should be noted and the wide slit (as much as 2 mm) between 
the two revolute portions of the edge of the upper lip: here 
the stigma protrudes just beyond the tip of the helmet. C 
shows the curving of the upper part of the style, in É is 
seen its natural and usual position in relation to the helmet 
and anthers. Å flower of somewhat different structure is 
shown in /: there the style is shorter, less curved, and does 
not reach down underneath the anthers. D and F are opened 
anthers, G stigmatic papillæ and H germinating pollen-grains. 
Honey is secreted by the three-lobed swelling on the front 
of the oblique ovary (P). 

Peloria may occur: such a flower is shown in N; there 
the corolla is irregularly 6-lobed, the stamens are of equal 
length, and the style protrudes far, and is erect. 

In very small-flowered specimens brought by NATHORST 
from Spitzbergen, E. WARMING found as close contaet of 
stigma and anthers, as shown in O. 

The corolla is of a pale rose-colour, with darker throat 
and tip of helmet, more seldom the colour is pure white 
(for instance in Upernivik). The flower is set almost hori- 
zontally. The corolla-tube is . . . . 7—8 mm long; the total 
length of the flower is about 13—14 mm. Slight protogyny 
appears to occur, and the stigma probably protrudes some- 
what more in the younger flowers than later on, so that 
cross-pollination may perhaps take place in these” (E. WaR- 
MING, 1890, p. 213). According to information given by M. 
PorsiLD it is common in Northern Greenland for the corolla 
to be of a whitish shade or, at any rate, of a pale pink colour. 
EKSTAM records this species from Nova Zembla as scentless, 
but from Spitzbergen, as having a strong perfume. Like all 
the species of Pedicularis mentioned here, it must be assumed 
originally to have been adapted to pollination by humble- 


466 Fr. J. MATHIESEN. 


bees, but insect-visitors are rare in Greenland (M. PorsiLD, 
in litt.), nor are they common in Northern Norway (EKSTAM, 
1897, p.175); from Spitzbergen humble-bees are absent (AURI- 
VILLIUS). In spite of this, the species regularly sets fruit 
everywhere; it must also be said to be the one of the AÅrc- 
tic Pedicularis spp., which is best fitted for self-pollination. 

In flowers like Æ and O (less, it is true, in /) self-polli- 
nation is inevitable. 

The seeds are described and figured in LANGE, 1871 
(Cab sl 2E2D 

Geographical Distribution: West Greenland from 
64% and northwards, rare between 64” and 68”; East Green- 
land from 65" and northwards (rare in the district of Ang- 
magsalik); Arctic North America (the continent and the 
islands), Arctic Siberia and Russia, Nova Zembla and Spitz- 
bergen, the mountain summits of northern Scandinavia. 

In Greenland, according to the numerous records in 
the literature, it appears that the species makes no great 
demands as regards the nature of its habitat. Thus it can 
be found on heaths, mossy flats, herb-slopes, in coppices and 
in fissures in fields where the surface is cracked into poly- 
gonal cakes (rudemark) (LANGE, WARMING, HARTZ, KRUUSE 
and M. Porsizn), according to the last author also in gravelly 
barrens (1920, p. 143). In Greenland this species can dis- 
pense with a snow-covering during winter, and it is there 
a decided sun-plant, (M. Porsirn), and has not its proper 
home in the snow-troughs. 

According to NorManN, in Northern Norway it prefers gra- 
velly, not grass-covered localities, and there it has not hitherto 
been found where the aspect is sunny, but only on the north- 
ern and indifferent (eastern and western)sides of the mountains ; 
it belongs there to the flora of the snow-troughs (TH. REs- 
VOLL). Å. CLEVE also remarks: "Bedarf sehr wenig Insolation 


Scrophulariaceae. 467 


und meidet sogar die sonnig trockene Wiese”; the species is 
otherwise characterized as "håufig in der Heide, insbesondere 
die frischere Andromeda-H., gern in schattigen, schneereichen 
Mulden, vom Plateau das Gebirge emporsteigend, Ferner in 
der Sumpfwiese und auf den Moorhimpelen, aber einerseits 
ins Moor nicht hinabsteigend, anderseits nicht auf trockener, 
vegetationsarmer Felsenflur. Ein verfilzter Humusboden ist 
unbedingt notvendig.” In Greenland this species is a decided- 
ly spring-flowering plant, but, as an exception, flowering 
specimens may be found throughout the summer according 
to the point of time at which the snow melts at the place 
in question. HarRTtz found it on Danmarks Ø with the flowers 
still in bud on the ist of September, and expressly remarks 
that it was ”on quite recently-bared ground, at the foot of 
the snow-drift.” 

Anatomy. The Root. Fig. 38, A and B, shows portions 
of the transverse section of a root, about 4 mm thick. The 
epidermis decays very quickly — as described by Hove- 
LACQUE for the root of Pedicularis palustris — the epidermal 
cells have already collapsed in roots not more than one mm 
in thickness, and in the root figured they have disappeared 
completely. The uppermost layer in Fig. 38, 4, is the outer- 
most layer of the primary cortex, which, on the whole, only 
consists of 4—5 cell-layers. The cells have been very greatly 
elongated tangentially during the secondary growth in thick- 
ness, and are divided by a great number of thin radial walls; 
the primary cortex persists for a long time, I found it even 
on roots about one cm thick. The outer walls of the outer- 
most layer are cuticularised. At the stage of development 
figured in Fig. 38, an endodermis is not recognizable; in quite 
young roots faint Casparian dots can be demonstrated. The 
structure of the secondary cortex is very characteristic, since 
there is found in it a circle of radiating clefts, of which two 


468 ; Fr. J. MATHIESEN. 


are seen in B (26 such clefts — most of them larger — occur- 
red in the root figured) by which all the phloém-tissue of 
secondary formation is split into a corresponding number of 

radi, which in their 














SEERE EIDE A outermost parts ap- 
ED pear to be some- 


what crumpled to- 
gether, evidently by 
pressure of the tis- 
sue later on formed 
by the cambium. In 
these radii the 
groups of  sieve- 
tubes are flanked 
on both sides by 
parenchyma. With- 
in the cambium 


(l 
ae 


i 
i 


some groups of ves- 
sels are seen; cor- 
responding to each 
ray of tissue in the 


IL 


cortex. 1 .or 2 raåys 


(7 

a 

Øl 

' 

co 
rr 
Be 
[VAN 






B of groups of vessels 


occur in the xylem, 
Fig. 38. Pedicularis hirsuta. 
A, Transverse section of the outermost layers 
of the cortex of the root. B, Transverse sec- by broad radii of 
tion of a portion of the stele of the root. parenchyma, in 
110 
(Greenland.) (Å and B about 119/,,) 0 HELE REEL 


from the phloém may be continued, and may reach almost 


divided reciprocally 


to the centre of the root. 

The portionof the stem with elongated internodes shows 
a thin-walled epidermis with stomata; the cortex is few- 
layered, and its cells contain chlorophyll-grains. When the 


Scrophulariaceae. 469 


stem has reached its full development (i. e. at the end of 
the flowering period), there is found in the stele a continuous, 
but narrow ring of vessels and a few layers of wood-fibres, 
strengthened internally by the outermost layers of the pith, 







SEJR 
CS K< 





md . Ø 
Boyes sr2, 





Fig. 39. Pedicularis hirsuta. 
A, Transverse section of the leaf. B, A non-glandular hair from the 
basal portion of a leaf. C, Epidermis of the upper, and D of the lower 
surface of the leaf. (Greenland.) (A about ""%/,; B about 7%/,; C and 
Dab ou FR) 


the cell-walls of which become somewhat thickened and ligni- 
fied; the central part of the pith dies away. In the basal 
part of the stem which bears the leaf-rosettes (the mesocorme), 
a transverse section shows a circle of vascular bundles with 


470 Fr. J. MATHIESEN. 


intervening parenchyma-rays. Secondary growth in thick- 
ness takes place, and in connection with this a similar cleft- 
formation in the phloém as in that of the root. The pith is 
wide, and does not die away. 

The Leaf. The epidermal cells of both the upper and 
lower surface have thin, finely porose and almost straight 
to slightly wavy lateral walls (Fig. 39, C and D). Stomata 
occur only on the lower surface; they are surrounded by 
4—6 cells. The transverse section shows 1—2 layers of well- 
developed palisades and a spongy tissue consisting in part 
of rather copiously branching cells (Fig. 39, A). Non-glan- 
dular hairs of the type shown in Fig. 39, B occur both on 
the stem and leaf; they are found in particularly great num- 
bers on the basal portions of the foliage-leaves of the portion 
of the stem with elongated internodes, and on the leaves 
subtending the flowers, but more sparingly on the basal 
portion of the rosette-leaves proper, and only singly on the 
leaf-lobes, and then always on the upper surface. On the 
epidermis of the lower surface, under the branches of the 
veins of higher order, glandular hairs of the usual type are 
found; the epidermal cells upon which the glandular hairs 
are seated, have straighter, thicker and more highly porose 
lateral walls, than have those which surround them. 

Chlorophyll was present in abundance in the whole of 
the mesophyll and was also found in the epidermis of the 
lower surface. 

No difference could be observed as regards the anatomical 
structure of the leaves of the rosette and of those on the 
portion of the stem with elongated internodes. 


Scrophulariaceae. 471 


Pedicularis lanata (Willd.) Cham. & Schlecht. 


Alcohol-material from Greenland (Amerdlok, 15. 7. 1884, 
leg. Tx. Horm; Kingartak, 25. 7. 86, leg. RyDEer; Proven, 
24. 6. 1888, leg. MYHRE). 

Herbarium-material from Greenland, King William's 
Land, Ellesmere Land, Arctic North America, Nova Zembla 
and Arctic Siberia. 


Lit.: LANGE, 1880,, p. 76; 1887, p. 262; NATHORST, 1883, p..10; 
WARMING, 1886, pp. VII, VIII, 21, 44, 47 and 54; 1888, pp. 59, 74, 
87 and 104; 1890, pp. 206, 208, 210 and 213; RosENnvinGE, 1892, 
p. 687; 1896 (I), p. 68; Hartz, 1894, p. 45; 1895 (I) p. 306; 1895 (II), 
p. 372; Kruuse, 1898, pp. 350, 373, 380, 394 and 398; 1906, p. 249; 
1911, pp. 103, 132, 196 and 207; ABROMEIT, 1899, p. 44; EKSTAM, 
1899, pp. 7, 32, 37, 40, 48, 49 and 50; Knurtx, 1899, p. 193; ANDERS- 
son and HESsSsELMAN, 1901, p. 16; EAasrwoon, 1902, p. 291; PorsILD, 
1902, pp. 114, 178; 1910, pp. 259, 267 and 270; 1912, pp. 382 and 385; 
1920, p. 143; Simmons, 1902, p. 29, tab. II, figs. 1—3; 1906, pp. 124, 
137 and 164. 


Spot-bound, sympodial hemicryptophyte of the semi- 
rosette type; like the foregoing closely related species, it has 
a thick, scantily branched main root of long duration; ad- 
ventitious roots are probably, as a rule, only slightly deve- 
loped. As P. hirsuta, this species can form large and dense 
tufts. 

It appears that the first vegetative-stage may be con- 
fined to two years, so that the main axis of the young plant, 
after the formation of two consecutive leaf-rosettes, can, in 
the third summer, form a shoot-portion with elongated inter- 
nodes, although a comparatively weak one, terminating in 
an inflorescence. Generally, the necessary vegetative-stage 
preceding the flowering is, no doubt, of longer duration. The 
vegetative stage of the innovation-shoots extends as a rule 
over 2 years. 

P. hirsuta and P. lanata differ essentially from each 
other by the fact that the latter has no specially-developed 


ATP, Fr. J. MATHIESEN. 


bud-scales, but protects its young buds during the winter 
with the broad, arched, very persistent bases of the rosette- 
leaves developed during the growth-period of the preceding 
summer, which bases are densely hairy along their edges; 
the rosette-leaves formed towards the end of the growth- 
period, are especially distinguished by their large basal-por- 
tions. Fig. 36, G shows such a leaf. On the other hand, the 
stalk and blade of the leaf illustrated in Fig. 36, G are es- 
sentially smaller than those in Fig. 36, F, which is a pre- 
viously developed leaf from the same rosette; the leaf-blade 
may be even smaller than in G, but I do not think it is ever 
wholly wanting. 

The leaves upon the uppermost portion of the shoots 
where the internodes become elongated, have not such broad 
bases (Fig. 36, H). 

The rosettes are rich in leaves; at the base of a fairly 
vigorous flowering shoot I counted 60 withered leaves or 
remnants of such; also the above-ground portion of the stem 
is abundantly covered with leaves, especially below, although 
it cannot be said that the shoots have any true radical rosette 
of fresh leaves in the flowering year. The rosette-leaves, 
when withered, are very persistent, and may remain for 
several years, black and crumpled, at the base of the shoots 
(Fig. 36, F and G are two such withered leaves); naturally, 
this contributes towards making the tufts denser, and the 
protection of the young buds more effective. 

The root is intensely yellow in colour, and has, as already 
stated by NATHORST, a sweetish taste, almost like that of 
carrots. It is used as an article of food by the Smith-Sound 
Eskimos? and by the Chuckes (Kjellman, 1882 (III), p. 366). 

The structure of the flower is shown in Fig. 40, A—D 


1 P, FREucHEN: Om Plantekost hos Smith-Sund Eskimoerne. 
Geografisk Tidsskrift. Kjåbenhavn. Bd. 24. 1917—18. p. 310. 


Scrophulariaceae. 473 


(the drawing is executed after a sketch by EuG. WARMING 
made from material from Greenland). In A and B the 
flower is seen from the 
side, the position of the 
stamens is indicated in 
A; in B the ridge reaches 
highest above the upper- 
most edge of the revolute 
portion of the margin of 
the upper lip; of the 





three lobes of the lower 


lip, the middle one is Fig. 40. Pedicularis lanata. 

the smallest. The lower A and B, Flowers seen in side view; 
Ade 3 in ÅA the position of the anthers and 
lip is not oblique; the the style is indicated; the hairs of the 
two convexities are quite = calyx are drawn only along the out- 
lines of the figures. C, The uppermost 
part of a flower in front view. D, A 
flower in front view, the pair of anthers. (Greenland.) (A, B and 


narrow furrow between  C about >/,; D about 5/,.) (After E. 
WARMING.) 


glabrous.  C shows a 


the convexities should be 
noted. D, a pair of stamens; here the filaments of both 
stamens are hairy; this feature varies, sometimes only the 
two longest are hairy. In this species also there occur nectary- 
protuberances at the base of the ovary. 

This species is still more conspicuous (than P. flammea 
and hirsuta) on account of its richly and densely flowering 
inflorescence, and the bright pink colour of the flowers; the 
tip of the upper lip and the throat are darker in colour 
than the rest of the flower. It has also a slight scent, and 
the flowers are considerably larger than those of the two 
foregoing species (P. flammea and hirsuta), having, namely, 
a total length of 2 cm, and a corolla-tube-length of 12—13 mm 
.…….… The flower stretches out considerably, it may even be 
almost horizontal. The stigma projects out of the flower, 


474 Fr. J. MATHIESEN. 


and will evidently be easily touched by an insect visitor. 
the anthers are placed immediately behind the stigma and, 
on account of the stretched-out position of the flower, partly 
above it, so that self-pollination will probably take place” 
(E. WARMING, 1890, p. 213). The same author describes a 
flower from Spitzbergen with pelorial development. 

On Spitzbergen (forma dasyantha) Exkstam found the 
end of the style to be rolled up (as in P. hirsuta, cf. Fig. 37), 
so that the stigma is in contact with the anthers. In my 
material I always found it to be protruding, as also shown 
in the figure. 

Varieties with entirely white flowers may occur, but 
they are very rare (M. Porsirn, 1920). 

The species is fragrant (EkstaM and WARMING). It sets 
fruit everywhere abundantly and regularly, and also in Spitz- 
bergen, where humble-bees, its natural pollinators, are want- 
ing (ÅURIVILLIUS); according to information given by M. 
PORSILD, insect-visitors are rare in Greenland. Among the 
Eskimos the plant is known for the abundance of honey 
contained in its flowers; at Cape York its popular name is: 
«The food of the humble-bees;” and children pick the flowers 
and suck the honey out of them or even eat them entire 
(FREUCHEN, |. c.). 

During the end of the flowering-period, the axis of the 
inflorescence elongates. 

Geographical Distribution. The species is common 
in West Greenland, from Disco Bay and northwards. "In 
N. Stromfjord it is restricted to alpine stations and northern 
slopes and rather scarce (P & E). South of Holsteinsborg 
observed several times down to Itivdlinguaq, 66730" (P & EY” 
(Porsirn, 1920); from East Greenland it is totally absent 
(PorsiLn, 1. c.). It is also found in Arctic North America 
(the continent and the islands), the western Subarctic North 


Scrophulariaceae. 475 


America, (the Rocky mountains), Arctic Asia, Nova Zembla 
and Spitzbergen. The species prefers fairly dry habitats. 
M. PorsiLn records that it is found ”on rocky flats and on 
heaths” (1912, p. 382). "In poor and open heath, often in 
gravelly barrens far away from other plants. When growing 


NE Kl 


SER ng 










IO 
BET, 
SY 





Fig. 41. Pedicularis lanata. ' 
A, Transverse section of the leaf. B, Epidermis of the lower, and 
C of the upper surface of the leaf. D and E, Non-glandular hairs from 
the lower surface of the leaf. F, A glandular hair from the stem. G, A 
non-glandular hair from the basal portion of leaf. (Greenland.) (A 
about FED MCÆDSE ander Fab our SEG about 2/GE) 


amongst other plants, however, the roots also of this species 
are provided with haustoria” (1920, p. 143). During the 
winter, although the plants in themselves are often bare of 
snow (1. c.), yet the dense tufts always catch some snow 
between the numerous decaying leaves, and this naturally 


476 Fr. J. MATHIESEN. 


affords some protection to the winter-buds (M. PoRrSsILD in 
litt... According to the same author, m the same locality, 
the species flowers 1—2 weeks before P. hirsuta (1920, 
p. 143). On Spitzbergen, according to EKSTAM, it is found 
"auf trockenen, starker Insolation ausgesetzten Abhången,” 

In anatomical respects, the root and stem agree closely 
with the corresponding organs in P. hirsuta. The anatomy 
of the leaf is shown in Fig. 41, AG. In the transverse 
section 1—2 layers of short and broad palisade-cells are seen, 
and a spongy parenchyma, which consists of distinctly 
branched cells. B shows a portion of the epidermis of the 
lower surface, and C a similar portion from the upper sur- 
face of the leaf. The lateral walls are thin and finely porose; 
stomata are found only on the lower surface of the leaf and 
the guard-cells occur on a level with the leaf-surface. On 
the basal portion of the leaf, the cuticle is thicker than on 
the stalk and the lobes, and it is furnished with fine stria- 
tions. Chlorophyll occur in abundance in the mesophyll, and 
also in the epidermis of the lower surface. 

The glandular hairs in this species are almost exclu- 
sively confined to the edges of the leaf-sections; in Å is seen 
exteriorly to the left, a group of the tracheidal tissue in 
which the branches of the veins end, and in connection with 
which the glandular hairs occur. The structure of these is 
similar to that in the other species; the epidermal cell from 
which the glandular hair is developed, has not, however, in 
P. lanata, much thicker or more highly porose walls, than 
have the epidermal cells, which surround it. Non-glandular 
hairs of the type shown in G occur in abundance on the 
edges of the leaf-bases, on the stem and, in the floral region, 
on the scale-leaves and calyx, more scantily on the leaf- 
stalk, and on the upper surface of the leaf-blade. On the 
lower side of the leaf-lobes there are small non-glandular 


Scrophulariaceae. 477 


hairs (Fig. 41, B, D and E) intermingled with the glandular 
hairs; on the petiole and stem are glandular hairs of the 
form shown in Fig. 41, F. 


Pedicularis flammea L. 


Alcohol-material from Greenland (Godthaab, leg. E. 
WARMING, 29. 6. 1884; Christianshaab, leg. S. HANSEN, 2. 7. 
1888; Prøven, leg. MYHRE, 2. 7. 1888; Danmarks Ø, leg. N. 
HaArTz, 18. 2. 1892), Iceland (Hvitarvattn, leg. ÅA. FEDDER- 
SEN, 3. 6. 1886), and Northern Norway (leg. NYHuus, 1885). 

Herbarium-material from numerous places in East and 
West Greenland, from Norway, Iceland and Arctic America. 

Lit.: LANGE, 1870, pp.254 et. seq., 266, tab. III," fig: 22. (the 
seed); 1880, p. 75; 1887, p. 262; WARMING, 1886, pp. VII, VIII, 44, 
Alands sær s sn p eN SÆR BORE OR ETERN NORS O ande 
1890, pp. 207, 208 and 211; RosEenNvInGE, 1892, p. 686, 1896 (II), 
pel 8 HARTE EL SAD pr LES 2 and AS RET 895 (LE p pE TOD ORE 
175, 188, 247, 256, 288 and 290; 1895 (II), pp. 335, 359 and 372; HarRTzZ 
andskkrR use lo ED DN SS 88542 34735735 93781319 3808385) 
409, 411, 423 and 428; BøRGESEN, 1895, pp. 223 and 225; NORMAN, 
1895, p. 458; Kruuse, 1898, pp. 350, 373, 379,.394 and 398; 1905, 
p. 176; 1906, p. 248; 1911, in part IV pp. 196, 243, 254, 255 and 274 
besides notes in the preceding parts; ABROMEIT, 1899, p. 43; KNUTH, 
SIE DNK GEEVEN 1 OOS pp TET SINDET and DUSEN 1201 
pr 39: Porsmep ed 02 pp ANSETE NELS RE DAGE MOTORER OLES 
pp. 382 and 387; 1920, p. 142; OSTENFELD and LuUNDAGER, 1910, 
p. 31; Simmons, 1913, pp. 124 and 140. 

Spot-bound, sympodial semi-rosette hemicryptophyte, 
with short, vertical mesocorme which dies away behind, and 
a rather abundant development of adventitious roots. 

The first vegetative-stage fairly commonly appears to 
extend over 4 years — in the case of one individual (Disco) 
I found it to extend over 3 years, and in the case of another 
(Iceland) 6 years. AÅ short, vertical mesocorme is developed 
during the first vegetative stage; the main root is still present 
during the first flowering-period, but dies away afterwards. 

XXXVII. 31 


478 Fr. J. MATHIESEN. 


The winter-buds are provided with a number of bud-scales; 
these as well as the foliage-leaves leave real leaf-scars on 
the mesocorme on decaying. Adventitious roots are devel- 
oped rather abundantly, especially from the basal part of 
the mesocorme, they generally occur in connection with 
innovation buds; the adventitious roots are somewhat 
swollen. 

The innovation-buds arise from the axils of the scale- 
leaves; the lower buds on the mesocorme do not generally 
develop any further, but remain as reserve-buds; as in P. 
Oederi it is the buds which occur in the axils of the inner- 
most scale-leaves formed during the summer previous to 
that in which the parent-shoot flowered, which are the most 
vigorously developed. Usually only two of these buds ex- 
pand, but the number may be more, in a single case I counted 
as many as 6. 

During their first summer the adventitious-shoots always 
bear two fully-expanded foliage-leaves; as in the main- 
shoot the terminal bud is protected by scale-leaves. In case 
flowering does not take place in the following year, — it 
may occur, but no doubt only exceptionally, — a rosette 
of foliage-leaves is developed, followed in turn by scale- 
leaves, and then, after passing through still another winter 
the shoot, in the majority of cases, succeeds in flowering. 

In a specimen from Ignerit (Greenland) in which 2 of 
the basal buds on the mesocorme had developed further, 
one of them had formed, rosettes of foliage-leaves for 4 suc- 
cessive summers, and the other for even 5. 

In a winter-bud (the specimen in question was collected 
by N. HarTtz, on the 18th of February 1892, on Danmarks 
Ø, East Greenland) there were found young foliar organs 
and flowers, all of which were highly developed; in the 
young flowers the separate parts were quite distinguishable 


Scrophulariaceae. 479 


under the microscope. The shoot which was terminated by 
this winter-bud, was one year old — consequently, the length 
of the purely vegetative period in this case will only have 
been one year — during the previous year it had, as is usually 
the case, unfolded 2 foliage-leaves; the bud-scales numbered 3. 

The maximum height of the flower-bearing shoot was 
found to be 28 cm; from Iceland and Greenland I found 
small individuals, flowering for the first time, which were 
only 3 cm high. ÅA specimen from Disco differed entirely by 
a total absence of foliage-leaves on the above-ground stem, 
which was 20 cm high, and by having flowers, the lowermost 
of which were long-stalked almost down to the base; the 
axis was comparatively very thick (about 8 mm in diameter). 

E. WARMING found the flowers from Norway (W. Fin- 
mark), Iceland and Greenland to agree entirely in structure; 
from his descriptions I quote the following: "The flowers 
are not very conspicuous; their lower part stands erect, 
almost parallel with the axis of the inflorescence, their upper 
part is bent slightly forward (Fig. 42, 4); the colour is yel- 
low, the upper part of the helmet is, however, more or less 
reddish-brown. The corolla-tube is only 7—8 mm long; the 
lower lip is very small, with three rounded lobes, and often 
appears smaller still by the fact of its edges being bent 
backwards (Fig. 42, C). The cleft of the upper lip is some- 
times very narrow (!/, mm, see Fig. 42, D), but sometimes 
wider, even so wide that there is a large entrance to the 
interior of the flower, as for example in flowers with an 
upper lip curved strongly forward. 

This species appears to be as highly adapted to self- 
pollination, as it is poorly adapted to insect-pollination, 
which also agrees with the fact that it is so little conspicuous. 
I never saw the stigma protrude from the blunt, untoothed 
tip; so it is placed immediately in front of, or slightly under 

31+ 


480 Fr. J. MATHIESEN. 


the anthers (Fig. 42, F). The fact that in many flowers I 
have found the stigma covered with pollen-grains, some of 
which were germinating, and have found abundance of pol- 
len around the anthers, even in a flower like A—B, in which 
the stigma is situated so unusually high above the anthers, 
is a justification for regarding this as brought about by self- 
pollination. I must, however, point out the fact that the 





Fig. 42. Pedicularis flammea. (Greenland). 
A, A flower in natural position in the inflorescence; B (5/1), the upper 
part of the same in longitudinal section. C, Lower lip and the part 
around the throat seen in front view. D, The upper part of a flower 
in front view. E, Lower lip, slightly magnified. F as B, but with 
the stigma in another position. G, A flower in natural position, larger 
than A, showing the venation of the upper lip. (E. WAarminG, 1890.) 


stigma is not placed in the same position relative to the 
anthers in all the flowers of the same inflorescence, and it 
appeared to me to be the rule for it to be situated slightly 
more forward in younger than in older flowers; . . .. the 
filaments are glabrous. In many places in West Greenland, 
as far as Upernivik (RYDER), it was found with abundant 
fruit. Even in July (1884) new fruit was set at 64” and 
GYS IN Tabte 


Scrophulariaceae. 481 


I can add to the above that in older flowers the helmet 
often appears to bend forward, as described by KERNER in 
the case of P. Oederi. 

In the material at my disposal, both from West and 
East Greenland, there were many fruiting specimens. That 
it really is the rule for the stigma to be enclosed in the helmet, 
is seen among other things, by the fact that Brytrt in his 
Haandbog i Norges Flora” gives this as a specific character. 

Geographical Distribution: West and East Green- 
land (rare in South-west Greenland according to RosEN- 
VINGE, in East Greenland even as far north as Germania 
Land according to OSTENFELD and LUNDAGER), Iceland, Arc- 
tic and Subarctic North America, Arctic Russia and the 
mountainous regions of Northern Scandinavia. 

Ås regards the habitat of P. flammea, many notes are 
to be found scattered in the literature on the subject. A. 
CLEVE states that in the district of Northern Sweden in- 
vestigated by her, it was found "nur auf den Moorhugeln 
und in Sumpwiesen zusammen mit der vorigen (P. hirsuta), 
welche diese Art nicht bis in die trockene Heide zu begleiten 
vermag, ebensowenig wird sie auf dem N. Abhang oder in 
der Nåhe von spåt schmelzendem Schnee gefunden.” In 
Northern Norway it is found, according to NorMan, only 
on the northern and on the indifferent (eastern and western) 
sides of the mountains. In West Greenland it grows, accord- 
ing to E. WARMING, in willow-coppices where it may be said 
to have its home, and also on herb-flats, on heaths (parasiti- 
cally on Vaccinum uliginosum and Salix herbacea) and in 
grassy bogs. Porsirn has also found it on slightly damp 
moss-heaths. N. Hartz and KRUUSE record it from East 
Greenland as growing on herb-slopes and rocky flats, in 
carpets of prostrate Betula nana and Vaccinium uliginosum, 
on knolls in bogs, and on damp heaths. According to HAarRTZ 


482 Fr. J. MATHIESEN. 


(1895 (II), p. 335), its habitat is snow-covered during winter; 
the plant is mentioned as common in the interior of the 
fjords, but absent further eastwards, out towards the open 
ea. DuSsEN records practically the same, and he also knows 
it only from a single habitat in the coast-land outside the 
fjords (Cape Borlase Warren). A. CLEVE considers it to belong 
to the "Gruppe der spåte- 
ren Fruhlingspflanzen.” In 
Greenland it is a middle- 
summer-flowering plant (M. 
PoRrsILD). 

The Anatomy of the 
root and stem agrees ex- 
actly with that of the cor- 
responding organs in P. 
Oederi. The parenchyma of 
the root contained a great 
abundance of amylodextrin- 
starch. . 





The  Ssurface-view of 
Fig. 43. Pedicularis flammea. 
A, Epidermis of the upper surface 
of the leaf; B, of the lower surface of the leaf is shown in Fig. 
of the leaf. C, D and E, Glandular i 
hairs; C and D from the upper ER 
surface and Æ from the lower sur- the lateral walls were al- 
face of the leaf. (Greenland.) (Å ways found to be rather 
and [B tabout 29%/,; C, D and E 
about 79/1.) 


upper and lower epidermis 


faint. The transverse sec- 
tion of the leaf agrees ex- 
actly with that of P. Oederi. Non-glandular hairs are ab- 
sent; in addition to the glandular hairs of the usual Pedicu- 
laris-type, such forms are also found as are shown in Fig. 
43, C, D and E; C and D occur scattered, especially on the 
upper surface of the leaf, and Æ on the lower surface of the 
segments, especially along their main vein and lateral veins 


Scrophulariaceae. 483 


of Ist order. Chlorophyll-grains occur abundantly in the 
whole of the mesophyll, and also in the epidermis of the 
lower surface. 


Pedicularis Oederi Vahl. 


Alcohol-material from Norway (Kongsvold (Dovre), leg. 
E. WARMING, 13. 7. 1887; Muggrubfjældet near Røraas, leg. 
THEKLA RESVOLL, 29. 7. 1918). 

Herbarium-material from Nova Zembla, Arctic Siberia 
(Khabarowa and from near the Taimyr river) and St. Law- 
rence Island. 


Lit': AXxXErLL, 18697 p2 102; KJELLMAN; 1882 (1), p/-257; 1882 (II), 
p. 325; 1882 (IV), p. 510; KJELLMAN and LuNDSTROM, 1882, p. 304; 
ÅURIVILLIUS, 1883, p. 451; WARMING, 1886, p. 47; 1890, pp. 207, 208, 
210 and 214; LinNnman, 1887, pp. 82 and 99, tab. IV, fig. 46; KERNER, 
Bd. II, 1898, p. 337; Knurtx, 1899, p. 186; SyLven, 1906, p. 89; ScHRO- 
TER, 1908, pp. 454 455 and 458; REsvoLL, 1917, p. 210. 

Spot-bound, sympodial semi-rosette hemicryptophyte 
with a short, vertical mesocorme which dies away behind. 
Adventitious roots are rather abundantly developed. 

According to SYLVEN the seed germinates in spring. 
The elongated hypocotyl unites with the main root, scantily 
branched throughout, into a peg-shaped food-storing organ 
which becomes rather thick at an early period. In more 
advanced young-plants adventitious roots, thickened by the 
storage of reserve food-material, like the main root, some- 
times appear to be developed from the hypocotyl, or from 
the base of the epicotyl.” Tx. Resvorr has studied the 
whole development from seed to flowering plant: During the 
first summer two small foliage-leaves expanded, the terminal 
bud was protected by a few scale-leaves, and contained the 
rudiments of the foliage-leaves of the following summer; the 
plant passed the winter in this stage. The foliage-leaves fell 
off late in autumn, and left leaf-scars. "After having passed 


484 Fr. J. MATHIESEN. 


through the winter, the young leaf-organs developed into a 
rosette of foliage-leaves, few in number, . .. even early in 
the summer the development of foliage-leaves stopped, in- 
stead of which the apex of the stem developed a terminal 
bud covered with scales. In this manner the plant continues 
through several years. The leaves, which are exceedingly 
small in the first summer, having blades only about 1cm 
long, become larger every summer, and the number of the 
leaves also increases with the age of the leaf-rosette. The 
last summer before the plant flowers I have seen 4—8 foliage- 
leaves.” 

The shortest first-vegetative-stage observed by the 
author in question, extended over 3 summers, in other 
individuals it proved to have extended over as many as 6 
summers; in one individual (from Nova Zembla) I found 3 
vegetative years, another showed 7. As the leaves fall off, 
leaving a regular leaf-scar (as already described by E. WAR- 
MING (1890, pp. 207 and 209) both for this species and for 
Pedicularis palustris) and the broader and more distantly- 
placed scars of the foliage-leaves can be easily distinguished 
from the narrower and more closely-placed scars of the 
scale-leaves, the individual year-growths of the mesocorme 
become very marked and easily observable (TH. ResvoLL). 

It is characteristic of this plant, that the development 
takes place, so to speak, suddenly, as long as the plant is in 
the rosette-stage. Thus all the leaves come out almost at once, 
and not one by one during the summer; and as soon as the 
rosette has unfolded, the bud for the next summer is already 
seen in the middle of it. The same is also the case as regards 
the rosette-stage of the lateral shoots.” (Tx. REsvoLL). 

Even in the bud, the rudiments of the organs which 
are to expand during the next summer, attain a high degree 
of development. In an individual which Tx. REsSVOLL in- 


Scrophulariaceae. 485 


vestigated on Knutshø (Dovre) on July 23rd, the terminal 
bud proved to contain "the young stem for the coming 
summer, covered by the rudiments of the stem-leaves; within 
these were the subtending leaves of the flowers; the separate 
parts of these were already distinctly discernable.” 

The flower-bearing stem-portion has, as a rule, no foliage- 
leaves at its base, but such may occur as an exception. 

In the interval between germination and flowering the 
plant develops a vertical mesocorme, as much as 1 cm long 
and 0.5 cm thick, from the lower part of which there proceed 
a varying number of adventitious roots (I counted as many 
as 8); as in P. flammea they are somewhat swollen, as a 
rule especially in their proximal half, but sometimes also 
along their entire length, but narrowing abruptly towards 
the base; they become as much as 10 cm long, and as much 
as 6 mm in diameter; most frequently they are furnished 
with slender, filiform branches, or, at any rate, scars 
left by such are present; more rarely with 1—2 vigorous 
root-branches. The main root dies away; in the majority 
of cases it probably does not survive the first flowering 
period of the individual. As mentioned by WARMING (1890, 
p. 207), the majority of the adventitious roots arise at the 
base of a bud on the mesocorme. According to the obser- 
vations of WARMING and TH. REsvoLL — which observations 
I am able to confirm fully — the lateral buds are developed 
especially in the axils of the scale-leaves, and are of the 
same age as these. The buds in the lower part of the stem 
develop two small foliage-leaves during their first summer, 
but usually do not develop any further; as a rule they are 
found as small formations covered with a few scale-leaves; 
they must be regarded as reserve-buds, for, as seen in a 
specimen from Nova Zembla, under certain circumstances 
they may develop further and even produce flowers. 


486 Fr. J. MATHIESEN. 


The most fully developed buds are always found (Tx. 
RESVOLL) in the axils of the uppermost set of scale-leaves, 
which are found to the number of 35—10 at the base of the 
flower-bearing stem-portion. It is commonly the innermost 
scale-leaves which subtend the innovation buds, and usually 
only two such are found, more rarely several (as many as 7, 
TH. ResvorLt). During their first summer they develop 2—3 
rather large foliage-leaves, often followed by 1—2 quite small 
ones, and lastly bud-scales for the protection of the winter- 
bud. The buds open almost at the same time as the flowers 
of the main shoot. 

According to TH. REsvoLL, the duration of the vegeta- 
tive-stage of the lateral shoots may be restricted to one year 
only; it is, however, longer, as a rule; for instance, on 
Knutshé (Dovre) the said author found 4-year-old lateral 
shoots, which had not as yet young floral organs in the 
terminal bud. In several cases also I found a vegetative 
stage of only one year's duration (among others in a specimen 
from Nova Zembla), more frequently, however, 2 or 3 years. 

As the parent-rhizome dies away behind, and the lateral 
shoots develop adventitious roots, there is a possibility of 
vegetative propagation, although, of course, the plant cannot 
spread much by this means. 

SCHROTER records this species from the Alps as occur- 
ring parasitically especially on Carex firma. 

The structure and biology of the flower was first des- 
cribed by LINDMAN; since that E. WARMING and KERNER 
have dealt with this species. Some unpublished drawings of 
the flower have been kindly placed at my disposal by E. 
WARMING; the illustrations in Fig. 44, with the exception 
of G and H, have been made after them. Å is characterized 
by its long corolla-tube, its densely-hairy calyx, (the hairs 
have been indicated only along the edge of the calyx), 


Scrophulariaceae. 487 


and by its but slightly protruding stigma; in B the main 
features of the venation of the helmet are seen, to which a 
system of raised ridges on the outer side of the helmet cor- 
responds. The dots on the side of the helmet in Fig. 44, B 
indicate small glandular hairs, which are especially numer- 
ous in the front part of the helmet. Fig. 44, H shows such 
glandular hair in lateral view, highly magnified; the head 
is 4-celled. D is the lower lip in surface view; the two con- 





Fig. 44. Pedicularis Oederi. 
A, B, C, D and E are drawn from flowers from Sweden; F is a flower 
from Kongsvold (Dovre). G, Transverse section of one of the warts 
from the revolute portion of the edge of the upper lip. H, A glandular 
hair from the helmet. (A, B, C, D, E and F slightly under ”/,; G and 
H about %9/,.) (Drawn by E. WARMING.) 


vexities are distinctly visible here, as also in Fig. 44, C and 
F. Fig. 44, F shows a flower with a very far-protruding 
style; in Æ the position of the stamens is seen, the filaments 
of the foremost pair of stamens are hairy along their upper 
part. One of the warts from the revolute portion of the 
upper lip is seen under high magnifying power in Fig. 44, G; 
on its outer surface it has strong cuticular striations, as 
have also the cells which surround it. The calyx, subtending 
leaf and axis of the inflorescence vary from glabrous to 


488 Fr. J. MATHIESEN. 


hairy, even in individuals from the same locality (Muggrub- 
fjældet near Råraas); the hairs are multicellular, pointed 
and glabrous. 

I quote the following from Linnman's exhaustive des- 
cription:— "The flower is 20 mm long, yellow or whitish- 
yellow in colour; the tip of the upper lip is dark-red on its 
inner side, and this colour more or less penetrates through 
and gives the corresponding spot on the outer side a greyish- 
red tint; the upper lip has often in addition on the outer 
side a dark-red spot on each side. The flower is quite erect 
and adpressed to the peduncdle..... The stamens agree with 
those in the other species; their pollen is consequently dry 
and. loose, is fully developed even in the flower-bud, and is 
contained in a receptacle formed by the four convergently 
dehiscing anthers, which are held together by the laterally 
highlyfcompressedt upper lip eee As in the other species 
the pollen is shed when an insect penetrates into the flower, 
and thereby widens the slit of the upper lip, which is rather 
narrow in this species..... The nectary is in the same place 
as in the other species; the corolla-tube is 10—13 mm high.” 
The position of the stigma in the flowers figured by Linn- 
MAN is very much like that in Fig. 44, B and F, given here; 
in addition LINDMAN mentions flowers in which the style 
was so short, that the stigma remained quite inside the 
helmet, and did not even reach the anthers; the stigma was 
normally developed, however, in these flowers. 

According to KERNER (p. 337) the whole of the upper 
lip curves forward so strongly at the end of the flowering 
period, that it looks as if it were broken; even if it originally 
stood as a continuation of the lower part of the corolla-tube 
enclosed in the calyx, it stands then at an angle of 70”, or 
even 90? with this, and by bending forward it drags down 
with it the style and the stamens, so that the stigma no 


Scrophulariaceae. 489 


longer stands in front of, but below the anthers; these are 
no longer united, and shed their pollen spontaneously, which 
is scattered down upon the stigma, and effects self-pollination 
in case cross-pollination should fail. Liunnman and WARMING 
mention nothing regarding this point; I observed it in many 
inflorescences but must, however, add that the curvature 
may also take place in the uppermost part of the corolla-tube. 

KERNER (Die Schutzmittel der Bluthen gegen unberu- 
fene Gåste, 1876) and after him H. Mirrer, describes the 
pollination of P. recutita, a species very similar to the pre- 
sent one, and, in so doing, states that the insect (humble- 
bee) on its visit thrusts its proboscis into the narrow furrow 
between the two convexities on the lower lip, and forces 
these and the edges of the slit of the upper lip apart from 
each other; while LINDMAN maintains that the furrow in 
P. recutita is too narrow an opening for the introduction of 
the proboscis of the humble-bee, as this is not even pos- 
sible in the larger-flowered P. Oederi, where the proboscis 
can only be thrust in higher up, at about the middle of the 
revolute portions of the edges of the upper lip ("die Rolle”). 
Visitors: Linnman observed Bombus nivalis and Bombus 
alpinus; AURIVILLIUS mentions frequent visits of humble- 
bees. 

According to WARMING, fruit is set abundantly in 
Scandinavia. 

Geographical Distribution. In the Carpathians and 
the Alps (fganze nårdliche Kette von St. Gallen bis Waadt — 
fehlt den Centralalpen,” ScHROTER), the mountainous districts 
of Scandinavia, Arctic Russia and Siberia (recorded by 
KJELLMAN from several localities along the north coast of 
Siberia), Nova Zembla, Central Asia and the mountainous 
districts of East Asia. 

Habitat: Bryrr (Håndbog i Norges Flora) records 


490 Fr. J. MATHIESEN. 


boggy places in mountainous districts, . ... from the birch- 
zone up to the lichen-zone; more rare below the pine- 
limit”; Ta. REesvorL has now and then observed it in 
snow-troughs; it is, however, no typical snow-trough-plant. 
It does not ascend very high in the Alps; ScHROTER (p. 255) 
mentions it as occurring in the transition-zone between his 
Agrostidetum-trifolietum and 
Curvuletum — (Carex-curvula- 
assoc.). In Arctic Eastern Asia 
KJELLMAN found it growing 
on damp coastal plains. 
Anatomy. In anatomical 
respects the root very much 
resembles that of P. hirsuta, 
the radiating cleft-formation 
in the stele — at any rate in 





the adventitious roots, which 


Fig. 45. Pedicularis Oederi. 
A, Transverse section of the are the only ones that have 


peripheral layers of the root. heen investigated — is not, 


B, Section through a leaf-scar. = É 
(A about 79/,; E about 140/,,) however, so decided as in P. 


hirsuta. 

The epidermis decays early; during growth the cells of 
the cortex are greatly elongated in a tangential direction, 
and undergo divisions by thin radial walls. The portion 
figured in Fig. 45, 4 is taken from a transverse section of 
an adventitious root, 6 mm thick; the endodermal region lies 
at a depth of only 3—5 cell-layers; the Casparian dots are 
still recognizable here and there in the endodermis, the cells 
of which, as those of the primary cortex, are seen to be 
very greatly elongated in a tangential direction, and to have 
undergone secondary division. In the secondarily formed 
cortex the parenchyma predominates over the sieve-tissue. 

In all the parenchymatous cells of the root and meso- 


. Scrophulariaceae. 491 


corme (except the outermost layer) reserve food-material 
was found in the form of grains, as much as 10% in size, 
round, or slightly angular, and with a small central cavity. 
With a solution of iodine-iodide of potassum they assumed 
a red-violet to brownish-red colour, which disappeared on 


Å ! Mm il 
ret 







Vs: 


SDNGE:D REE AE 
ES es ESSLASE 


some ET BRIGADER 
() 
Mg Sd, HU ZL 








Fig. 46. Pedicularis Oederi. 
A, Transverse section of the leaf. B and C, Epidermal cells, B, from 
the upper, and C from the lower surface of the leaf. D, A non-glandular 
har H(AFaboutt 0, Band] about 220" FED about") 


heating, but reappeared after cooling; they stand out after 
treatment with potash-lye, and at last disintegrate altogether, 
Judging from the above-mentioned reactions, they must be 
assumed to consist of amylodextrin-starch. 

The structure of the above-ground stem is fundamentally 
as in the other species. The thickening and lignification of 
the wood-fibres between and outside the vessels, and of the 


492 Fr. J. MATHIESEN. 


peripheral cells of the pith, does not appear to take place 
until towards the time of ripening of the fruit. 

The Leaf: Some of the cells of the upper and lower 
epidermis are shown in surface view in Fig. 46, B and C. 
Here, the great thickenings on the lateral walls of some of 
the epidermal cells of the upper surface and, as regards the 
lower surface, especially the walls of the cells which bear 
the glandular hairs, is very striking. This feature, however, 
is not constant in the species; as regards this point the 
individuals from Dovre were, on the whole, as shown in the 
figures in question, but the thickenings were far fainter, or 
even entirely wanting, in the specimens from Rårås. Ås 
usual in the Pedicularis spp. the stomata occur only on the 
lower surface of the leaf, and the epidermal cells which 
surround them have highly undulating lateral walls. 

Å transverse section of the leaf is shown in Fig. 46, A. 
The spongy parenchyma consists of copiously-branched cells, 
especially in its lower layer. As seen in the figure, the epi- 
dermis of the lower surface, in the spaces between the branches 
of the veins, is highly convex and in places without direct 
connection with the spongy parenchyma, so that, on the 
lower surface of the leaf, there are hollow spaces — air- 
chambers — in the roof of which the stomata occur. In 
the figure, a glandular hair is seen under each of the veins; 
two such hairs are shown in surface view in Fig. 46, C. Dis 
a non-glandular hair. Chlorophyll-grains occur abundantly 
in the whole of the mesophyll, and in the epidermis of the 
lower surface. 

AÅ section through one of the scars which the leaves at 
their fall leave on the rhizome, showed that here a cork of at 
least six layers was formed; outside the leaf-scars no cork- 
formation takes place. Å small portion of such a section is 
shown in Fig. 45, B. 


Scrophulariaceae. 493 


Summary and General Remarks. 


A. Account of the Growth-form and remarks on the 
StructureoftheShoot, the Vegetative Reproduction 
and the Winter-stage. 

I. Undershrub-Chamæphytes: Veronica fruticans and 

alpina. 

The shoot-development extends over 2 years; especially 
in V. alpina the first-year portions of the shoots often assume 
the character of runners; this, in conjunction with the 
abundant development of adventitious roots, and the dying 
away of the older portions of the stem, makes abundant 
vegetative reproduction possible in this species. Special bud- 
scales do not occur. 

Both species may form lax tufts. Nanophyllous winter- 
greens. In the Arctic regions they require a snow-covering 
during their winter-rest; V. fruticans requires also a habitat 
which is freed from snow rather early in spring. 

II. Hemicryptophytes. 

a. Proto-hemicryptophytes: Castilleia pallida and 
Bartschia alpina. 

The former species has a main root of long duration; 
in Bartschia it dies away early, and adventitious roots are 
abundantly developed; this in conjunction with the forma- 
tion of runners allows rather abundant vegetative reproduc- 
tion. When Bartschia grows in comparatively dry and firm 
soil the formation of runners is checked and it forms tufts; 
in that case it evidently forms a transition to the under- 
shrub-chamæphyte type. Nano-microphyllous. During the 
winter-rest the growing-point is protected by the scale-leaves 
on the portion of the shoot formed the foregoing summer. 
In the Arctic regions Bartschia (and probably also Castilleta) 
requires a snow-covering during its winter-rest. 


REX VER: Jen 


494 Fr. J. MATHIESEN. 


b. Rosette- or semi-rosette plants: Pedicularis spp. 

1. With Runners: P. lapponica and P. capitata. 

In P. lapponica the runners are most frequently mono- 
podial, and during a limited number of growth-periods they 
form few-leaved rosettes at their apex, while the floral 
shoots, without any preceding vegetative stage, arise later- 
ally on the rosette-axis. Sometimes the runners may ter- 
minate their growth by the formation of a floral portion. 
Floral shoots may also arise laterally on the basal portion 
of older shoots of the same kind, so that a small rhizoma 
multiceps is developed. 

In P. capitata the runner terminates its growth after a 
vegetative period of 1—3 years, by the formation of a floral 
portion, often without foliage-leaves (a peduncle). 

The winter-buds of both species are protected by scale- 
leaves. Adventitious roots are developed and vegetative 
reproduction takes place. 

In the Arctic regions P. lapponica (and certainly also 
P. capitata) requires a snow-covering during its winter-rest. 

2. Without Runners: P. sudetica, euphrasioides, Scep- 
trum carolinum, hirsuta, lanata, flammea and Oederi. 

All these species have a short vertical rhizome ("meso- 
corme”, E. WARMING]); with reference to the duration of 
the main root, they may be divided into two groups, as 
stated by E. WarmInG (1890, p. 206 et seq.), viz., those in 
which it is of long duration, a fact connected with its vigour, 
and often rather copious branching (P. euphrasioides, hir- 
suta and lanata), and those in which it is of short duration 
(P. flaemmea and Oederi, to which may be added P. sudetica 


and Sceptrum carolinum). 


1 E. WARMING, 1918: Om Jordudlåbere (Underground Runners). 
With a résumé in English. Kgl. Danske Vid. Selsk. Skrifter. 8. Række. 
MNÆNOS6: 


Scrophulariaceae. 495 


In those species in which the main root is vigorous 
and branched, the development of adventitious roots is 
repressed, and does not, at any rate, occur except in old 
individuals (cf. TH. ResvoLz, 1917, p. 215); in the species 
of the other group adventitious roots are more or less abun- 
dantly developed, certainly according to the nature of the 
substratum. The length of the first vegetative-stage varies, 
as a rule, it may be of several-years duration. The variations 
must be assumed to be due to the varyingly favourable 
conditions afforded by the habitat in question, as for in- 
stance, early disappearance of snow in spring. 

In P. lanata the winter-buds are protected by the large 
persistent bases of the foliage-leaves of the year's rosette; 
in the other species special bud-scales occur. 

P. euphrasiotides appears, perhaps exceptionally, to be 
able to occur as a biennial hapaxanth. The growth of the 
main shoot — in all the species which have been investigated 
— terminates by the development of a floral portion, fre- 
quently furnished with a few or several foliage-leaves, but 
sometimes quite devoid of these (individuals of P. sudetica 
and Sceptrum carolinum). The adventitious shoots arise most 
frequently in the axils of the scale-leaves; previous to their 
flowering they usually pass through a vegetative stage of 
some years duration. Only in P. euphrasioides the shoots 
usually appear to be able to achieve flowering in their second 
year. After fruit-setting the upper part of the shoot dies 
away; the complex of the persistent båsal-portions forms a 
rhizoma multiceps (a mesocorme), which is especially large 
and vigorous when the main root is of long duration, as in 
P. hirsuta and lanata; these species also form the largest tufts. 

In those species in which the main root and rhizome 
die away quickly behind, and adventitious roots are abun- 
dantly developed (especially in P. sudetica and Sceptrum caro- 


324 


496 Fr. J. MATHIESEN. 


linum) some vegetative reproduction takes place; but natur- 
ally by this means the plant can spread only to a very limited 
extent. 

According to M. Porsirn's observations P. hirsuta and 
lanata can dispense with a snow-covering during winter; all 
the other species of Pedicularis here mentioned no doubt 
require a snow-covering, but at the same time certainly also 
require that the snow in their neighbourhood shall melt 
rather early in spring; therefore, in the Arctic regions these 
species hardly belong to the real snow-trough flora, as is 
for instance the case with P. hirsuta in Norway. In southern 
Greenland, however, several of the species ascend rather far 
up the mountains (E. WARMING, 1888, p. 87). 

III. Therophyte: Euphrasia arctica. 


Germinates in spring. 


B. The Structure and Biology of the Flower. 

In the large winter-buds of P. sudetica, hirsuta, flammea 
and Oederi the floral organs are found to be highly developed 
in the year previous to that in which they expand, and the 
same is probably the case in the rest of the species of Pedi- 
cularis. Also in Veronica alpina and Bartschia alpina floral 
organs have been demonstrated in the winter-buds. 

The corollas are small and.only slightly showy in Vero- 
nica alpina and Euphrasia arctica; in the former species the 
colour is deep blue, and in Euphrasia pale lilac to white, 
with yellow and violet markings. Veronica fruticans has 
bright blue corollas, which are even as much as 14 mm in 
diameter. Bartschia alpina, Castilleia pallida and the Pedi- 
cularis spp. have rather large flowers with brightly coloured 
corollas (deep bluish violet, red, yellow, and yellow with red 
or brown markings), collected into crowded and often large 


Scrophulariaceae. 497 


inflorescences; Bartschia and P. lapponica have in addition 
coloured bracts. 

Forms with white corollas have been found in Veronica 
alpina, fruticans, P. hirsuta and lanata. 

Honey-secretion, either by a secretory ring around the 
ovary (Veronica) or by protuberances at its base, has been 
demonstrated in the majority of the species. Perfume is 
present in P. lapponica, sudetica, euphrasioides and lanata. 
The species which have been investigated varied from homo- 
gamous to slightly protogynous. 

In Veronica alpina and Euphrasia arctica self-pollination 
is no doubt customary. 

In the Arctic regions self-pollination no doubt plays an 
important part as regards the Pedicularis spp. since, accord- 
ing to several investigators (EksTtAM, PORSILD, etc.), insect- 
visitors are very scarce or even totally wanting there, and 
in Spitzbergen, as mentioned by AuRIVILLIUS, and as has 
since often been pointed out by others, none of the natural 
pollinators of these flowers (humble-bees) are found; in spite 
of this absence, P. hirsuta and lanata regularly set fruit there. 

When, as in P. flammea, the stigma is always included 
in the helmet on account of the shortness of the style, cross- 
pollination by insects seems to be impossible. In P. hirsuta 
(no doubt frequently) and lanata (sometimes) the upper part 
of the style can be found to be so strongly curved that the 
stigma is pressed against the under-side of the anthers; per- 
haps this is due to movement which has taken place during 
a later stage in the flowering, so that originally cross-pollina- 
tion has been possible; in the majority of the flowers of 
P. lanata which have been investigated, the stigma was 
found to protrude slightly beyond the tip of the helmet, as 
also in P. sudetica. In these three species the flowers are set 
almost horizontally, and the stigma is therefore placed under 


498 Fr. J. MATHIESEN. 


the anthers, so that at any rate the dry and light pollen, 
when it falls out of the anthers by accidental shaking of 
the plant, will easily be able to fall upon the almost globular 
stigma.  Self-pollination must take place with somewhat 
more difficulty in P. euphrasioides and lapponica owing to 
the great obliquity of the flower. The latter species, which 
has in addition the most protrusive stigma, is no doubt 
also the worst self-pollinator, and according to Porsirn it 
is also the poorest fruit-setting species in Greenland. 

In P. Oederi, in which the flower, as in P. flammea, 
stands almost in a vertical position, the upper lip curves 
forward and downward at the end of the flowering period, 
by which means the stigma is brought in a position that 
enables it to receive the pollen when it falls. 

Pedicularis Sceptrum carolinum, as Castilleta pallida, no 
doubt normally requires insect-visits for pollination. Bart- 
schia alpina is characterized by great variation in the length 
of its style; im the short-styled forms, especially in those 
in which the stigma is included in the helmet, self-pollination 
must easily be able to take place; any appreciable growth 
of the corolla tube has not been demonstrated in the Arctic 
regions, so the variations must be regarded as individual. 
On the other hand, in several instances of older flowers, the 
anthers were found to project outside the corolla so that 
in these cases we may perhaps assume with KERNER that 
some amount of cross-pollination takes place between older 
and younger flowers by the agency of the wind. 

With the exception of P. capitata I found, in my material, 
ripe fruit in connection with all the species. 


The seeds were small and light in all the species. 


sid 


Scrophulariaceae. 499 


CC: Paårasitism. 


As is well-known the Euphrasia spp., Bartschia alpina and 
the Pedicularis spp. are green semiparasites. 


Ås regards their capability of assimilating the carbon 
dioxide of the air, HE1NnrIcHER (1910, VI, p. 574) states that 
in the Euphrasia spp. this is almost normal, and the same 
author also attributes active assimilation to Bartschia. VOL- 
KART (1899, .p. 30) sums up his results as regards the Pedi- 
cularis spp. (P. recutita, foliosa and verticillata) investigated 
by him as follows: "Die Versuche zeigen, dass die Stårke- 
speicherung der Blåtter der untersuchten Pedicularisarten 
normal und ohne grosse Abweichung von denjenigen anderer 
Pflanzen vor sich geht. Eine weitgehende Herabsetzung der 
Assimilationståtigkeit durch den Parasitismus, wie sie BON- 
NIER aus seinen Versuchen ableitet, findet nicht statt.” 

In this connection I wish to draw attention to the fact 
that the mesophyll, in all the Rhinantheæ spp. investigated 
by me, was found to be normal in structure, and its cells 
contained a normal supply of chlorophyll grains. 

Bartschia alpina according to HEINRICHER's investiga- 
tions, is an obligate parasite. According to VOLKART, the 
Pedicularis spp. may show gradations in their dependency 
on other plants; from P. palustris, which he describes as 
"Stårkster Parasit mit stark entwickelter Neigung zur sapro- 
phytischer Ausnutzung der Nåhrpflanze” (by the destruction 
of the roots attached), to P. comosa, which is said to be a 
Relativ selbeståndig entwichlungsfåhige Art”; as regards 
P. Oederi he writes ”Saprophytische und hydroparasitische 
Ausnutzung der Nåhrwurzel gleichgestellt.” 

From the Arctic regions we lack accounts of thorough 
investigations regarding this matter; but as regards P. lanata 
there is a note to hand which might suggest that under 


500 Fr. J. MATHIESEN. 


certain circumstances this species may at times lead an 
independent life, since M. Porsirn remarks (1920, p. 142) 
that he has found P. lanata growing "in gravelly barren, 
far away from other plants”; where it occurs among other 
plants, this species, also, has haustoria on its roots (1. c.). 

According to VOLKART, in the Alps the Pedicularis spp. 
do not appear to be very particular as regards host-plants. 
From the Arctic regions ROSENVINGE and WARMING records 
P. hirsuta and flammea as parasitic on Vaccinium uliginosum 
and Salix herbacea; Hartz (1905, p. 219) writes that in East 
Greenland P. lapponica "is closely connected with Betula, 
and probably occurs parasitically on its roots.” A rather 
interesting circumstance connected with the present question 
is recorded by Car. KRUUSE (1911, p. 61) from the district 
of Angmagsalik. On some slopes facing north and cut into 
terraces, where a rich vegetation of "espalier-shrubs” was 
growing on the vertical sides of the terraces, he found the 
platforms to be "covered with coarse gravel which bore at 
great intervals (3—5 metres) wind-affected tufts of Salix 
(Salix glauca)”, and in these tufts a Pedicularis (hirsuta or 
flammea) always occurred. "Where the platforms consist of 
sand, Salix herbacea is always predominant, and the latter 
also, with its crooked branches 5—10 cm long, protects a 
Pedicularis.” 


D. Anatomy. 


The Root. Both the species of Veronica have a thin- 
walled, cuticularised exodermis; V. fruticans has a peripheral 
cork-formation in the root (as in the stem), whilst this is 
lacking in V. alpina. In both species the primary cortex 
persists a long time and, especially in V. fruticans, consists 
of rather thick-walied cells; the original endodermal cells are 
elongated during growth and undergo divisions by radial 


Scrophulariaceae. 501 


walls which are quite thin in V. alpina, but of the same 
thickness as in the other cells of the cortex in V. fruticans. 
In the latter species there are distinct growth-zones in the 
xylem-portions of the stele, whilst such growth-zones are 
lacking in V. alpina. 

It is common to the roots of Euphrasia, Bartschia and 
the Pedicularis spp. to have the primary cortex few-layered 
and its cells frequently very thin-walled; root-hairs are 
usually wanting (in some specimens of Euphrasia arctica, 
alone, I found a few scattered root-hairs). 

In the species of Euphrasia the epidermis persists a long 
time; according to HovELAcQUE the same is the case in 
Bartschia, but in this I found that it may sometimes die 
away early. With the exception of P. Sceptrum carolinum 
it dies away very quickly in the Pedicularis spp. which have 
been investigated; even in roots a millimetre thick, I found 
the cells collapsed and partly thrust off; the outer walls of 
the layer below the epidermis become cuticularised, and 
where the growth in thickness of the root is considerable 
(P. euphrasioides, hirsuta, lanata, flammea and Oederi) the 
cells of the exodermis as well as of the other 3—6 layers 
of the primary cortex become greatly elongated tangentially 
and divided by thin radial walls. 

The primary cortex persists a long time. 

In Bartschia alpina and P. capitata a cuticularised 
lamella arises in the wall of the endodermal cells, in the 
former species all the way round, in the latter only in the 
inner and radial walls. 

The Casparian dots were very distinctly discernable in 
Euphrasia arctica, and in P. lapponica, whilst they were 
quite faint in P. hirsuta and lanata. 

Lacunæ, such as are found, for instance, in the primary 
cortex in P. palustris (HovELAcQUE), were, in the species 


502 Fr. J. MATHIESEN. 


investigated, found well-developed in the adventitious roots 
of P. sudetica only. 

In the roots of P. Euphrasioides, hirsuta and lanata, 
which had a vigorous growth in thickness, the rather thick 
secondary cortex was found to be traversed by radiating 
clefts (Fig. 38, B); this was less well-marked in the adventiti- 
ous roots of P. flammea and Oederi. In P. hirsuta and lanata 
in which the thin-walled parenchyma — as is also the case 
in P. flammea and Oederi — is also dominant in the xylem, 
the clefts are often continued far into the latter. 

In the xylem of P. euphrasioides distinct growth-zones 
were seen, each probably corresponding to one year's in- 
crement. 

The thick roots and rhizomes in several of the Pedi- 
cularis spp. certainly contain a considerable amount of 
reserve food material, no doubt especially during the winter- 
rest; the sweet taste of these organs in P. hirsuta and lanata 
is well-known. In P. flammea, Oederi and in the specimen 
of P. hirsuta mentioned on page 462, which was collected in 
late summer, I found the roots full of amylodextrin-starch. 

The condition of the material at my disposal did not 
allow of a closer study of the haustoria. 

The Stem. Ås has frequently been pointed out (Hove- 
LACQUE and SOLEREDER) the absence of parenchyma-rays 
from the part of the xylem formed by the cambium, is a 
character widely distributed among the Scrophulariaceæ. 

Parenchyma-rays were also absent from all parts of the 
shoot in the Veronica spp., Castilleia pallida, Euphrasia and 
Bartschia alpina, and from the above-ground parts of the 
axes in the Pedicularis spp. (in P. lapponica also from the 
runners). In these cases the cambium very quickly develops 
inwardly a continuous ring of vessels and wood-fibres, which 
bridges over the leaf-gaps so that these become demonstrable 


Scrophulariaceae. 503 


on the inner side of the xylem-ring only as thinner portions 
of the latter among "les faisceaux reparateurs”. 

A transverse section of the rhizome in P. sudetica, 
Sceptrum carolinum, hirsuta, lanata, flammea and Oederi 
showed, on the other hand, a circle of vascular bundles 
mutually separated by parenchyma-rays, as is commonly 
the case in fleshy dicotyledonous rhizomes, especially such 
as are abundantly covered with leaves. With respect to 
P.Sceptrum carolinum this feature has been already described 
and figured by CHartin.? In -P.capitata the runners had two 
rather broad diametrically opposite parenchyma-rays. 

In the above-ground parts of the axes in the Pedicularis 
spp. the peripheral portions of the pith consisted usually 
of thickened, lignified and porose cells; this was most decided- 
ly the case in P. Sceptrum carolinum, in the up to one- 
metre-high stem of which the stereom is, on the whole, very 
vigorously developed. When an elongation of the axis takes 
place during flowering and fruitsetting, the lignification of 
the stereom naturally does not begin until this is accom- 
plished. The central portions of the pith are very thin-walled 
and die away early. 

Hard bast was present in the periphery of the stele in 
Veronica fruticans, and very abundantly in the above-ground 
parts of the axis in Bartschia alpina; in the rhizome of the 
latter species the hard bast was present more sparingly, and 
sometimes totally absent. 

Continued growth in thickness through several growth- 
periods could be observed in the persistent basal portions 
of the shoots of the two Arctic species of Veronica (in V. 
fruticans the growth in thickness is continued throughout 
many years — as many as 16 were counted), in Castilleia 


1 CHAtTIN: Anatomie comparée des vegetaux. Plantes parasites. 
Dicotylédones. P. 192. Pl. XLII. 


504 Fr. J. MATHIESEN. 


pallida (the increment was small and irregular) and some- 
times in Bartschia; the runner-like portions of the shoots in 
P. lapponica occasionally show two annual rings; in the 
rhizomes in P. Sceptrum carolinum the xylem of the vascular 
bundles always showed two growth-layers, since in the 
second year of the rhizome-portion a group of vessels (and 
stereom) is developed outside that produced in the first 
year; the two groups are distinctly separated, some layers 
of thin-walled unlignified parenchyma being found between 
them. Thin-walled and unlignified cells are also present in 
the Veronica spp. at the boundary between the annual rings. 

Cork-development occurs sub-epidermally in Veronica 
fruticans in the basal, persistent portions of the shoots, and 
in P. Sceptrum and capitata further within the primary 
cortex; the cork is always only few-layered, but P. Sceptrum 
carolinum develops, in addition, cork-cambiums in the paren- 
chyma between the annual increments in the vascular 
bundles and also within the pith. In the majority of cases 
the endodermis was demonstrable by the presence of Cas- 
parian dots, or else a cuticularised lamella was found in its 
walls, either all the way round (as in the runners in Bart- 
schia) or only on the inner and radial walls (runners of P. 
capitata). 

The Leaf. All the species investigated have leaves 
mesomorphic in structure. The cuticle is thin; in Veronica 
fruticans a moderate thickening of the walls of the epidermal 
cells takes place, the other species have thin walls. The 
lateral walls of the epidermal cells of Castilleia pallida are 
almost straight, of the other species more or less decidedly 
undulating; in Euphrasia arctica and Bartschia alpina the 
difference between the epidermis of the upper and lower 
surfaces is in this respect inconsiderable; im the species of 


Veronica and Pedicularis I found that the epidermal cells of 


Scrophulariaceae. 505 


the lower surface had the most decidedly undulating lateral 
walls. 

The lateral walls of the epidermis in Veronica alpina, 
Castilleta pallida and Euphrasia arctica are without sculp- 
ture; in the other species they are porose; in Veronica fru- 
ticans and Pedicularis euphrasioides, flammea and Oederi the 
pores are distinct and alternate with flanged thickenings, 
whereas they are fine in Bartschia alpina and the other 
Pedicularis spp. It should be noted, however, that the epi- 
dermal cells of the lower surface which bear the large glan- 
dular hairs have (except in P. lanata) very vigorously devel- 
oped flanged thickenings in the Pedicularis spp., and usually 
also in Euphrasia and Bartschia; also their walls are, on the 
whole, distinctly thicker than those of the surrounding epi- 
dermal cells. 

Stomata occur exclusively on the lower surface of the 
leaf in the Pedicularis spp., nearly equally on both surfaces 
in Veronica fruticans, V. alpina, Castilleia pallida and Eu- 
phrasia arctica, and more abundantly on the lower surface 
in Veronica officinalis and Bartschia alpina. The guard-cells 
are on a level with, or raised only slightly above the leaf- 
surface; they are surrounded by 3—8 cells. 

The mesophyll in all cases is rather lacunose and consists 
of thin-walled cells. In Castilleia pallida it may be almost 
homogeneous; 2—3 layers of short palisade-cells occur in the 
Veronica spp., Euphrasia arctica and Bartschia alpina. Of the 
Pedicularis spp., P. euphrasioides, capitata, hirsuta, flammea, 
Oederi, and frequently lapponica, had 1—2 layers of typically 
developed palisade-cells; they were shorter and broader in 
P. Sceptrum carolinum, sudetica and lanata. 

In P. flammea and Oederi there were large air-chambers 
between the spongy parenchyma and the epidermis of the 
lower surface (cf. Fig. 46, A). In all the species investigated, 


506 Fr. J. MATHIESEN. 


the cells of the mesophyll contained a normal number of 
chlorophyll-grains. In the majority of the cases chlorophyll- 
grains were also present in the epidermis of the lower surface. 

Only P. lapponica, Sceptrum carolinum and flammea 
appear to be totally devoid of non-glandular hairs. The 
leaves were densely covered with hairs in Castilleia pallida, 
Euphrasia arctica, Bartschia alpina, P. hirsuta, lanata (in the 
last-mentioned two species especially along the edges of the 
petiole on the part near the base of the leaf) and capitata 
(on the lower leaf-surface and the petiole). The non-glandular 
hairs are from one to several-celled; in Veronica fruticans 
and Pedicularis capitata they are rather thick-walled, in the 
others thin-walled, smooth or with fine cuticular striations; 
only Veronica fruticans had hairs with coarser cuticular 
warts. 

Glandular hairs were present in all the species: small 
glandular hairs with a one-celled stalk and two-celled head 
in the Veronica spp., Castilleia pallida and in especially great 
numbers in Euphrasia and Bartschia alpina; the last-men- 
tioned two species had in addition large, peltate glands with 
a low sunk stalk-cell and a 2—4 celled flat head, like those 
in Lathræa (SCHERFFEL, HABERLANDT and GOEBEL; com- 
pare my Figs. 14 and 19). As previously mentioned by Hove- 
LACQUE and PEercY GROOM, large glandular hairs of peculiar 
structure occur also in the Pedicularis spp.; here the low 
stalk-cell is not sunk, and the head is 2-celled (see for in- 
stance" Fig.23: "Aand CC). Al Fthelspecies "or Pedicularis 
which have been investigated had such glands. 

The above-mentioned special forms of large glands are 
found only on the lower surface of the leaf and, as pointed 
out by the above-mentioned authors and as may be seen 
from my figure 24 and from most of the transverse sections 
of leaves figured, they are localised under the vein-branches 


Scrophulariaceae. 507 


of higher order which at their tips consist only of a few thin- 
walled tracheids. According to information to hand it seems 
unquestionable that these glands are organs for the secretion 
of water. 

As to the Pedicularis-glands it should be noted that 
special water-paths from the tracheids to the basal cell of 
the glandular hair, limited by the cuticularisation of cell- 
walls, as have been demonstrated in P. palustris by PERCY 
Groom, have not been able to be demonstrated in any of 
the species investigated here. AÅ bursting and crumbling- 
away of the cuticle on a small circular spot in the middle 
of the glandular head, such as the author in question men- 
tions in connection with P. palustris, was very common in 
P. lapponica and P. Sceptrum carolinum, and was occasion- 
ally also found in other species. 

Glandular hairs with a 2 to several-celled stalk and a 
2—4 celled head were present in Castilleia pallida, Euphrasia 


arctica, Bartschia alpina, and also in P. flammea. 





TYP. BIANCO LUNO, KBHVN. 


4. 


IEHESBRYOPEYEPAOFTEEBAND 


BY 


AUG"HESSEEBO 


WIEHS39PFIGURESSINSEHESPEXT 


Arbejder fra den botaniske Have i København. 








INTRODUGTION. 7; 


er to the time when Grånlund made botanical collections 
in Iceland (1868 and 1876) the Bryophyta of the island was 
very little known. Earlier botanists, who occasionally collected speci- 
mens also of the Bryophyte Vegetation, have only exceptionally given 
information as regards localities, and the older lists consist merely of 
an enumeration of species without any information as regards dis- 
tribution or frequency. To this must be added that the older deter- 
minations of species are very uncertain, and in many cases un- 
doubtedly erroneous or quite improbable, and that the lists are 
compiled quite uncritically without stating the name of the collector 
or the source of the information. For instance, more than half of 
the 149 mosses and 54 hepatics enumerated in Lindsay's list are 
undoubtedly wrongly determined, and the majority of them beyond 
doubt do not occur in Iceland. Therefore, in preparing the following 
list of the Bryophyta of Iceland, older records have been taken 
into consideration only when it has been possible to verify them 
by means of specimens in the collections. 

The material dating from older collections (previous to Gron- 
lund”s) is but very scanty. Mårch was the first to contribute 
anything of importance to our knowledge of the Bryophyta of 
Iceland. In 1820 he collected a rather considerable number of 
species, among which are many that were not found again until 
quite recently, and even several which have not since been found 
by others. Moårch's collections are in the Botanical Museum in 
Copenhagen, but unfortunately he has in only quite a few cases 
recorded the habitat on the wrappers. W. J. Hooker (1809), 
Lindsay (1860) and Caroll (1861) collected mosses and hepatics 
in Iceland, but as mentioned above, Lindsay's list, in which 


398 A. HESSELBO 


these finds åre enumerated, is so full of mistakes that it is quite 
worthless. 

Japetus Steenstrup collected (1839 —40) some Bryophyte 
samples in West and North Iceland and in the interior of the 
country. 

But it was not until Gronlund's investigations that the foun- 
dation was laid of a real knowledge of the Bryophyte Vegetation 
of Iceland, he not only critically revised the older lists, but also 
prepared lists of his own collections. It was also Gronlund 
who was the first to give real descriptions of moss societies. 
Groånlund's journeys in 1868 and 1876 extended over West and 
North Iceland, from Reykjavik across Esja to Borgarfjérdur and 
thence to Skagafjérdur, Eyjafjordur and Myvatn. He did not visit 
East, South and North-west Iceland. 

In recent times the Iceland botanists Olafur Davidsson, 
Stefån Stefånsson and Helgi Jønsson have contributed con- 
siderably to our knowledge of the Bryophyta of Iceland. 

Olafur Davidsson (died 1902) made, during 1899—1900, very 
considerable and valuable collections in North Iceland around 
Eyjafjéordur and Skagafjérdur and on the small island of Grimsey 
north of the Arctic Circle. His collections, which belong to the 
Reykjavik Museum, have only now been determined. 

Stefån Stefånsson, the head of a school in Mådruvellir and 
member of the Althing up to 1916, has made collections during 
many years, especially in North and North-east Iceland. The 
greater part of his materia! is published in Gronlund 's list. 

Helgi Jonsson, Dr. phil. and teacher in Reykjavik, on his 
numerous journeys to almost all the different parts of the island, 
has everywhere collected specimens of the Bryophyte Vegetation, and 
in his descriptions of the vegetation of Iceland has usually men- 
tioned the Bryophyta also. His collections, which are preserved 
in the Botanical Museum in Copenhagen, are determined by C. Jen- 
sen. Besides the botanists mentioned above, a few others have 
also occasionally collected specimens of the Bryophyte Vegetation. 
C. Ostenfeld, during a few short visits, has especially investigated 
the lava-fields and the hot springs near Reykjavik and on the 
peninsula of Reykjanes. A. Feddersen (1884—86) made some 
collections in South and West Iceland. Dr. C. Hansen, Professor 
Th. Thoroddson and Pétur Sophoniasson collected mosses 


THE BRYOPHYTA OF ICELAND 399 


in several places. The majority of these collections are determined 
by C. Jensen and included in Grénlund's list (1895). 

My own investigations were made on three journeys, viz. in 
1909 and 1912 by the aid of the Carlsberg Fund, and in 1914 by 
the aid of the Botanical Travelling Fund, and Japetus Steen- 
strup's Legacy. In 1909 I investigated in the beginning of June 
the region around Reykjavik and Hafnarfjordur, and made a five 
day's trip to the Esja mountains. . On June 9th I sailed to Horna- 
fjordur in South-east Iceland, and after staying a few "days there 
went straight through East Iceland to Seydisfjåordur, where I arrived 
on June 30th. On the way I stopped at several farms, viz. Stafa- 
fell, Hof, Djupivogur, Berufjérdur, Håskulstadr and Vallanes. On 
July 9th I took the coast steamer to Husavik in North Iceland, 
and after some days sojourn there went into the interior of the is- 
land across Ås and Svinadal to Reykjahlid near Myvatn, and after 
having stayed a few days there to Akureyri. Unfortunately investigation 
in this interesting region was almost entirely prevented by continual 
rain and fog. I stayed five days in Akureyri and then continued 
my journey westwards through North Iceland to Stadur near 
Hrutafjordur, stopping at the farms Tyverå in Øxnadalur, Vidimyri, 
Vidivellir, Geitaskard, Hnausar and Lækjarmot. From Stadur I 
rode across Haukadalsheidi and Brattabrekki to the farm Dals- 
mynni in Nordredalur, where I arrived on August 2nd. After stay- 
ing a few days there in continual rain, I was obliged to abandon 
my original plan of travelling by land to Reykjavik and, instead, 
to ride to Borgarnes and take the steamer to Reykjavik. 

In 1912 I arrived at Reykjavik on June 2nd and rode thence 
past Kolvidarhol to Reykir, where the vegetation around the "hot 
springs was investigated, and thence to Skålholt, Laugardalur and 
Thingvellir, staying a few days at each place, and then back again 
to Reykjavik, whence I sailed on June 1åth to Dyrafjérdur in 
North-west Iceland. On the way I had an opportunity during a 
24 hours stay in Stykkisholmur to investigate the immediate neigh- 
bourhood. After a three days stay at Dyrafjordur I rode on June 
20th to Isafjordur, whence, during the following three weeks, I 
made numerous excursions, partly, on foot, to the surrounding 
country and to Sugandafjordur, and partly to the north-west side 
of the fjord and the district around Jåkulsfjéordur, to Hesteyri and 
the whole tract of land from the head of Jåkulsfjérdur to Stadur, 
and from Bæir to Arngerdareyri. 


400 A. HESSELBO 


On July 10th I returned to Reykjavik by the steamer "Botnia”, 
and rode thence through Esja and around Hvalfjårdur to Reykholt- 
dalur, where I stayed some days and investigated the hot springs, 
and made an excursion to tbe farm Gilsbakki. On my way back 
I passed through Reykholtdalur and Lundurreykjadalur to Thing- 
vellir, and thence to Reykjavik. 

On my last journey in 1914 I stayed on the island of Vest- 
mannaey, which I had previously had opportunity to visit only for 
a few hours, on my way to Reykjavik, from the 4th to 9th July. 
I went to Reykjavik on the 10th, and rode on the same day to 
Kolvidarhol, and thence further across Reykir and Thjorsårtun to 
Fljotshlid near the Markarfljot, where I stayed some days at the 
farms Breidabolstadr and Barkarstadr. On the 17th I rode across 
the great river Markarfljot to the vicarage of Holt, south of Eyja- 
fjallajåkull, and stayed partly there and partly at the farms Drångs- 
hlid and Seljaland until July 24th, when I recrossed the Markar- 
fljot on horseback, and rode past the farms Thjorsårtun and Skål- 
holt to Geysir, and thence across Thingvellir to Reykjavik, where 
my journey ended abruptly on August l1st on account of the out- 
break of war. 

Ås may be seen from the above survey, Iceland is as yet very 
incompletely investigated as regards bryology. All previous collec- 
tors, with the exception of Gronlund, had no, or almost no, know- 
ledge of the Bryophyta, and have, therefore, only been able to 
take chance specimens, which must, therefore, often give a mis- 
leading idea of the composition of the vegetation. The great 
distances, together with a slow and toilsome journey on horse- 
back, render investigation difficult, and rain and fog often greatly 
delay the work, and make it almost impossible to travel in the 
mountains. 

The most thoroughly investigated regions are South-west Iceland 
from Eyjafjallajåkull to Borgarfjordur, East Iceland from Horna- 
fjéørdur to Seydisfjordur, the districts around Eyjafjérdur in the 
north and around Isafjardardjup in the north-west. Throughout 
large parts of the country møsses have, practically, never - been 
collected; thus, in the north-east from Seydisfjordur to Axarfjérdur, 
in South Iceland from Vik to Hornafjérdur, in the greater part of 
North Iceland and in almost the whole of the large interior plateau. 
Therefore this 'list of the Bryophyta found in Iceland can by no 


THE BRYOPHYTA OF ICELAND 401 


means be regarded as complete, and future investigations will un- 
doubtedly effect great alterations in our present knowledge of the 
distribution of the species in the different parts of the country. 


ABBREVIATIONS OF NAMES. 


O0.D. = Olafur Davidsson 
F. = Feddersen. 

Grl. = Grønlund. 

Ho. =—< Hornemann. 

HJV FHelsi Jonsson; 
CF = 6. Jensen: 

Ostf.  — C. H. Ostenfeld. 

SE — Stefån Stefånsson. 
Stp. =— Japetus Steenstrup. 
For (l="ASHesselbo: 

Q! = The specimen seen by A. Hesselbo. 


M. B,H. = The Botanical Museum of Copenhagen. 


402 A. HESSELBO 


-FISTFOFSTIE BROEN EA 


IEEE P ASE TE: 


Fam. RICCIACEÆ. 


1. Riccia bifurca Hoffm. 


N. Iceland: Hrossaberg (St.)!; Vidimvri (Grl.)!; Myvatn!. S. Iceland: 
Near Reykjanes lighthouse (Ostf.)!; Kverkfjall (Wegener.)!. 

In all these localities it grows on warm, damp clayey flats near 
hot springs. Near Myvatn — where it grew abundantly around fuma- 
roles, from which issued aqueous vapours containing sulphuretted hy- 
drogen — the temperature of the soil was about 40”! about one cm. 
below the surface. Near Reykjanes lighthouse Ostenfeld (Bot. Tids- 
skrift, vol. XXII, 1899, p. 239) gives the temperature of the soil as 20” 
—30". All the plants investigated bore capsules. 


2. Riccia sorocarpa Bischoff. 


N. Iceland: Reykjalaug in Fnjoskadal (O.D.)!. S. Iceland: Grafarbakki 
near the Minni Laxå (F.)!; Laugaråshver!. 

Like the preceding species, RØW. sorocarpa grows on warm, damp 
clayey flats. Near Laugaråshver it was found abundantly on clayey 
slopes stretching down towards the boiling hot spring. There the tem- 
perature of the soil was about 257—30". All the plants investigated 
bore capsules which in the plants from Laugaråshver were fully ripe 
at the end of July. 


3. Riccia crystallina L. 


S. Iceland: Thoørlakshver near Skålholt!. (27. 7. 1914). 

The plants, which had fully developed capsules, grew widely scat- 
tered on. clayey flats, otherwise bare of vegetation, around some small 
holes from whence issued boiling water accompanied by vapours slightly 
impregnated with sulphuretted hydrogen. 

Note. Riccia glauca L. is mentioned by Zoéga and is recorded 
by Hornemann as occurring abundantly near Geysir. But as no speci- 
mens of it are to be found in the collections, and during the search 
made in 1914, no species of Riccia were found near Geysir, it is im- 
possible to determine what species is meant. 


" Here and throughout the book the centigrade scale is meant. 


THE BRYOPHYTA OF ICELAND 403 


Fam. MARCHANTIACEÆ. 
4. Sauteria alpina Nees. 


N. Iceland: Grænuhnukr in Brattafjallgardi between Mådruvellir and 
Bru (St.). 


5. Reboulia hemisphærica (L.) Raddi. 


SW celand:" Hafmarfordur FE) SFS Med and FDrångsblidt (PI: 
Holt!; near Skågafos!; Seljaland'!. 

This plant was found only in the southern part of the country, 
but it appears to occur there rather frequently. Near Hafnarfjordur it 
was found in a lava cave; in S. Iceland it was collected everywhere 
from the somewhat damp faces. of tuff rocks, especially from the base 
of the sides of illuminated caves in tuff cliffs; in this situation it usually 
grew in company with Preissia commutata, Fegatella, Marchantia, Anoec- 
tangium compactum, Distichium montanum, etc. It was found every- 
where in fruit which was almost ripc at the end of July. 


6. Fimbriaria pilosa (Wahlb.) Taylor. 


N. Iceland: Hof (0. D.)!; Boggverstadadalur (0. D.)!. NW. Iceland: 
Bæir on Snæfellsstrand!. 

Near Bæir it grew abundantly on a stony slope facing south and 
stretching down towards the sea, upon rather dry, humus-covered rocks. 
It was everywhere collected in fruit which near Bæir had just ripened 
opf June 6 MOT 2 


-= 


1... Fegatella conica Corda. 
Hepatica conica Lindb. 


S. Iceland: Drångshlid (H. J.)!; Merkjåfoss (F.)!; Breidabolstadr!; 
Barkastadr!; Seljaland!; Holt!; Drångshlid!; Skågafos!. 

This plant was not found except in S. Iceland, but there it was 
common in all the parts which were investigated. It grew there every- 
where on faces of tuff rocks, especially on those of caves and clefts, 
and more particularly on the faces with a southern exposure where it 
may occur very abundantly, sometimes on the damp tuff rocks them- 
selves and sometimes creeping over mosses, for instance Hypnum fili- 
cinum, Eurynchium Swartzii, etc. Sterile specimens only have been found. 


8. Preissia commutata (L.) Nees. 
Chomocarpon quadratus (Scop.) Lindb. 


N. Iceland: Hof, fr. (0. D.)!: Modruvellir (0. D.)!; Hrossaberg on 
warm clayey flats, fr. (St.)!; Stora Gjå near Myvatn!.  NW. Iceland: 
Stadr on Snæfellsstrand, fr. (Stp.)!.  W. Iceland: common near the hot 
springs in Reykholtdalur!; Hafnarfjordur!. S. Iceland: Thingvellir (Grl.;! 


404 A. HESSELBO 


Laugardal (Grl.;”); Reykjanes near hot springs (Ostf.)!; common every- 
where on faces of tuff rocks in Fljotshlid and below Eyjafjallajokull!. 
Preissia commutata occurs both on more or less damp ground and 
on damp rocks, and also on warm clayey flats near hot springs. In 
the last situation, as in Reykholtdalur, it covers large arcas on slopes 
stretching down towards the boiling hot water. In E. Iceland it has 
not yet been found; in N. and NW. Iceland it is rather rare and grows 
ihere on damp rocks, along river-banks or in rock-clefts. In S. and 
SW. Iceland it is frequent in lava-clefts near Thingvellir and Hafnarfjordur, 
and "in Ss Iceland proper it us very common Theretitterowstatktle 
base of, or some way up the rock faces, especially upon those with a 
southern exposure, or in sheltered clefts, and often occurs in great 
abundance, and fruiting richly. The fruit was not quite ripe even at 
the end of July. 


9. Marchantia polymorpha L. 


Common all over Iceland on damp soil, on rocks, by rivers, in 
marshes and in moss bogs, and very often found with fruit or anthe- 
ridiophores. It is extremely common especially in NW. Iceland, and 
covers banks of streams often in great abundance, to a height of about 
300 metres. In S. Iceland it is also very common on damp tuff rocks. 

This species is most widely' distributed in. the birch "zone"toa 
height of about 300—400 metres above sea-level where it usually grows 
on marshy or gravelly ground along small streams, or occurs in abun- 
dance in moss bogs or in marshes among mosses. But it is also met 
with upon mountain heights, as for instance near Berufjéordur, Seydis- 
fjordur and Isafjéordur, where it has been collected abundantly up to a 
height of 600—700 metres. 


Fam. ANEUREÆ. 


10. Aneura pinguis (L.) Dum. 
Riccardia pinguis (L.) Gr. 


Very common on damp ground both in marshes and on gravelly 
soil by rivers and also on damp rocks. It usually grows intermixed 
in the tufts of other Bryophyta, rarely forming tufts by itself; it has 
been collected in fruit in a few localities only (near Eyjafjordur, and 
near Seljaland at an altitude of about 500 metres). It is most widely distri- 
buted in the lowlands up to about 300 metres, but it is also frequently 
met with upon mountain heights. 


11. Aneura multifida (Lindb.) Dum. 


Riccardia multifida (Lindb.) Lindb. 


E. Iceland: Lon!, on swampy ground among Sphagnum rubellum; 
W. Iceland: Reykholtdal!, frequent near hot springs intermixed in the 


THE BRYOPHYTA OF ICELAND 405 


tufts of other Bryophyta such as Sphagnum rubellum, Scapania irrigua, 
and Pellia Neesiana. In one place it was growing on a stone which 
protruded slightly above hot water of a temperature of about 50: 
there it was growing intermixed in a tuft of Enthostodon ericetorum, 
Scapania irrigua and Anthoceros punclatus. 

This species is recorded by Morch from Iceland, but no speci- 
mens of it are to be found in the collections. 


12. Aneura latifrons Lindb. 


Riccardia latifrons (Lindb.) 


N. Iceland: several places near Eyjafjordur (0. D.)!; Akureyri!; 
Husavik!. NW. Iceland: Laugarland!. SW. Iceland: Reykjavik!; Kolla- 
fjordur!. 


This plant usually grows on peaty soil intermixed in the tufts of 
other Bryophyta such as Dicranella crispa, Lophozia Kunzeana, Sphagnum 
spp. and Aneura pingvis. Near Husavik it was growing among Lepto- 
bryum pyriforme on damp gravelly ground. 


FAM METZGERIEÆ 


13. Metzgeria furcata (L.) Lindb. 


Commonly distributed all over Iceland in dry rock-clefts and on 
rock-faces, usually associated with Radula complanata. In S. Iceland, 
where it is very common on rock-sides in clefts of tuff rocks, it is 
sometimes found also in rather damp localities. Only sterile specimens 
have been found. 

Metzgeria furcata is a typical lowland-plant. In Iceland it has not 
been found at a higher altitude than about 300 metres. 


Fam. HAPLOLÆNEÆ. 
14. Pellia Neesiana (Gottsche) Limpr. 
Marsilia Neesiana Lindb. 


Very common on wet ground especially along river-banks, in moss 
bogs and on inundated ground; somewhat more rare in marshes. It 
grows almost always intermixed in the tufts of other Bryophyta, and 
usually as an erect, elongated form among Philonotis, Mnium cinclidioides, 
Acrocladium cuspidatum, Marchantia, etc., more rarely it grows in un- 
mixed tufts on damp ground. Fruit appears to be produced rather 
rarely and also rather sparinglv. 

This species has its main distribution in the lowlands up to about 
300—400 metres, and it is only by exception that it has been found 
above this level. 

Pellia epiphylla (L) Lindb. is enumerated in the majority of the 


406 A. HESSELBO 


older lists, and Groånlunds records it from many localities in Iceland, 
but all the specimens found in the collections under this name are 
those of Pellia Neesiana. 


15. Blasia pusilla L. 


N. Iceland: Reykum (Grl.)!; Melar (Grl.)!; several places near Mådru- 
vellir near Eyjafjérdur (0. D.)!; Reykjalaug in Fnjoskadalur (0. D.)!: 
Oxnadalur!; Stadr near Hrutafjordur!. W. Iceland: Gilsbakki!; common 
in Reykholtdalur!. S. Iceland: Reykirdalur!. 

Only the &g plant has been found in Iceland. 

This species occurs in rock-clefts and along the banks of streams, 
on wet gravelly ground, and also on clay near hot springs. For instance, 
it grows abundantly in Reykholtdalur, and more scantily in Reykir- 
dalur on the damp, steaming clayey flats, the temperature of which 
just below the surface is about 257—40”; it is found associated with 
Gymnocolea inflata, Haplozia crenulata and Fossombronia Dumortieri. 

On banks of streams it usually grows rather scantily and associated 
with Dicranella crispa, Pellia Neesiana, Didymodon rubellus, Angstromia 
longipes, Scapania subalpina, etc. 

Note... Mårchia Blyttii (Mérch) Brockm. is recorded by Gronlund 
to occur in Brynjudalur, but the specimens preserved in the Botanical 
Museum in Copenhagen belong to Blasia pusilla. 


BEAM TCODONIEÆ: 


16. Fossombronia Dumortieri (Hub. et Genth) Lindb. 


N. Iceland: near Myvatn!. S. Iceland: near hot springs on Rey- 
kjanes (Ostf./); Reykir!; Thorlåkshver!; Laugaråshver!; Sydri Reykjahver!; 
Thingvellir!; Kolvidarhél!; Geysir (H0.;!). W. Iceland: common near 
hot springs in Reykholtdalur!. 

Fossombronia Dumortieri is one of those species which are rarely 
absent from the neighbourhood of any hot springs if only the water 
does not contain too much sulphuretted hydrogen. There it grows on 
warm clayey flats with a temperature of about 40”, and especially on 
damp slopes along the outlets of springs; as a rule it sets fruit which 
ripens during June—July. 

In Reykholtdalur it is common near all the hot springs, also near 
ihe springs on Riskupstungur, near Geysir and on Reykjanes. In Reykir- 
dalur, where the majority of the springs give out abundance of sul- 
phuretted hydrogen in their vapours it occurred less frequently, but 
nevertheless was found near several springs, also around some small 
fumaroles near Kolvidarhol from which issued aqueous vapours im- 
pregnated with sulphuretted hydrogen. It grows in all these localities 
associated with other Hepaticæ, for instance Gymnocolea inflata, Haplozia 
crenulata and Anthoceros punctatus. Near Myvatn it was growing around 
2 fumarole, on a damp clayey flat with a temperature of about 40”, 
associated with Riccia bifurca and Haplozia crenulata. The only place 


THE BRYOPHYTA OF ICELAND 407 


where the plant has been found on other than warm ground is near 
Thingvellir where it was growing in a ditch along the road to Hrafnagjå, 
on clay washed together into a heap; there it had for companions 
Dicranella crispa, Didymodon rubellus, Eucalyx subellipticus and tiny 
plants of Pohlia, Bryum, etc. 


Fam. EPIGONANTHEÆ. 


17. Gymnomitrium corallioides Nees. 


S. Iceland: Thingvellir (Stp.; Grl.;!); Seljaland (Stp.;!). SW. Iceland: 
Reykjavik (Grl.;!); frequent in Esja!; Hafnarfjordur!. E. Iceland: frequent!. 
N. Iceland: Akureyri!. 

Occurs rather commonly, but as a rule, not abundantly in SW., E. 
and probably also N. Iceland; in NW. Iceland it has not yet been found. 
It usually grows on the top of blocks of basalt and lava in small greyish 
cushions, and rarely on the dry gravelly soil of Grimmia-heaths. On 
the heath in Seljaland it occurred on blocks of basalt up to a height 
of about 400 metres; in Esja it has bcen found up to about 500 metres. 
Only sterile specimens have been found. 


18. Gymnomitrium concinnatum (Ligthf.) Corda. 


Commonly distributed over the whole of Iceland. 

In the lowlands it is especially common in the lava-fields, where 
it grows both on the top of blocks of rock and in crevices and caves, 
generally mixed with other Hepaticæ such as Lophozia alpestris, L. quin- 
quedentata and Pleuroclada albescens v. islandica; but it is also frequently 
met with in clefts mixed with other Bryophyta and creeping over stones. 
But it has its greatest distribution from about 300—400 metres upwards 
towards the snow-line, where it grows both on more or less damp 
ground and on rocks either in low, extensive cushions or mixed with 
other Bryophyta such as Lophozia alpestris, L. ventricosa, Pleuroclada 
albescens, Dicranum Blyttii and D. fulvellaum, usually also woven into 
dense cushions of Conostomum boreale. 

In NW. Iceland especially, where it is one of the most frequently 
occurring Bryophyta, it grows on gravelly flats on mountain heights in 
extensive carpets associated with Anthelia nivalis, Alicularia minor, Poly- 
trichum sexangulare and the other above-mentioned Bryophyta. Near 
Seljaland, at about 650 metres, it was growing in great masses on 
weathered basalt rocks associated with Marsupella emarginata, Poly- 
trichum sexangulare and Dicranum Blyttii. 

Fruit not rare; it ripens, according to altitude, during June—July. 


19. Gymnomitrium varians (Lindb.) Schiffner. 


NW. Iceland: Dyrafjérdur! (at an altitude of about 350 metres) on 
damp gravelly ground; Reykjaheidi, in a lava-cleft!. Near Dyrafjérdur it was 
growing abundantly in low, blackish-brown mats associated with Anthelia 
nivalis and Alicularia minor on ground saturated with melting snow. 


408 A. HESSELBO 


20. Gymnomitrium revolutum (Nees) Phil. 


NW. Iceland: Sugandafjordur! (at an altitude of about 200 metres). 
It occurred here, on wet gravelly soil upon a slope, scantily among 
Hypnum callichroum, Dicranum Starkei, Scapania uliginosa and Lophozia 
Floerckei. 


21. Marsupella Funckii (W. et M.) Dum. 


E. Iceland: Hof!, in brownish-black cushions about one cm. high, 
in rock-clefts associated with Lophozia alpestris and Dicranum Andersoni; 
Seljaland! (at about 350 metres), on the ground between blocks of basalt 
and mixed with Marsupella emarginata and Dicranum Blyttii. 


22. Marsupella emarginata (Ehrh.) Dum. 


NW. Iceland: Dyrafjordur on damp rocks at an altitude of about 
250 metres!. S. Iceland: Holt! (at about 400 metres), on marshy ground 
among Hypnum sarmentosum, Oncophorus virens, etc.; Seljaland!, in 
several places from an altitude of 350 to 650 metres, both on gravelly 
soil and on rocks. 

This species is recorded by Morch from Iceland (figured in Flora 
Danica. tab 1945), and by Grønlund from Thingvellir and Hafnarfjordur. 
The specimen from Hafnarfjårdur has proved to be Anthelia julacea. 
Mårch's specimens are not to be found in the collections. 


23. Marsupella aquatica (Lindb.) Schiffner. 


NW. Iceland: Kaldalon! by the bank of a small lake intermixed in 
a tuft of Hypnum sarmentosum and H. exannulatum; Isafjérdur!, abun- 
dantly in an almost dried up river, at an altitude of about 250 metres. 


24. Alicularia scalaris (Schrader) Corda. 


Very common over the. whole of Iceland. 

Alicularia scalaris is one of the most commonly occurring Hepaticæ 
in Iceland, and is of almost equal frequency in the lowlands as in the 
higher mountainous regions. Fruit is rather common. 

It grows especially on a somewhat damp substratum, both on rocks 
and on gravelly ground and also by streams and in bogs, sometimes 
as pale green or — in exposed localities — brownish mats, sometimes 
sprinkled in the tufts of other Bryophyta. On peaty ground it often 
forms extensive, continuous carpets in company with Pogonatum urni- 
gerum. On warm clayey flais it also occurs abundantly, and forms 
together with Haplozia crenulata dense, reddish-brown or pale green 
mats along the warm water near both sulphurous and alkaline springs. 


THE BRYOPHYTA OF ICELAND 409 


25. Alicularia geoscypha De Not. 
Nardia minor (Nees) Arnell, N. haematosticta Lindb. 


Commonly distributed all over Iceland and, as a rule, fruiting. 

This species doubtless occurs as frequently in the lowlands as on 
mountain heights upwards to the limit of plant-growth. It is generally 
met with on damp gravelly ground associated with other Hepaticæ, for 
instance by streams with Cephalozia bicuspidata, Scapania subalpina, S$. 
curta, Lophozia alpestris and L. Wenzelii, or on damp gravelly ground 
on rocky flats with Pohlia gracilis, P. commutata, Aongstroemia longipes 
and several other species. On mountain heights, especially in NW. 
Iceland, it generally grows in company with Anthelia Juratzkana on gra- 
velly flats irrigated by melting snow. 

Var. insecta (Lindb.) K. M. was collected near Isafjårdur (at about 
300 metres alt.) together with the type. 

Note. Alicularia compressa (Hooker) Nees is figured in Flora Danica, 
tab. 1772, fig. 2, and is recorded to have been found near Gronnefjord (?) 
by Morch, but no specimens of it are to be found in the collections. 


26. Eucalyx subellipticus (Lindb.) Breidler. 


E. Iceland: Berufjordur!; Seydisfjordur!. N. Iceland: Hof near Eyja- 
fjordur (O. D.)!; Oxnadalur. NW. Iceland: Kaldalon!; Laugarland!; Grun- 
navik (at about 350 metres alt.); Gnupsdalur!. VW. Iceland: Gilsbakki : 
Esja, several places!; Kolvidarh6l!. S. Iceland: Thingvellir, several places !. 
It has been collected everywhere in fruit. Å 

It is no doubt rather common but often overlooked on account of 
its diminutive size and resemblance in habit to the far more frequently 
occurring small species of Alicularia and Haplozia, with which it is 
generally associated. It is usually found on damp gravelly ground, 
more rarely on damp rocks or in lava-clefts; now and then in unmixed 
tufts, but generally mixed with Alicularia scalaris and A.geoscypha, Sca- 
pania curta, Dicranella crispa, Pohlia spp., etc. Near Hof it grew mixed 
with Haplozia pumila; in Almannagjå it occurred in many places at the 
bottom of clefts or on humus-covered ledges associated with Lophozia 
alpestris, Alicularia geoscypha and Scapania subalpina. Near Seydisfjordur 
it occurred creeping over stones by the river and mixed with Haplozia 
atrovirens. 


27. Haplozia crenulata (Sm.) Dum. 
Nardia crenulata. 


Near Geysir (Morch; Stp.!); figured in Flora Danica, tab. 1774, 
fig. 1, "in uliginis Islandiæ frequens, A. Mårch.” Common near all the 
hot springs in Reykholtdalur!, Reykirdalur!, on Biskupstungur!, on Rey- 
kjanes (Ostf.)!, near Laugarvatn!, in Lundurreykjadalur! and near Mvyvatn'!. 
It was found also in W. Iceland: Borgarnes!, at several places along the 
road; Lækjarmot!, on wet ground in bogs. NW. Iceland: Laugarland!, 
on boggy ground. 

The Botany of Iceland. Vol. I, part II. 


[NS] 
<Q 


410 A. HESSELBO 


Haplozia crenulata is in Iceland a decidedly warm-soil species which 
is hardly absent from any hot spring, whether sulphurous or alkaline. 
It grows there in abundance on warm clayey flats which have a tem- 
perature of 207—35”, near Myvatn even at a temperature of about 40”. 
It occurs far more rarely on peaty soil, and then only scantily. Curi- 
ously enough, these habitats, also, are situated in districts where there 
are hot springs. Near Laugarland, for instance at the edge of the 
marsh, there is a small spring with lukewarm water, but the heat from 
it can exert no influence on Haplozia crenulata, nor does the latter 
occur in the immediate neighbourhood of the spring. 

This species varies considerably in colour, size, thickness of the 
cell-walls and as regards the leaf-margin. In the low, reddish-brown 
forms which grow on warm ground nearest to the hot water the cell- 
walls are more highly thickened, and are especially distinctly collen- 
chymatous, while the more vigorous, green forms which grow at some 
distance from the spring on more boggy ground, among other Bryo- 
phyta, have thin-walled cells which are indistinctly collenchymatous. 
The marginal cells are sometimes large and thick-walled, sometimes 
scarcely larger than the other leaf-cells and then only slightly thickened ; 
in such cases it may be difficult to distinguish the plant from the forms 
of Haplozia sphærocarpa and Alicularia scalaris with which it is often 
found associated. Under high magnifying powers the marginal cells 
will however always be seen to be somewhat papillose, which is never 
the case in Alicularia. 


28. Haplozia sphærocarpa (Hook.) Dum. 


S. Iceland: Grafarbakki near a hot spring (F.)!; Thorlåkshver among 
Catharinea undulata!; Sydri Reykjahver among Oligotrichum hercynicum !; 
Laugarvatnshver among Sphagnum cymbifolium!; Isafjordur on a rocky 
flat (300—400 metres above sea-level)!. 

All the forms found, which are quite sterile, stand slender and 
erect among other Bryophyta and must doubtless be referred more 
particularly to the type. 2 

This species, like its companion Oligotrichum, has a very peculiar 
distribution in Iceland, having two such widely different areas of distri- 
bution as the warm clayey flat with a temperature of 257—30” and 
the rocky flat. 

Note. A liverwort is figured in Flora Danica, tab. 2195, under the 
name Jungermannia pumila; it is recorded to have been found by 
Morch near Lejrå. Lindberg, in his critical revision of the mosses 
in Flora Danica, refers it to Jungermannia cæspiticia, but the specimens 
are not to be found in the collections. 


29. Haplozia cordifolia (Hook.) Dum. 


Very common and often fruiting richly. It grows especially on 
rocks, in or by waterfalls; or submerged in rivers, where it sometimes 
covers large surfaces of the firm rocky bottom with its blackish-green 
mats. But it may also be met with on irrigated gravelly ground or in 


THE BRYOPHYTA OF ICELAND 411 


moss bogs associated with species such as Philonotis seriata, Chiloscyphus 
polyanthos v. fragilis and Scapania undulata. 

It is most widely distributed in the lowlands up to a height of 
about 300 metres, but is also frequently met with up to about 500 
metres, for instance in Esja. 


30. Haplozia riparia (Tayl.) Dum. 


Vestmannaey. S. Iceland: Flokastadagil!; Klitnafoss!; Barkarstadr'!; 
Holt!. 

In all the above-mentioned localities the plant was growing on tuff 
and was, as a rule, fruiting. On Vestmannaey it was growing on "Stora 
Klit,” on the surface of dripping tuff-rocks facing north, associated with 
Hymenostylium curvirostre, Hypnum filicinum and Anomobryum filiforme. 
In S. Iceland it is fairly common in damp ravines on Fljotshlid and 
below Eyafjall, and grows there partly associated with the above-men- 
tioned species, partly in company with Fegatella conica, Preissia commu- 
tata, Bryum oenum, etc. 

Haplozia riparia is recorded by Gronlund from Brynjudalur and 
Seydisfjordur, but the specimens from these localities must he referred 
to Haplozia atrovirens. 


31. Haplozia atrovirens (Schl.) Dum. 


Common on damp rocks, more rare on damp gravelly ground. 

It occurs sometimes in low, blackish-green cushions, sometimes 
intermixed in the tufts of other Bryophyta. Male plants and fruiting 
plants are frequently met with, sometimes in separate tufts, sometimes 
mixed in the same tuft. Like Haplozia cordifolia it occurs most com- 
monly to a height of about 300 metres, but is also frequently met with 
up to about 500 metres. Var. sphærocarpoidea (De Not.) Mass. has a distri- 
bution similar to that of the type, and occurs in the same localities as 
the latter. Both forms pass evenly into each other. 

The cuticle of the leaves, in the majority of the plants, is more 
or less distinctly rough with striæ, and rarely smooth. 


32. Haplozia pumila (With.) Dum. 


N. Iceland: Hof, near Eyjafjordur (0. D.)!. It was growing there 
intermixed in a tuft of Eucalyx subellipticus. 

The rest of the specimens referred by Gronlund and Jensen to 
this species all belong to Haplozia atrovirens. 


33. Jamesonielia autumnalis (De Cand.) Steph. 
S. Iceland: Nuphlidarhåls (Stp.)!. 
34. Sphenolobus minutus (Crantz) Steph. 


SE. Iceland: Lon, scantily among Sphagnum rubellum and Fissidens 
osmundoides. NW. Iceland: Hesteyri, in a tuft of Dicranum elongatum. 
W. Iceland: Grund in Skorradalur, among Polytrichum strictum. 


PÅ (cz 


412 A. HESSELBO 


This plant was first found by Morch and figured in Flora Danica, 
tab. 2190, without a more precise notification of its habitat. 

Grénlund's record of the occurrence of this species in Esja is 
due to an erroneous determination. 


35. Sphenolobus saxicola (Schrad.) Steph. 


fin Islandia leg Morch” (M. B. H.)!.. Figured in Flora Danica, tab. 
2693, fig. 1, but no habitat is given. 


36. Sphenolobus politus (Nees) Steph. 


Isafjordur! c. coles. 

It was growing there in several localities at an altitude of about 200 
—300 metres, sometimes along the banks of streams associated with Har- 
panthus Flotowianus, sometimes in bogs associated with Sphagnum teres, 
Lophozia quinquedentata and L. Kunzeana, 


37.Lophozia quinquedentata (Huds.) Cogniaux. 


Very common all over Iceland. 

Lophozia quinquedentata is doubtless the liverwort of most fre- 
quent occurrence. It is met with, up to the snow-line, in all possible 
localities which are somewhat damp, now and then in unmixed tufts, 
but generally woven into the tufts of mosses. Female plants and fruit 
are rather rare, male plants are far more frequent; plants bearing 
gemmæ are also often met with. 

It varies extremely as regards size, habit and leaf-form. The extreme 
members of the variation-series are the large var. turgida Lindb. which 
is common on boggy ground and t e delicate, often only one mm. 
broad, var. fenera C. Jens. which is frequent in dry localities, as for in- 
stance on humus-covered rocks and in lava-fields. 


38. Lophozia lycopodioides (Wallr.) Cogniaux. 


Almost as common as the preceding species on drier ground. It 
is found especially on stony or grass-covered slopes, creeping over 
humus-covered rocks, in birch coppices, and on the top of knolls in 
bogs, sometimes in large, unmixed, yellowish-green tufts, sometimes inter- 
mixed with other Bryophyta. Only sterile specimens have been found. 
It varies considerably in size. Delicate forms (var. parvifolia), often 
only one mm. broad, grow on dry rocks woven into the tufts of other 
Bryophyta, while the large, vigorous forms are especially met with on 
slopes overgrown with Hylocomium. Now and then forms also occur 
which approach very closely to Lophozia Hatscheri (Evans) Steph. But 
plants which can with certainty be referred to this species have not 
been found in Iceland. 

Lophozia lycopodioides is most widely distributed in the birch-zone, 
up to about 400 metres above sea-level, and is especially wide-spread 
towards the north-west while it occurs more sparingly in S. Iceland. 


THE BRYOPHYTA OF ICELAND ALS 


In the district around Isafjérdur it is also met with abundantly on the 
rocky flat, 500—600 metres above sea-level, and near Akureyri it was 
collected, although sparingly, near the snow-line (at about 770 metres alt.). 


39. Lophozia Floerckei (W. et M.) Schiffner. 


NW. Iceland: Dyrafjérdur; Isafjéordur; Hesteyri!; Grunnavik!; Kal- 
dalon!; Arngerdareyri!. SW. Iceland: Svinahraun!. 

In the majority of the older lists of Bryophyta from Iceland L. 
Floerckei is recorded from rather a considerable number of habitats, 
but all the specimens from these habitats preserved in the Botanical 
Museum in Copenhagen, have been wrongly determined; the majority 
of them must be referred to Lophozia lycopodioides. The only specimen 
in the collections, correctly determined, was gathered by Mårch, but 
no habitat is given. 

This species has a very peculiar distribution as it is very common 
in the north-west, and is also found abundantly on the highly situated 
Svinahraun in the south-west (about 300 metres), but has not otherwise 
been found in Iceland. In NW. Iceland it ascends from the low land 
to the rocky flat, but is most frequently met with from 200 to 400 
metres above sea-level. There it grows on fairly dry ground in bil- 
berry heaths, both on the ground and creeping over rocky blocks, as- 
sociated with Dicranum fuscescens, D. scoparium, D. molle, Lophozia lyco- 
podioides and L. quinquedentata. At a higher level it grows especially 
on the Salix-herbacea-flats, associated with Lophozia ventriosa, L. alpestris, 
Dicranum Starkei, Pleuroclada albescens, etc., but also occurs on a more 
damp bottom. 

In Svinahraun it occurred commonly between the lava-blocks and 
also in a damp depression at the edge of the lava-field, associated with 
Polytrichum commune. 


40. Lophozia quadriloba (Lindb.) Evans. 


E. Iceland: Berufjérdur!; Lon!; common near Seydisfjordur; Val- 
lanes!. N. Iceland: Husavik!; Akureyri!; about Eyjafjordur (O.D.;!). NW. 
Iceland: Kaldalon!; Isafjérdur!. W. Iceland: Melar (Grl.)!; Esja, several 
places!. S. Iceland: Svinahraun!; Holt!; Hornafjérdur!. The Vestmanna- 
eyjar!. 

This plant is no doubt common, but usually it occurs so scantily 
that it is only found by close search of the Bryophyte collections under 
the microscope. It grows both on damp humus-covered rocks and on 
boggy ground, from the low land upwards to the mountain-tops. Thus, 
near Isafjordur it was collected at about 440 metres. Near Seydisfjordur 
it is frequent up to about 500 metres and near Akureyri it was found 
on cliffs at an altitude of 900 metres. 

Lophozia quadriloba hardly occurs in unmixed tufts, but always 
mixed with other species, for instance with Dicranum spp. or on rocks 
among Distichium montanum, Tortella fragilis, Oncophorus, or with other 
Hepaticæ such as Cephalozia, Lophozia alpestris and Aneura pinguis. On 


414 A. HESSELBO 


boggy ground it is found interwoven with Hypnaceæ in their tufts, but 
rarely in those of Cinclidium, Lophozia Kunzeana or other bog-mosses. 

The majority of the specimens investigated are small and delicate, 
and must partly be referred to forma heterophylla Bryhn et Kaalaas, and 
partly to forms transitional between this and the type. Only sterile 
plants have been found. 


41. Lophozia Kunzeana (Hub.) Evans. 


Very common in bogs, on damp rocks, damp gravelly ground, etc. 
It is most widely distributed in bogs where it grows woven into, 
or creeping over tufts of other Bryophyta, especially Sphagnum. Near 
Akureyri, for instance, it commonly occurs in bogs up to a height of 
about 600 metres. In S. Iceland it is rather rare in the bogs of the 
lowlands, but is met with abundantly at an altitude of 200—400 metres. 


42. Lophozia barbata (Schmid.) Dum. 


E. Iceland: Årbær, among Grimmia canescens (H.J.)!.. W. Iceland: 
Bjarnarhåfn (H. J.)!;' Berserkjahraun (H. J.)!; Reynivellir (Grl.)!; Nordre- 
dalur in the district of Borgarfjordur!; Ålafoss!. 

In addition to the above-mentioned localities Gronlund and Helgi 
Jønsson record this species from several other localities, but some of 
ihe specimens which have been investigated proved to be wrongly deter- 
mined (being, as a rule, forms of L. lycopodioides), so only those loca- 
lities are enumerated here from which authentic specimens are known. 

L. barbata appears to be rather rare. It has as a rule been col- 
lected, intermixed rather sparingly in tufts of other Bryophyta especially 
Grimmia and Hylocomium spp. and belongs to the Grimmia-heaths or 
to the mossy bottoms of birch coppices; thus in Nordredal it frequently 
occurred in birch coppices among Hylocomium spp., Dicranum scoparium, 
Ptilidium ciliare, etc. 

Note. Lophozia gracilis (Schl.) Stephani is recorded from several 
localities, but all the specimens which have been investigated have been 
wrongly determined, being mostly Lophozia lycopodioides var. parvifolia. 


43. Lophozia ventricosa (Dicks.) Dum. 


Common on more or less damp ground, both boggy and gravelly 
ground, and on rocks on mountain heights. 

This species is very common especially in NW. Iceland, and occurs 
there on a heathy bottom woven into tufts of Dicranum; along the 
banks of streams associated with Harpanthus, Lophozia quinquedentata 
and Cephalozia bicuspidata; and in bogs among Spagnum. It appears, 
on the whole, to be more frequent the higher it ascends; undoubtedly 
its main distribution is from a height of about 300—400 metres upwards. 

Var. porphyroleuca (Nees) Hartm. is likewise commonly distributed 
and occurs, for instance, frequently interwoven with Conostomum in its 
compact tufts, and also in tufts of Dicranum and Sphagnum on damp ground. 


THE BRYOPHYTA OF ICELAND 415 


It is hardly possible to draw any distinct boundary line between 
this form and the type. the thickenings of the leaf-cells showing all 
transitional stages from quite thin-walled to decidedly collenchymatous 
cells, and the colour also varying according to the habitat. Fruit is 
rare; gemmæ are as a rule present. 

Var. confertifolia (Schiffn.) syn. Lophozia confertifolia Schiffner. Plants 
agreeing exactly with the description of this form and with the figure 
in Rabenhorst's Kryptogamenflora, fig. 314, occur rather commonly 
on damp gravelly soil in NW. Iceland associated with all the transitional 
forms of L. ventricosa; I think they can only be regarded as Alpine 
forms of the present species. 


44. Lophozia Wenzelii (Nees) Steph. 


E. Iceland: Seydisfjérdur!; Vallanes!. N. Iceland: Tverå in Oxnadalur!. 
NW. Iceland: Isafjérdur, (at about 200 metres alt.)!. SW. Iceland: Svi- 
naskard (at about 400 metres alt.). 

It grows on wet gravelly soil or peat, especially along streams 
where it forms low, dense carpets or grows mixed with Alicularia sca- 
laris, Dicranella crispa, Distichium inclinatum and other Bryophyta. Near 
Isafjordur it was growing by a small stream together with Sphagnum 
Squarrosum. 


45. Lophozia alpestris (Schleich.) Evans. 


Very common all over Iceland. 

It generally grows woven into tufts of other Bryophyta, but now 
and then forms tufts by itself. It is found on almost every kind of 
substratum, from the sea-level to mountain heights. It occurs most fre- 
quently on rocks, especially those which are somewhat damp, but is 
also met with on damp ground along the bank of streams, on the gra- 
velly flats of mountain heights, in lava-clefts and woven into the moss- 
carpets of Grimmia-heaths. 

Fruit and calyces occur rarely and have been found only in NW. 
Iceland: Gnupsdal! and in S. Iceland: Seljaland (at about 350 metres)!; 
Svinahraun!. Male plants are rather common, and gemmæ are very 
often present. 


46. Lophozia excisa (Dicks.) Dum. 


S: Tceland: Reykir!; fr. 
It grows there on damp gravelly soil by a small stream associated 
” with Eucalyx subellipticus, Scapania curta and Dicranella crispa. 

This species is recorded from Iceland by Moårch, but of the two 
specimens found in the collection in the Botanical Museum in Copen- 
hagen under this name, one is Lophozia ventricosa var. porphyroleuca and 
the other L. heterocolpos. 

Note. Lophozia bicrenata (Schmid.) Dum. is recorded to have been 
found near Krisuvik by Steenstrup, but the packet contained only a 
mixture of several other species of Lophozia. 


416 A. HESSELBO 


47. Lophozia Schultzii (Nees) Schiffner. 


E. Iceland: Seydisfjordur; Vestdalur (at an altitude of about 200 
metres)!.  N. Iceland: Akureyri!. W. Iceland: Stykkisholmur!. 

It was collected everywhere only in small quantities together with 
bog-mosses such as Hypnum revolvens, H. stellatum, Mnium. Seligeri, 
etc.; it was as:a rule sterile. Near Stykkisholm it was. collected in 
unmixed tufts with antheridia and young calyces. 


48. Lophozia Milleri (Nees) Dum. 


Common, but occurred almost always in small quantities inter- 
mixed in tufts of mosses. 

This species is most frequent in the lowlands up to about 300-400 
metres above sea-level, but it also occurs at great altitudes in the Alpine 
region, for instance, at about 600 metres on Berufjérdurskard. It is 
almost always sterile. 

Lophozia Miilleri occurs on very different substrata, most frequently 
on rocks, but it may also be found on damp ground along streams, or 
even in bogs. It varies exceedingly in size according to the degree of 
dampness of the habitat. Xerophilous forms from rocks are usually 
small and delicate, while bog forms which grow in the tufts of Hypnaceæ 
are considerably larger and often approach so closely to L. Hornschu- 
chiana that they are difficult to distinguish from the latter. On the 
damp, vertical sides of the clefts of the tuff rocks of S. Iceland, for 
instance on Fljotshlid, a vigorous, blackish-green form is frequently 
met with forming large, pure mats. 


49. Lophozia Hornschuchiana (Nees) Macoun. 
Jungermannia bantryensis, Hooker. 


E. Iceland: Seydisfjordur, frequent up to about 400 metres!. N. Ice- 
land: Akureyri!; Hof; Eyjafjordur (0. D.)! cc. coles.  NW. Iceland: Isa- 
fjordur. OW. Iceland: Dalsmynne in Nordredal!; Mulakot!; Botusdalur!; 
Reykjavik!; Kollafjérdur. 

Occurs rather frequently, but as a rule scantily, on boggy ground 
among mosses (Hypnum, Sphagnum, etc.), more rarely on stones in rivers. 
In Nordredal, for instance, it was found abundantly in a small river 
on Submerged stones. 


50. Lophozia heterocolpos (Thed.) Howe. 


N. Iceland: Spénsgerde near Eyjafjordur (O.D.)!. NW. Iceland: Bæir!; 
Kaldalon!. SW. Iceland: Hafnarfjérdur!; Esja!, at an altitude of about 
400 metres; Thingvellir (Grl.;!). 

In all the above localities it was found on humus-covered rocks, 
especially in clefts. Near Kaldalon it grew on damp rocks woven into 
a tuft of Meesea trichoides and Dicranella subulata; on Esja it grew upon 
damp tuff-rocks. In Thingvallahraun it is widely distributed in clefts 


THE BRYOPHYTA OF ICELAND 417 


and caves in the lava-fields, usually mixed with Blepharostoma tricho- 
phyllum, Amblystegium Sprucei, Diplophyllum albicans and Plagiothecium 
denticulatum. The characteristic gemmiferous shoots occur always, but 
plants in fruit have not been collected. 


51. Gymnocolea inflata (Huds.) Dum. 


In Islandia” leg. Morch! and figured in Flora Danica, tab. 1945, 
fig. 2; SW. Iceland: Krisuvik (Stp.)!; Laugarvatn (Stp.!); Reykholt (Grl.;!); 
Reykirdalur!; Kolvidarhol!. NW. Iceland: Laugarland '!. 

Gronlund records this species from several other localities be- 
sides those mentioned above, but the specimens in the Botanical Museum 
in Copenhagen are all wrongly determined and must be referred to 
Lophozia Miilleri or to L. alpestris. 

In Iceland Gymnocolea inflata is a decidedly warm-soil plant which, 
with the exception of a single locality, was found exclusively on the 
steaming clayey flats around hot springs in SW. Iceland. Near Laugar- 
land it was growing rather sparingly in a bog. Haplozia crenulata was 
growing in its neighbourhood, and as there was a small spring with 
lukewarm water at the edge of the bog it is possible that the occur- 
rence of the two species is connected with the spring, although there 
was no characteristic vegetation around the spring, but only the com- 
mon bog-plants. 

In Reykirdalur, where it is found in abundance near almost all 
the hot springs, it forms extensive brownish-black or almost entirely 
black mats on warm ground with a temperature of as much as 40” 
around fumaroles or on slopes stretching down towards the boiling hot 
basins. At some distance from the spring it will grow woven into the 
tufts of Polytrichum commune or mixed with Cephalozia bicuspidata which 
likewise occurs in brownish-black forms. 

It was collected with calyces only at Laugarvatnshver. 


52. Plagiochila asplenioides (L.) Dum. 


Very common everywhere. 

Plagiochila asplenioides occurs on almost every substratum, on damp 
or on dry rocks, in bogs, on earth, etc. It is most frequent up to a 
height of about 300 metres above sea-level, but often occurs as far up- 
wards as about 600—700 metres. Usually it grows intermixed in the 
tufts of other Bryophyta, more rarely it forms tufts by itself. The 
majority of the specimens which have been collected are small, only 
1—2 mm. broad, with leaves slightly dentate or entire. Larger forms 
with leaves typically dentate occur mostly on somewhat damp ground, 
for instance among stones on a talus of loose blocks and debris (Urd) 
or in caves. Only sterile specimens have been found. 


53. Leptoscyphus anomalus (Hook.) Lindb. 
E. Iceland: Ståd (H. J.)!. 


418 A. HESSELBO 


54. Lophocolea cuspidata Limpr. 


Vestmannaey!. S. Iceland: Holt!. 

On Vestmannaey it was found in several places at the base of cliffs 
and in specially large quantities in Heljusdal where it grew abundantly 
among stones on an Urd usually mixed with Eurynchium, Stockesii 
and Mnium undulatum. Near Holt it was found in several places, but 
everywhere only in small quantities in clefts of tuff-rocks, woven into 
the tufts of mosses. Only sterile specimens have been found. 


55. Lophocolea minor Nees. 


N. Iceland: Hålsskågur (O.D.)!. S. Iceland: Reykirdalur!; Vestman- 
naey: Heljusdal!. , 

In Reykirdalur it was growing rather sparingly on a grass-covered 
slope stretching down towards a stream. On Vestmannaey it occurred 
in several places associated with L. cuspidata sometimes intermixed in 
its tufts. In all the localities only sterile specimens were collected, but 
bearing the characteristic gemmæ. 


56. Chiloscyphus polyanthus (L.) Corda. 


Common in wet localities or in water, on gravelly ground and 
occasionally in bogs and on wet rocks. Only sterile specimens were 
found. 

It occurs sometimes in unmixed cushions, which are usually blackish- 
green in colour in water and yellowish-green in boggy ground, some- 
times mixed with other Bryophyta such as Marchantia, Philonotis, Dicra- 
nella squarrosa and Mnium spp. It is most widely distributed in the 
lowlands up to about 300 metres and does not appear to ascend much 
higher than 400 metres. 

The typical form was collected only in S. Iceland on damp tuff- 
rocks near Holt. All the other plants which have been investigated be- 
long to var. fragilis (Roth) K. M. It varies considerably as regards 
size and colour, but the form of the leaves and the size of the leaf- 
cells are almost always the same. 

Note. Chiloscyphus pallescens is recorded from several places by 
Moårch, Gronlund and Helgi Jonsson, but all the specimens refer- 
red to this species belong to Chiloscyphus polyanthos var. fragilis. 


57. Harpanthus Flotowianus Nees. 


E. Iceland: Seydisfjordur, several places!. N. Iceland: several places 
around Eyjafjordur (O. D.)!; Stadr near Hrutafjérdur!. NW. Iceland: Dyra- 
fjordur; Sugandafjérdur; Isafjordur; Grunnavik; Kaldalon; Arngerdareyri !. 
W. Iceland: Olafsdalur in Dalasysla (H. J.)!; Esja, several places!. 

In NW. Iceland this species is of the most frequent occurrence 
among all the Bryophyta and is met with everywhere on wet ground 
up to a height of about 300 metres above sea-level. In the other parts 
of the country it is wanting or rare; on Esja it is however rather fre- 


THE BRYOPHYTA OF ICELAND 419 


quent. It occurs both in bogs and along the banks of streams, and 
also on inundated gravelly ground, usually intermixed in tufts of mosses 
such as Cinclidium, Mnium cinclidioides, Sphagnum and Hypnum spp.; 
also on wet gravelly soil associated with Lophozia quinquedentata, Ce- 
phalozia bicuspidata and other Hepaticæ. Only sterile specimens have 
been found. . 


Fam. TRIGONANTHEÆ. 


58. Cephalozia bicuspidata (L.) Dum. 


Very common on earth, damp gravelly soil, humus-covered rocks, 
and in bogs up to a height of about 600 metres above sea-level, espe- 
cially in NW., N. and in parts also of E. Iceland, while it appears to 
be more rare in S$S. Iceland. It grows partly in large cushions, for 
instance on damp ground by streams, partly intermixed in tufts of 
møosses especially Sphagnum and Dicranum spp. and thrives best on a 
somewhat damp substratum. Found as a rule in fruit. It varies con- 
siderably as regards size and colour. 

Var. Lammersiana (Hub.) Breidler grows on very wet ground, for 
instance along banks of streams, where it forms large green mats in 
association with other Hepaticæ. It is especially common in NW. Ice- 
land. The bog-forms, which grow erect among Sphagnum, are slender 
with spreading leaves. On damp slopes a dark-brown-to-black form 
frequently occurs, often forming very extensive carpets; it corresponds 
most closely with var. Loeskeana (Schiffner) K. M., in previous lists it 
was referred to var. Lammersiana. At the head of deep lava-caves 
and at the bottom of lava clefts it occurs in association with Alicularia 
scalaris as much elongated shade-forms. 

Note. Cephalozia Francisci is recorded by Mårch from Hafnar- 
fjordur, but the specimen in the Botanical Museum is C. bicuspidata. 


59. Cephalozia ambigua C. Mass. 


NW. Iceland: Laugarland, fr.!. 
It grew there in small brownish tufts on the damp ground at the 
bottom of the valley. 


60. Cephalozia pleniceps (Aust.) Lindb. 


Common in all parts of Iceland except in the southern part, where 
it appears to be rare. It is especially common in NW. Iceland and 
forms there one of the most frequently occurring Hepaticæ. 

It occurs especially on damp ground and in bogs, woven into tufts 
of Sphagnum, Dicranum and Mnium hornum or associated with Lophozia 
Kunzeana, L. quinquedentata and other Hepaticæ and as a rule bears 
calyces or fruit. It occurs most frequently up to a height of about 
300 metres, but it is also often met with at far greater altitudes. 

Var. macrantha (Kaalaas et Nichols.) K. M. is widely distributed on 
wet ground, and grows in association with similarly elongated forms of 


420 A. HESSELBO 


C. bicuspidata and Cephaloziella Hampeana, woven into the tufts of 
Sphagnum spp., but is as a rule sterile. 

Note. Cephalozia connivens is recorded by Moårch and Grånlund 
as found in Iceland, but all the specimens which have been investigated 
belong to C. pleniceps. 

W 


61. Cephalozia media Lindb. 


N. Iceland: Arnarfellsaurar (St.;!); NW. Iceland: Grunnavik!, inter- 
mixed in tufts of Dicranum fuscescens. 


62. Pleuroclada albescens (Hook.) Spruce. 


E. Iceland: Seydisfjérdur!, common above a height of about 400 
metres  N. Iceland: Akureyri!, common above 500 metres; Hestahraun 
(St.)!.… NW. Iceland: Very common above 300 metres!. Var. islandica 
(Nees) Spruce. N. Iceland: Reykjaheidi!. S. Iceland: Thingvallahraun 
(MorebsFSIps Gr EN Ham arfjordur! 

The typical form is no doubt common on mountain heights, at 
any rate in N., NW., and E. Iceland. In S. Iceland it has not yet been 
found. On mountain heights it grows on stony flats associated with 
Dicranum spp. (D. Starkei, D. molle and D. fuscescens), Lophozia spp. 
and Cesia concinnata or associated with Polytrichum sexangulare, Lophozia 
ventricosa, Cesia concinnata, etc., on Salix-herbacea-flats. 

The variety belongs especially to the lava-fields and grows there 
at the bottom of deep clefts, creeping over blocks of lava, often in 
abundance. It occurs there either in large whitish-green cushions or 
mixed with Hylocomium spp., Rhacomitrium hypnoides, Polytrichum alpinum, 
P. sexangulare, Lophozia spp., etc. Only sterile specimens of each form 
have been found. 


63. Cephaloziella Hampeana (Nees) Schiffn. 


Very common, but occurs only as scattered shoots intermixed in 
the tufts of mosses, sometimes, however, also in tiny, yellowish-green, 
brownish or reddish tufts upon these. 

It occurs on highly different substrata. On rocks and on earth it 
grows in tufts of Dicranum spp., Ditrichum flexicaule, Distichium montanum, 
Tortella fragilis, etc.; in bogs it has been found woven into tufts, of 
Sphagnum, Sphaerocephalus palustris, Hypnaceæ, Oncophorus spp. and 
several others. Bog-forms, with elongated stems and widely spreading, 
very much projecting leaves, agreeing most closely with var. erosa Warnst., 
occur very commonly in Sphagnum cushions. 

The leaf-tissue varies very considerably, the leaf-cells, even on the 
same plant, being sometimes quite thin-walled, sometimes more or less 
thickened; the leaves also are more or less outspread, so that all pos- 
sible transitional forms between this species and Cephaloziella rubella 
are known. 


THE BRYOPHYTA OF ICELAND 421 


64. Cephaloziella rubella (Nees.) Warnst. 


Typical specimens of this species were collected in the following 
localities: 

S. Iceland: Hornafjérdur!. E. Iceland: Seydisfjordur!. N. Iceland: 
Hafrardalur (St.)!; Akureyri!, at an altitude of about 500 metres!; Hof 
(O. D.!). SW. Iceland: Kollafjérdur!. 'S. Iceland: Holt!. 

It was found everywhere woven into tufts of Sphagnum spp. and 
Dicranum spp. (D. fuscescens and D. anguslum) and usually mixed with 
Cephalozia bicuspidala and Lophozia spp., now and then also with C. 
Hampeana. Near Isafjordur it grew sparingly in a tuft of Mnium hornum, 
which was densely interwoven with Cephalozia pleniceps. It was col- 
- lected everywhere in fruit. 

C. rubella is an uncertain species which when sterile is in many 
cases hardly distinguishable from C. Hampeana. 

C. divaricata and C. bifida are recorded from several places in Icc- 
land, but the specimens belong either to C. Hampeana or to C. rubella. 
Groénlund's C. divaricata from Melar is Lophozia quadriloba. 


65. Odontoschisma Sphagni (Dicks.) Dum. 


W. Iceland: Snældubeinstadahver in Reykholtidalur!. 

lt was found in the above locality in small quantity woven into a 
tuft of Sphagnum rubellum on a warm substratum. 

This species is recorded from Iceland in several old lists and is 
figured in Flora Danica, tab. 2251 (leg. Mårch), but all the specimens 
found in the collections under this name are those of O. elongatum. 


66 Odontoschisma elongatum (Lindb.) Evans. 


In Islandia” (Mérch!). E. Iceland: Hof!; Lon!; frequent near Sey- 
disfjéordur!. N. Iceland: Hvarf (St.)!; Akureyri!. W. lceland: Budahraun 
(H.J.)!; Grund in Skorradalur!; common around Reykjavik!. S. Iceland: 
Holt! at an altitude of about 400 metres. 

It grows usually woven into tufts of bog mosses such as Sphagnum, 
Hypnum and Oncophorus spp., Meesia trichoides, Lophozia Kunzeana, 
more rarely upon moss-tufts or on wet ground, in small brownish tufts. 
Only sterile specimens have been found. 

A specimen collected by Mårch, having tiny, inconspicuous under- 
leaves and less thickened cell-walls bearing fine warts, forms a transi- 
tional form to O. Sphagni. 


67. Odontoschisma denudatum (Mart.) Dum. 


W. Iceland: Braudarholt near Reykjavik! dg. 
It grew in the above locality on boggy ground among other Bryo- 
phyta. 


422 A. HESSELBO 


68. Odontoschisma Macouni (ÅAust.) Underwood. 


S. Iceland: Bergarfoss near Hornafjérdur!; E. Iceland: Grottafoss 
near Seydisfjordur! at an altitude of about 200 metres. N. Iceland: Hof 
(O.D.)!. SW. Iceland: Svinahraun!. 

This plant grows partly in small unmixed tufts, partly mixed with 
other Bryophyta such as Anoectangium lapponicum, Distichium montanum, 
Pohlia cruda, Blepharostoma trichophyllum on damp humus-covered rocks. 
In Svinahraun it grew in lava-clefts filled with humus. 


69. Calypogeia Trichomanis (L.) Corda. 


N. Iceland: near Myvatn (Grl.”); Reykjahlid!. W. Iceland: near se- 
veral hot springs in. Reykholtdalur!; Tunguhver (probably Deildatungu- 
hver) (Grl.;”); Kollafjérdur!. S. Iceland: Laugaråshver!; Geysir!.  More- 
over, Grénlund records this species from Hvammur, but no specimens 
of it are to be found in the collections. 

The plant grows in all the above localities on a warm substratum. 
Near Reykjahlid it covered the roof of a lava-cave which had a tem- 
perature of about 25%. In the other localities it occurred on a warm 
damp substratum woven into mosses such as Sphagnum, Hypna and 
Polytrichum, or in pale green to brownish cushions above them. In 
Reykholtdalur where it occurred abundantly in several localities, the 
temperature in the tufts was from 25” to 27". 


70. Lepidozia setacea (Web.) Mitten. 


S. Iceland: Merkjåfoss (Feddersen) !. 
It was found very sparingly in the above locality associated with 
Cephalozia pleniceps and woven into a tuft of Dicranum scoparium. 


FAM PET DIOIDEÆ 


71. Blepharostoma trichophyllum (L.) Dum. 


Very common from the lowlands up to about 500—600 metres 
above sea-level. 

This species rarely forms unmixed tufts, but grows almost always 
woven into tufts of other species and is met with in the most varied 
localities, on boggy ground and damp gravelly ground along rivers, in 
lava-fields, on damp rocks, and especially in abundance in rock-caves 
where it often covers the roof and walls in association with Ambly- 
stegium Sprucei, with a thin dark-green layer. 

Fruit occurs rather rarely; but it is found plentifully in Alman- 
nagjå!, and near Hof in N. Iceland (0. D.)!. 


72. Chandonanthus setiformis (Ehrh.) Lindb. 


In the herbarium of the Botanical Museum there is a specimen, 
labelled by Mérch "In Islandia, among Trichostomum canescens, Aug. 1820.” 


THE BRYOPHYTA OF ICELAND 423 


73. Anthelia julacea (L.) Dum. 


Common in the lowlands on damp gravelly ground and creeping 
over wet stones in streams, occasionally also on damp boggy soil; fruit 
is not rare. It does not appear to ascend higher than about 400 me- 
tres above sea-level. 


74. Anthelia Juratzkana (Limpr.) Trevis. 
AÅnthelia nivalis (Sw.) Limpr. ex. p. 


Very common on damp gravelly flats irrigated by melting snow, 
somewhat rarer on damp gravelly ground along streams or on damp 
rocks. 

This plant has its main distribution on mountain heights where, 
on the clayey or gravelly flats next to the snow-covered areas, it forms 
large, continuous carpets of a peculiar bluish-black colour. often mixed 
with Salix herbacea or with other Bryophyta such as Alicularia geo- 
scypha, Lophozia ventricosa and Polytrichum sexangulare. In NW. Ice- 
land, where the snow on slopes with a northern exposure may remain 
during the greater part of the summer, these Anthelieta often descend 
to sea-level. The plants growing on. damp rocks or gravelly ground in 
lower levels differ from the typical form from mountain heights in being 
lighter in colour, usually yellowish green, and in the tufts being as a 
rule higher, with less close-set leaves; or it may also grow intermixed 
with other Bryophyta especially Hepaticæ. 

A. Juratzkana and ÅA. nivalis are so closely related that in many 
cases it is impossible to determine them when sterile. While the typi- 
cal AÅ. Juratzkana, as it occurs in abundance on mountain heights, is 
very characteristic because of its extensive low carpets, quite typical 
specimens of Å. julacea are far more rarely met with on the damp soil 
of the lowlands. Of far more frequent occurrence are forms which 
pass, more or less, in habit, size of leaf-cells and thickness of cell-walls 
from A. julacea towards Å. Juratzkana. 


75. Ptilidium ciliare (L.) Hampe. 


Very common both on dry and on somewhat damp ground. 

The plant grows both among Hylocomium spp. on slopes and inter- 
mixed in heaths of Rhacomitrium hypnoides and R. canescens on more 
or less damp rocks coated with soil, and also in clefts in lava-fields 
and on knolls in bogs. It occurs most frequently in low-lying regions 
up to about 300—400 metres, but may also occasionally be met with 
on mountain heights. Only sterile specimens have been found. 


Fam, SCAPANIOIDEÆ. 


76. Diplophyllum albicans (L.) Dum. 


"In Islandia” (Morch!). W. Iceland: Budahraun (H.J.)!; Hafnarfjordur 
(Grl.; H. J.;); Svinahraun!; Modruvellir in Kjos!. S. Iceland: Thingvalla- 


424 A. HESSELBO 


hraun (Grl.; Stp.;!); Krisuvik (Stp.)!; Holt!; Reykirdalur!, at an altitude 
of about 260 metres. 

Diplophyllum albicans is a plant characteristic of the lava-fields of 
West and South Iceland and occurs there in great abundance on ver- 
tical rock-sides in clefts and caves. while it appears to be entirely ab- 
sent from the lava-fields of North Iceland. Outside the lava-fields it 
has been gathered only extremely sparingly among stones on heaps of 
debris at foot of cliffs (Urd). In Reykirdalur it grew scantily on a warm 
substratum among large blocks of basalt. Only sterile specimens have 
been found. 


77... Diplophyllum obtusifolium (Hook.) Dum. 


SIn Islandiae montibus leg. Méårch” and figured in Flora Danica, 
tab. 1831, fig. 2. No specimens of it are to be found in the herbarium 
of the Botanical Museum. In the district of Seydisfjordur!, on a damp 
rock-face by the river it occurred in a small quantity in a tuft of Di 
cranum molle and Lophozia Kunzeana. In N. Iceland: Åsbyrgi!, it was 
found on the ground among fallen blocks of lava. Only sterile specimens 
have been found. 


78. Scapania subalpina N. ab Es. 


Common on damp ground along streams, on damp rocks, in clefts 
and on damp slopes of the low land. It does not appear to ascend 
much higher than about 300 metres. 

& plants are very common; calyces and fruit occur occasionally. 

The plant varies in having deeply toothed to almost entire leaves. 
It is usually green or yellowish-green in colour, but sometimes reddish 
or brownish to almost blackish-brown in more exposed localities. 


79. Scapania remota Kaalaas. 


N. Iceland: Akureyri!. 

The plant, which agreed exactly with the description and figure 
in Karl Muller's Monographie der Lebermoosgattung Scapania, grew 
in the above locality, in a bog 400 metres above sea-level, intermixed 
in tufts of Oncophorus virens. 


80. Scapania irrigua (Nees) Dum. 


Very common on damp ground, in bogs, along streams and on 
damp rocks. 

This species is most widely distributed in the lowlands up to 
about 300 metres above sea-level, but may also frequently be met with 
at high altitudes in the Alpine regions. In wet localities, especially 
in bogs, where it occurs everywhere woven into the tufts of other 
Bryophyta, it is usually yellowish green in colour. The leaves are en- 
tire or fewly toothed towards the apex with thin-walled or slightly 
collenchymatous cells. But occasionally vigorous, brownish-green forms 


THE BRYOPHYTA OF ICELAND 425 


also occur with more highly thickened cell-walls. In somewhat drier 
localities it is usually brownish in colour and more compact with more 
deeply toothed leaf margins, sometimes almost as in $. subalpina and 
with more thick-walled cells. Small forms from damp sandy soil often 
constitute transitional stages to S$. curta, while vigorous aquatic forms 
approach closely to S$. undulata. Å very vigorous form with leaves 
which were as deeply toothed as in $. subalpina was found in abun- 
dance in lava-clefts near Thingvellir. 

cd" plants are common; calyces and fruit occur rather rarely. 

Note. Scapania compacta is recorded by Gronlund from Esja, 
but the specimen in the Botanical Museum is really Scapania irrigua. 


81. Scapania uliginosa (Sw.) Dum. 


E. Iceland: Stéd (H.J.)!.: NW. Iceland: Gnupsdal in Dyrafjordur!; 
Sugandafjérdur!; Isafjardarheidi!; DyYnjandi!; Arngerdareyri. 

With the exception of a single locality in E. Iceland this species 
was found only in NW. Iceland where it is rather common on very 
wet or irrigated soil along streams, often occurring in great abundance 
up to a height of about 400 metres. Near Arngerdareyri it grew abun- 
dantly in a bog in reddish-brown tufts associated with Hypnum spp. 
The leaf-form and habit in this form were typical, but the leaf-cells 
had brownish and rather thickened walls and a roughly granular 
cuticle. 

The plant from Sugandafjordur differed from the type in its leaves 
being sometimes distantly toothed and the cells very slightly thickened 
at the angles, in which respects it approached S$. irrigua. 


82. Scapania paludosa C. M. 


NW. Iceland: Gnupsdal in the district of Dyrafjordur, c. coles.! Grun- 
navik g!; Isafjordur g!. 

In Gnupsdal the plant grew abundantly in the water in small 
streams, where it formed large, loose, green to brownish mats. The 
leaves were highly decurrent, with indistinct teeth or entire margins, 
and thin-walled cells. 

Near Isafjordur and Grunnavik it grew on wet boggy ground. The 
leaves in both these forms were shortly decurrent and more or less 
toothed with a broadly reniform or (often in the same plant) cordate 
postical lobe which was occasionally furnished with a small point as in 
S. irrigua; the leaf-margin was often reflexed as in S$. undulata; the 
cells were thin-walled, but towards the margin occasionally somewhat 
thick-walled. The commissure is typically short and strongly curved 
with broad wings. 


83. Scapania dentata Dum. 
Scapania purpurascens (Hook.) Pearson. 


E. Iceland: Berufjérdur!; Seydisfjordur!. common up to about 400 
metres. Kirkjubél (H. J.)!. N. Iceland: Stora Gjå!. "W. Iceland: Hvammur 
The' Botany of Iceland. Vol. I, part II. 98 


426 A. HESSELBO 


(Grl.)!; Esja!, common up to about 400 metres above sea-level; Hafnar- 
fjordur!. S. Iceland: Krisuvik (Stp.)!; Krékr (H. J.)!; Almannagjå!; Holt!:; 
Seljaland!; Austerhlid near Geysir!. 

This species has its main distribution in E. and SW. Iceland and 
it has not been found in NW. Iceland. It grows especially on wet rocks 
and often in the water itself, but also on gravelly soil along the margin 
of streams. Thus, near Seydisfjordur it occurred abundantly along several 
rivers. In such localities it forms extensive reddish-brown cushions 
several centimetres deep. 

On damp soil and on rocks it forms low cushions, green, rose-red 
or reddish-brown in colour. In the lava-fields near Hafnarfjårdur and 
Thingvellir and in Stora Gjå it usually grew at the bottom of the com- 
paratively dry caves and clefts in small, flat tufts of a peculiar 
vellowish-green or beautiful rose-red colour. Such forms from com- 
paratively dry localities have always deeply-toothed leaves while the 
aquatic forms have less deeply-toothed, sometimes almost entire leaves, 
in which feature they approach $. undulata very closely. The cell-walls 
are however always more highly thickened than in the latter species, 
especially at the angles, and the postical lobe of the leaves are longer 
and narrower obovate (in $. undulata almost circular). &å plants and 
fruit are of frequent occurrence. 


84. Scapania undulata (L.) Dum. 


Very common on stones and gravelly soil in rivers, occasionally 
also on inundated ground in swamps and moss bogs. 

This plant occurs most frequently in the low land up to about 
300 metres, and forms there, especially creeping over stones in ihe 
shallow water along. rivers, extensive growths in association with Chilo- 
scyphus polyanthus var. fragilis and Haplozia cordifolia. Not rarely it is 
also met with at higher levels, for instance near Dyrafjérdur, where it 
grew abundantly on slopes irrigated by melting snow, 400—500 metres 
above sea-level; and near Holt below Eyjafjall, where it was found 
abundantly in a stream even at an altitude of about 600 metres... The 
aquatic forms are usually green; in. swamps anå moss bogs it often 
forms extensive, thick carpets, reddish-brown or almost purple in colour. 
It is frequently found in fruit. 

The leaves are entire or have a few obtuse teeth at the margin, 
but forms sometimes occur with more deeply toothed leaves or even 
with the leaf-margin entirely toothed, in the latter case it may be 
difficult to distinguish the plant from $. dentata and from the large 
aquatic forms of S$. irrigua. The latter, however, have always cells more 
thick-walled especially at the.angles, while S$. undulata has always thin- 
walled leaf-cells. 


835. Scapania curta (Mart.) Dum. 


Common on rocks coated with soil, damp sandy or gravelly ground 
and on peat, occasionally also on boggy ground up to about 400 metres 
above sea-level. Near Isafjordur and in Esja it was collected at an 


THE BRYOPHYTA OF ICELAND 427 


altitude of almost 500 metres. &g plants are common, fruit rather rare. 
The majority of the plants bear gemmæ. 
Var. geniculata (Mass.) C. M. i 

S. Iceland: Reykir!; Vestmannaey!. NW. Iceland: Armuli'!. 

Var. rosacea (Corda) Carr. is only exceptionally met with typically 
developed, while forms transitional between the present variety and the 
type are rather common. 

The plant is usually green or brownish-green, more rarely reddish. 
On damp ground it becomes larger and approaches $. irrigua. The 
leaves vary somewhat in form. The antical lobe is usually pointed, in 
the lower leaves it is, however, often rounded; it is more rare for. all 
the leaves to be rounded. The leaf-cells are as a rule thin-walled or 
somewhat thickened at the angles. Forms with highly thickened, red- 
dish-brown cell-walls are more rare, and occur always in exposed localities. 


86. Scapania Bartlingii (Hampe) Nees. 
Scapania Carestiae de Not. 


S. Iceland: Drångshlid associated with Amphidium Mougeottii and 
Bryum archangelicum (H. J.)!. 

Note. Scapania nemorosa is enumerated in the majority of the 
older lists, but all the specimens in the collections belong to Sca- 
pania undulata. 


Fam. RADULOIDEÆ. 
87. Radula complanata Dum. Gottsche. 


S.Iceland: Hornafjérdur!. E.Iceland: Hof!; Djupivogur!; Seydisfjordur !. 
NÆTeeland: Fhrastarholsårgsil fr" (OSDY)!N WS Iceland: Flatey (Grc)!- 
Budahraun (H. J.)!. Common in SW. and S. Iceland!. 

The plant is common in E., W. and S. Iceland, while it appears to 
be absent from NW. Iceland and to be rare in N. Iceland. 

It occurs only in the lowlands and rarely ascends higher than 
about 300 metres. In a few localities in Esja and near Kolvidarhol it 
was collected at an altitude of about 400 metres. It grows especially 
on dry, or only slightly damp, shaded rock-sides, mostly in clefts and 
caves, in company with Metzgeria furcata, Lejeunea cavifolia, etc., and 
often bears gemmæ and in S. Iceland fruit also. It was sometimes col- 
lected on the bark of trees, viz. on birch on Flatey and on mountain 
ash in Budarhaun. 


Fam. MADOTHECOIDE Æ. 
88. Madotheca Cordæana (Hibener) Dum. 
Madotheca rivularis (Dicks.) Nees. 


"In Islandia” (Morch)!. E.lIceland: Hof!; Seydisfjordur (H. J.)!; Beru- 
fjordur!. NW. Iceland: Dyrafjérdur! at an altitude of about 150 metres; 
28+ 


4928 A. HESSELBO 


Dynjandi!; Grunnavik! at an altitude of about 270 metres. W. Iceland: 
Nordredalur!;- Brynjudalur (Grl.)!; Reykjavik (Grl.;!); several places in 
Esja!. Common in S. Iceland!. Vestmannaey ! 

This plant is most widely distributed in S. Iceland where, as on 
Vestmannaey, it is common on more or less damp rock-sides especially 
in clefts of the tuff mountains on Fljétshlid and below Eyjafjall. On 
Vestmannaey it grew sometimes in abundance among fallen stones in 
Heljusdalur, and sometimes on slopes associated with Thuidium abie- 
tinum and Hylocomium spp. In E. and NW. Iceland it was rarer and 
generally occurred only sparingly on faces of basalt rocks immediately 
above the surface of the water of rivers or on damp slopes among 
stones. This species has not yet been recorded from N. Iceland. 


Fam. JUBULEÆ. 
89. Frullania dilatata Nees. 


W. Iceland: Flatey, on mountain ash (name of finder not stated); 
Dalasysla, Melar, on sea-fowl cliffs (H. J.)!. Only sterile specimens have 
been found. 


90. Frullania Tamarisci Nees. 


Common both on dry and on somewhat damp rocks in company 
with Hypnaceæ, Grimmia spp., Antitrichia curtipendula, etc.; in Rhaco- 
mitrium heaths (both R.canescens and R. hypnoides); on slopes among 
Hylocomium spp.; on the top of knolls in marshes, etc. 

I do not think the plant occurs at a much higher altitude than 
about 300 metres and it is always sterile. 


91. Frullania fragilifolia Tayl. 


SW. Iceland: Hafnarfjérdur, in a lava cave (Grl.)!; Kolvidarhol!; on 
a rock-face at an altitude of about 400 metres associated with Radula 
and Lejeunea. 


92. Lejeunea cavifolia (Ehrh.) Lindb. 
Lejeunea serpyllifolia (Dicks.) Spr. 


In Islandia” (Mérch)!. W. Iceland: Snæfellsnes!; Sandur in a lava 
cave (H.J.)!; Hafnarfjårdur in lava caves!; Kolvidarhol!. 'S. Iceland: 
Seljaland (Stp.)!; common on Fljåtshlid and below Eyjafjall!: Thingvalla- 
hraun!. 

This plant is common in S. Iceland where it grows especially on 
shady rock-sides in the clefts and caves of tuff mountains, associated 
with Metzgeria furcata, Madotheca Cordæana, Amphidium Mougeottii, etc. 
The habitat of greatest altitude is near Kolvidarhol at about 400 metres 
above sea-level, there it grew on a rock-face associated with Radula 
and Frullania fragilifolia. 


THE BRYOPHYTA OF ICELAND 429 


Besides in the tuff districts, it is also rather frequent in the lava- 
fields where it grows at the bottom of deep, damp clefts, but as a rule 
only in small quantities. Sterile specimens only have been found. 


Fam. ANTHOCEROTACEÆ. 
93. Anthoceros punctatus L. 


W. Iceland: Reykholtdalur (Ho.; Grl.;!). S. Iceland: Reykirdalur!; 
Sydri Reykjahver!. This species was first collected by Koenig and 
figured in Flora Danica, tab. 396. 

It occurs near the majority of the hot springs in Reykirdalur. It 
grows there abundantly on damp clay-flats with a temperature of 20” 
—30”, associated with Fossombronia, Haplozia crenulata and Archidium, 
especially on slopes immediately above the hot water. It almost always 
bears numerous capsules. 


430 A. HESSELBO 


IF SPHÅAGNAFES: 


1. Sphagnum imbricatum (Hornsch.) Russow. 
S. Austini Sull. 


S. Iceland: Thorlåkshver and Laugaråshver near Skålholt!. lt grew 
there on the warm substratum along the outlets from the hot springs 
in immense yellowish-brown cushions as much as 25 cm. deep. 


2. Sphagnum cymbifolium (Ehrh.) Hedw. 


W. Iceland: near several hot springs in Reykholtdalur!. S. Iceland: 
Thorlåkshver!; sydri Reykjahver!; Laugarvatnshver!; Hurdarbak (H. J.)). 

In all the above localities it grows on damp ground around hot 
springs. 


3. Sphagnum medium Limpr. 


NW. Iceland: Bæir!; Årmuli!. $. Iceland: Common in the district 
around Skålholt!. W. Iceland: Reykjavik! 
The plant grows everywhere in wet bogs. 


4. Sphagnum papillosum Lindb. 


W. Iceland: Reykjavik!, common on boggy ground; Reykholtdalur 
(Grl.;!), around numerous hot springs. $S. Iceland: Hraun; Grimstadr 
(H. J.)!: Hraundalshraun (H. J.)!; Sydri Reykjahver!. 

In Reykholtdalur Sphagnum papillosum constitutes the bulk of the 
vegetation around the hot springs and, for instance at Deildatunguhver, 
covers large areas of the warm, damp ground with cushions as much 
as about 20 cm. deep. It appears to be rather widely distributed in 
S. Iceland and is found there, as near Reykjavik, on the knolls in the 
bogs, sometimes on a rather dry substratum. 


5. Sphagnum inundatum Russow. 


W. Iceland: Grundarfjérdur (H. J.!). S. Iceland: Thorlåkshver!; Sydri 
Reykjahver'!. 

In the two latter localities it grew along the outlets from the hot 
springs, near Sydri Reykjahver it grew also on the knolls in a small 
bog near the spring. 


THE BRYOPHYTA OF ICELAND 431 


6. Sphagnum Gravetii Russow. 


NW. Iceland: Hesteyri!. 
It grew there in pools in company with a very large form of 
Hypnum revolvens v. Cossoni. 


7. Sphagnum compactum De Candolle. 


S. rigidum Schimp. 


S. Iceland: Nuphlidarhåls near Krisuvik (Stp.)!. 


8. Sphagnum teres (Schimp.) Ångst. 


Very common everywhere on damp ground up to a height of about 
600 metres above sea-level. 

S. teres is the most commonly occurring species of this genus and 
is almost as frequent in the lowlands as in the Alpine region, at high 
levels. It grows both on knolls in bogs and on more or less damp 
slopes- and is also common on warm ground. 

It is almost always green or pale yellowish-green in colour. Forms 
with appressed leaves are by far the most frequent; f. squarrosulum 
(Lesq.) is met with here and there, e. g.: Esja, Akureyri, Stadr near 
Hrutafjårdur and Laugaråshver. The walls of the chlorophyll-cells are 
very often distinctly papillose. Only sterile specimens have been found. 


9. Sphagnum squarrosum Crome. 


N. Iceland: Myvatn (Grl.!); Litlu Borgarkatlar (St.)!; Grimsey (0. D.)!. 
NW. Iceland: Isafjordur!; Grunnavik!; Sugandafjérdur!. 

In NW. Iceland it is rather common up to about 300 metres above 
sea-level. It grows there sometimes on knolls in bogs and sometimes 
on more dry ground, thus, near Isafjordur it grew in several places on 
slopes stretching down towards the rivers. Only sterile specimens have 
been found. 


10. Sphagnum fimbriatum Wils. 


N. Iceland: Akureyri!; Geitaskard!. NW. Iceland: Dynjandi!; Laugar- 
land!. W. Iceland: Ketilstadr (H. J.)!; Stykkisholmur (H. J.;!); Nordredalur 
in the district of Borgarfjérdur!. S. Iceland: Geldingsey (H. J.)!. 

This plant grows usually on very wet ground in bogs and is no 
doubt commonly distributed in northern and western Iceland. Thus, 
near Stykkisholm it was common in bogs, and also around Akureyri 
where it occurred abundantly up to about 400 metres above sea-level. 


11. Sphagnum Girgensohnii Russow. 


E. Iceland: Eskifjérdur (H. J.)!; common near Seydisfjordur!.  N. Ice- 
land: common near Eyjafjordur (O.D.;!); Geldingafell (St.)!; Hafrardalur 


432 A. HESSELBO 


(St.)!… Hvaneyrardalur (St.)!.. NW. Iceland: common near Isafjérdur!; 
Dyrafjordur!; Dynjandi!. W. Iceland: Budir (H. J.)!; Nordredalur in Bor- 
garfjordur!. S. Iceland: Almannagjå!; Laugarvatn!; Ingjaldshéll (H. J.)!; 
Heidi in Myrdalur (H. J.)!; Seljaland!. 

Commonly distributed both on knolls in bogs and on drier ground. 
Thus, in lava-clefts near Thingvellir, it is frequently met with on humus- 
covered blocks associated with Rhacomitrium hypnoides and Hylocomium 
spp. It is of most frequent occurrence in the lower country, but near 
Eyjafjordur it ascends to about 600 metres and near Isafjérdur to about 
400 metres above sea-level. Fruit was found in several places. It 
occurs especially in orthocladous forms. 


12. Sphagnum Russowii Warnst. 


S. Iceland: Thingvellir !. 


13. Sphagnum Warnstorfii Russow. 


Common on knolls in bogs and on damp ground on slopes up to 
a height of about 500 metres. 

Reddish forms are of most frequent occurrence, violet, green and 
yellowish-green forms are far more rarely met with. Only sterile spe- 
cimens have been found. 


14. Sphagnum rubellum Wils. 


Common on knolls in bogs and on wet slopes especially in the 
lowlands, but it also frequently ascends to about 500 metres above sea- 
level. Red and violet forms are frequent, green ones are rarer. Only 
sterile specimens have been found. 


15. Sphagnum acutifolium Ehrh. 


ln Islandia legs Morch UØN STCeland FH oOo OSDY) NEN WÆTeeland: 
Isafjordur!; Armuli!. W. Iceland: Reykholtdalur on warm ground (Grl.;)); 
Reykjavik!; between Thingvellir and Geysir (Grl.)). 

Occurs here and there on boggy ground in the lowlands and as 
a rule rather sparingly. Near Isafjérdur it was found in a bog at an 
altitude of 250 metres. Only sterile specimens have been found. 


16. Sphagnum subnitens Russ. et Warnst. 


NW. Iceland: Bæir!:; Thorvaldsdalur (St.). W. Iceland: Grund!: 
Snældubeinstadahver in Reykholtdalur!; Reykjavik (H.J.;/). SS. Iceland: 
Laugaråshver!; Borg (H. J.)!; Krisuvik (Stp.)!. 

Found here and there in NW., W. and S. Iceland where it occurs 
both in bogs and along the margin of hot springs. It is frequent around 
Reykjavik where it was also gathered in fruit. It is as a. rule pale 
yellowish-green in colour. Near Laugaråshver a f. violascens was found 
growing in large cushions on warm ground. 


< 


THE BRYOPHYTA OF ICELAND 433 


17. Sphagnum Lindbergii Sch. 


W. Iceland: Ketilstadr among $. fimbriatum (H. J.)!. 


18. Sphagnum riparium Ångst. 


NW. Iceland: Thoroddstadaengjar (St.)!; common in pools and in 
very wet parts of bogs near Jåkulsfjordur!. 


19. Sphagnum angustifolium C. Jens. 


N. Iceland: Akureyri!. W. Iceland: Reykholtdalur near a number of 
hot springs!. S.lceland: Reykirdalur!; Laugaråshver'!. 

This species occurs abundantly along outlets from the hot springs 
in deep cushions of a pale green, yellowish or light brownish-yellow 
colour. The leaves are distinctly wavy in all the plants from a warm 
substratum. Its. occurrence at Akureyri is peculiar; there it grew at 
an altitude of about 500—600 metres on wet boggy ground as a slender, 
pale-green, but otherwise typical, form. Only sterile specimens have 
been found. 


20. Sphagnum Dusenii C. Jens. 
W. Iceland: Snældubeinstadahver in Reykholtdalur!. 


In the above locality it grew in large yellowish-brown tufts in 
warm Water of a temperature of 25”, near a hot spring. 





434 A. HESSELBO 


MUSCI VERI 
Fam. ANDREÆACEÆ. 
1. Andreæa petrophila Ehrh. 


Very common all"over Iceland. In the lowlands it grows in small 
round cushions on blocks and on rocks in dry situations and in such 
localities it is often the dominant species. On mountain heights, where 
it is frequently met with right at the upper limit of vegetation, it often 
descends upon the ground, even upon damp gravelly flats, where it 
forms extensive cushions almost black in colour. Fruit is almost al- 
ways present. 

Note. Andreæa rupestris L. = A. Rothii W. M. is recorded by Morch 
from Iceland and is figured in Flora Danica, tab. 2125, but Mårch's 
specimens must be referred io A. petrophila. 


Fam. ARCHIDIACEÆ. 
2. Archidium phascoides Bridel. 


W. Iceland: Reykholtdalur!, near all the hot springs (Grl.;!); Englands- 
hver!. S. Iceland: Reykirdalur!; Reykjanes (Ostf.)!; Kolvidarhol!; Thorlåks- 
hver!; Laugaråshver!; Sydri Reykjahver!; Geysir!. 

Archidium phascoides occurs in the collections under different names. 
It was first collected by Grånlund and determined by Zetterstedt 
as Leskea nervosa. Afterwards this specimen with several others were 
re-determined by Grånlund as Catoscopium nigritum var. Grånlundii 
established by C. Jensen. All the plants collected by Ostenfeld 
from Reykjanes are determined by C. Jensen as Pohlia nutans v. 
filicaulis. The reason for all these erroneous determinations will be found 
in the fact that the piant is almost always found barren and in leaf- 
form and cell-tissue presents a certain resemblance to mosses belonging 
to quite different groups, which often leads to the belief that the spe- 
cimens in question are stunted forms or have been affected by the warm 
substratum: thus, certain forms bear a striking likeness to a slender 
Amblystegium compactum. The plant is typical of warm clay flats, and 
in S. and W. Iceland it is met with around every hot spring. On" the 
warm clay flats with a temperature of 207—40%, it forms extensive 
vellowish-green mats 1—2 cm. high. On slopes stretching down towards 
the boiling water and along outlets from the springs it is also frequently 
found growing within a few centimetres from the water, usually mixed 


THE BRYOPHYTA OF ICELAND 435 


with Haplozia crenulata and Fossombronia Dumortieri, more rarely with 
Gymnocolea inflata. It is almost always sterile; on plants from Thor- 
låkshver flowers as well as a few ripe capsules were found, besides, 
here and there at the bottom of the tufts, the characteristic large 
spores (9. 6. 1912). 

Note. Phascum cuspidatum, Ephemerum serratum and Sphærangium 
muticum are enumerated on older lists, but no specimens of these 
species are to be found in the collections and their occurrence in Ice- 
land is very improbable. 


Fam. WEISIACEÆ. 
3. Gymnostomum rupestre Schleich. 


E. Iceland: Geithellir!.  N. Iceland: Akureyri!; Modruvellir; Hof 
(0: D.):  W. Iceland: Brynjudalur (Grl)!; Esja!… -S. Iceland: Skålbolt!; 
Thingvellir! 

Around Eyjafjordur it is common on damp rocks up to a height 
of about 300 metres; it was also found in numerous localities in SW. 
and S. Iceland and is common, for instance in Esja. It occurs especi- 
ally on damp rock-sides near waterfalls and in clefts, often forming 
extensive mats; now and then it is also found in dry localities. Only 
sterile specimens have been found. : 


4. Hymenostylium curvirostre (Ehrh.) Lindb. 
Barbula curvirostris Lindb. 


N. Iceland: Myvatn (Grl.)!. "W. Iceland: Reykholt (Grl.)!, in both 
localities sparingly and also sterile. S. Iceland: Merkjåfoss (F.)!; Bar- 
karstadr | fr: Holt fr! SVestmannacy,, fr:l. 

In S. Iceland the plant is no doubt common on faces of damp tuft- 
rocks. In several localities near Barkarstadr and Holt it occurred 
abundantly in clefts along rivers or in caves and in depressions in tuff 
rocks; it was usually richly in fruit, and associated with Preissia com- 
mutata, Fegatella, Haplozia riparia, etc. On "Kleven” on the island of 
Vestmannaey it grew on the face of a tuff rock, through which water 
was percolating, forming with Haplozia riparia large cushions which 
were saturated with water. 


5. Anoectangium compactum Schwgr. 


Pleurozygodon æstivus (Hedw.) Lindb. 


SIn Islandia” leg. Mérch! and figured in Flora Danica, tab. 1893, 
fig. 2, under the name of Gymnostomum curvirostre. 

E. Iceland: Seydisfjordur, common!; Berufjårdur, common!. W. Ice- 
land: Grundarfjordur (H. J.)!; Hafnarfjordur (Grl.;”; Esja!; Svinahraun!. 
S. Iceland: very common!; Vestmannaey !. 

This plant is very common in S., SW. and E. Iceland, but it does 
not appear to occur in N. and NW. Iceland. It grows especially on damp 


436 A. HESSELBO 


rocks, e€. g. on rock-sides along waterfalls and in clefts, where it forms 
large cushions, often almost one square metre in size, of a peculiar 
verdigris colour. In the lava-fields of SW. Iceland it is one of the most 
frequent species on the walls of caves and the sides of clefts. It hardly 
ascends higher than about 300 metres above sea-level. It is almost 
always sterile. The only fruiting specimen was collected by Morch, 
but no habitat is given. 


6. Weisia crispata (Br. germ.) Jur. 


SW. Iceland: Reykjavik, fr.!. SE. Iceland: Hornafjérdur!; SW. Ice- 
land: Ålafoss!. 

Near Reykjavik it grew in crevices of basalt rocks associated with 
Encalypta rhabdocarpa and Bryum elegans var. carinthiacum. The fruit 
had just ripened on June 6, 1909. Near Hornafjordur also it grew in 
small crevices of basalt. Only sterile specimens have been found. 


7. Weisia viridula (L.) Hedw. 


S. Iceland: Vik, associated with Brachythecium velutinum (H. J.)!. Vest- 
mannaey, Helgafell!, on a humus-covered block of lava. 

Only sterile specimens have been found. 

The species is recorded by Mårch from Iceland, but no specimens 
of it are to be found in the collections. 


8. Weisia Wimmeriana (Sendtn.) Br. eur. 


N. Iceland: Hof near Eyjafjordur (O. D.)!, fr. 
It grew there mixed with Eurynchium diversifolium on rocks coated 
with soil. 


9. Dicranoweisia crispula (Hedw.) Lindb. 


Very common all over Iceland. 

Dicranoweisia crispula is one of those species which is of equal 
frequency in all parts of Iceland, both at the coasts and in the high- 
lands, and at all altitudes up to the snow-line. In the low land it 
grows, like Andreæa, on dry stones and rocks, but like the latter, it 
descends to the ground on mountain heights where it forms, both on 
the ground or on blocks of rock, dense, jet-black cushions. It is almost 
always found in fruit. 

Note. Dicranoweisia cirrata is recorded by Hornemann, but has 
undoubtedly been confused with D. crispula. 


10. Dicranoweisia compacta (Schleich.) Schimp. 


N. Iceland: Hof near Eyjafjordur (O. D.)!. Only sterile specimens 
have been found. 


mA] 


THE BRYOPHYTA OF ICELAND 43 


Fam. RHABDOWEISIACEÆ. 
11. Cynodontium polycarpum (Ehr.) Schimp. 


Seydisfjordur (C. Hansen) !. 

This species together with the var. strumiferum, is recorded by 
Vahl, but has without doubt been confused with another species, pro- 
bably a Dicranum or Oncophorus. Morch also records both forms 
from Iceland, but of his specimens in the Botanical Museum in Copen- 
hagen some are Dicranum Starckei, and some are Ceratodon purpureus. 

Cynodontium gracilescens var. tenellum is recorded by Carrington 
from Akureyri, but has undoubtedly been confused with another species. 


12. Dichodontium pellucidum (L.) Schimp. 


Very common everywhere on damp ground. 

This plant is most widely distributed in the lowlands and is found 
there everywhere on damp ground, both in bogs and on gravelly soil 
along streams, and also on soil-covered rocks. In the lowlands it usually 
grows mixed with all kinds of other mosses in their tufts and is, as a 
rule, low in growth and short-leaved. On rocks near the water, where 
it is also frequent, more vigorous forms occur in unmixed tufts, 2—4 cm. 
high. At higher levels it occurs more sparingly, but is nevertheless fre- 
quently found up to about 600 metres above sea-level. Only sterile 
specimens have been found. 


Fam. ÅONGSTRÆMIACEÆ. 
13. Aongstræmia longipes (Sommerf.) Br. eur. 


E. Iceland: Høornafjérdur!; Hof!; Seydisfjordur!; Vallanes!. N. Ice- 
land: Akureyri (O. D.;!); Husavik!; Vidimyri!; the Tverå!; Dettifoss!; Stadr 
near Hrutafjordur!.  NW. Iceland: Årmuli!. OW. Iceland: Haukadal!:; 
Lundur!; Alafoss!.  S. Iceland: Breidabolstadr!; Barkarstadr!; Seljaland '!. 

This species is doubtless common everywhere, but is often over- 
looked on account of its minute size. It grows on damp ground, most 
frequently as single specimens scattered among other mosses; more 
rarely, as for instance near Hornafjérdur and Breidabolstadr, abundantly 
on peat along ditches. Near Lundur it grew abundantly associated with 
Dicranella Schreberi, Bryum lacustre, Dichodontium and other species in 
places where the peat had been pared off. 

In the majority of the cases the specimens were not found until 
the material brought home had been investigated, because in habit they 
exactly resemble the tiny Pohlia-forms of wide occurrence, or a small, 
sterile Anomobryum. 

On boggy ground it was usually collected together with Dichodon- 
tium pellucidum, Oncophorus Wahlenbergii, Meesia trichoides, Bryum spp., 
etc. On gravelly soil near Stadr it was found associated with Dicranella 
crispa, Didymodon rubellus, Blasia pusilla and Haplozia atrovirens and in 
several places with Distichium inclinatum. It occurs both in the low 


438 A. HESSELBO 


lands and up to 500—600 metres above sca-level. Near Akureyri it 
was found on a damp flat at about 500 metres. Near Barkarstadr it 
occurred abundantly on boggy ground at about 530 metres, and near 
Seljaland it was also frequent at an altitude of about 500 metres on a 
moss-grown gravelly flat. Only sterile specimens have been found. 


Fam. DICRANACEÆ. 
14. Oncophorus virens (Sw.) Brid. 


Very common on damp ground. It grows both on wet humus- 
covered rocks and in bogs, and also on damp gravelly ground, somec- 
times as unmixed tufts and sometimes mixed with other mosses; it is 
of equal frequency at all altitudes as far upwards as 600—700 metres. 
It often occurs in great quantities and usually fruits richly. In the 
losmverdistricts the fruer pens about ny est nESlcelandets 
species is either wanting or rare in the lowlands up to a height of 
about 300 metres (it has not been found on Vestmannaey), but from 
this height up to the snow-line it occurs abundantly in bogs. 

Var. serratus Br. eur. occurs occasionally on wet rocks. 


15. .- Oncophorus Wahlenbergii Brid. 


One of the most commonly. occurring mosses; it is found every- 
where in damp localities just like O. virens. It is very common espe- 
cially in bogs, except in S.Iceland where, like O. virens, it has its dis- 
tribution from an altitude of 300 metres upwards. 

Var. f compactus (Funch) Br. eur. E. Iceland: Skreiddal!; Seydisfjårdur!. 
It grew .in both localities on knolls in bogs. 

Var. elongatum Hagen.  W. Iceland: Nordredalur!. It grew there on 
boggy ground in association with Sphagnum teres, S. rubellum and S$. fim- 
briatum in loose tufts about 6 cm. high. 

Although typically developed forms of Oncophorus Wahlenbergii and 
O. virens differ so widely from each other that they are not likely to 
be -confused, yet so many transitional forms occur, especially in Alpine 
habitats, that very often it is a matter of opinion to which species such 
a form should be referred. On the same plant leaves may be found 
which, as regards the cells of the leaf-base, resemble sometimes O. 
Walhlenbergii and sometimes O. virens; also the leaf-margin is more or 
less recurved or plane. 


16. Dicranella squarrosa (Starke) Schimp. 


E. Iceland: Seydisfjérdur!; Skorastadr (H. J.)!.. N. Iceland: Husavik!; 
near Eyjafjérdur (O. D.;!), fr.; Lækjarmot!. NW. Iceland: Very common?!. 
W. Iceland: Vogur (H. J.)!: Esja!; ;Reykholtdalur!. S. Iceland: Holt! 

This plant grows almost always in water, especially on inundated 
gravelly ground by streams, on stones and in moss bogs, more rarely 
on wet marshy ground. In NW. Iceland it is one of the mosses of 
most frequent occurrence being found in abundance associated with 


THE BRYOPHYTA OF ICELAND 439 


Philonotis seriata, Oncophorus Wahlenbergii, Scapania uliginosa, Haplozia 
cordifolia and Chiloscyphus polyanthus var. fragilis on inundated gravelly 
ground up to about 300—400 metres above sea-level. In N. Iceland it 
is also rather common. Near Seydisfjordur it was found only at a 
height of about 300 to 500 metres above sea-level, but was common 
there in small streams associated with Scapania undulata, Hypnum ex- 
annulatum, Haplozia cordifolia, etc. In Reykholtdalur this species was 
found in several places near hot springs, even on stones in warm water 
with a temperature of 30”. In SW. and S. Iceland it occurs rarcly and 
more sparingly; near Holt it grew in several places at an altitude of 
150—600 metres. 

The plant is almost always found sterile. One specimen collected 
by O.D. near Hof in Eyjafjérdur (12. 6. 1897) bore a few capsules which 
had thrown off their lids. The gg plant was found only near Bæir in 
NW. Iceland growing in low tufts, about 1 cm. high. 


17. Dicranella Schreberi (Sw.) Schimp. 
Anisothecium crispum (Schreb.) Lindb. 


N. Iceland: Hof near Eyjafjordur (0. D.)!; Akureyri Q!. W. Iceland: 
Lundur!. 

Near Akureyri it grew in damp, peaty ground immediately north 
of the town and near Lundur, associated with Bryum lacustre, AÅong- 
stræmia longipes and Dichodontium pellucidum, on a wet flat from which 
the peat had been pared off. 


18. Dicranella crispa (Ehrh.) Schimp. 


Very common on damp ground, both on peat and on sandy or 
gravelly soil by streams. It occurs only in the lowlands, and has hardly 
been found higher than at an altitude of about 200.-metres. Fruit is 
always present in abundance and ripens about August ist. At this time 
also the setæ of the capsules of the ensuing year have reached their 
full length. 

Note. Dicranella heteromalla is recorded by Zoéga and Horne- 
mann from Iceland, but was undoubtedly confused with another species. 


19. Dicranella subulata (Hedw.) Schimp. 
Dicranella secunda (Sw.) Lindb. 


Iceland (Mårch)!, some of the specimens are correctly determined, 
some must be referred to Ditrichum homomallum. &Krisuvik (Stp.)!. N. 
Iceland: Husavik!. NW. Iceland: Kaldalon!; Gnupsdalur near Dyrafjérdur'!. 
Moreover, Groånlund records it from Hvammur and Thingvellir, but 
both specimens are wrongly determined, being Dicranella crispa and 
Dicranum Blyttii. 

This species was collected in small quantities only, but everywhere 
in fruit. Near Husavik it grew on peat, and on July 11, 1909 it bore 


440 A. HESSELBO 


both quite young capsules and also old lidless capsules. In Gnupsdalur 
it grew along the banks of streams with Scapania subalpina and Cepha- 
lozia bicuspidata. 


20. Dicranella cerviculata (Hedw.) Schimp. 


Iceland: (Morch)!. NW. Iceland: Stadr on Snæfjallastrénd (Stp.)!; 
Dynjandi!. W. Iceland: Borgarnes!; Kollafjérdur!; Reykjavik (Grl.;!). S. 
Iceland: Thjorså (Stp.)!. 

The plant was found everywhere in fruit. Near Borgarnes and 
Reykjavik the fruit was almost ripe at the beginning of August. It grows 
on peat, especially on the rather dry margins of turf pits and ditches 
and occurred abundantly especially in several places near Reykjavik. 


21. Dicranum fulvellum (Dicks.) Sm. 


Collected in Iceland by Mårch (no doubt near Middalur) and 
figured in Flora Danica, tab. 2002, under the name of D. Mårchianum 
Hornsch. S. Iceland: Holt!; Seljaland!; Thingvallahraun!. SW. Iceland: 
Svinahraun !. 

This species is recorded by Gronlund from Svinaskard and Thing- 
valla, but the specimens from both localities are wrongly determined, 
being really D. falcatum and D. Blyttii respectively. 

Dicranum fulvellum has been only found in the south-western part 
of Iceland, where it appears to be frequent. In Svinahraun, where the 
altitude hardly exceeds 300 metres, it was found rather sparingly (June 
3rd) on the top of lava-blocks, in fruit which was almost ripe. On 
Eyjafjall it occurred in several localities above Holt and Seljaland at 
an altitude of 300—700 metres. It grew either on blocks of basalt or 
more rarely on gravelly ground in company with Gymnomitrium concin- 
natum. At the end of July the fruit was ripe everywhere, and generally 
the capsules had thrown off their lids. 


22. Dicranum Andersonii (Wich.) Schimp. 


SE. Iceland: Hof!; coastal cliffs near Alftafjordur!. S. Iceland: Holt!; 
Vestmannaey!; Thingvellir!. 

Near Hof it occurred rather commonly on low basalt ridges, where 
it usually grew in crevices in small cushions about 3 cm. high. Here 
the fruit had just ripened on June 15th and the lids still persisted. 
Near Holt it was found rather frequently on tuff rocks up to a height 
of about 300 metres. Thus, it grew abundantly, for instance on the 
vertical northern face of a tuff rock sometimes in small tufts, sometimes 
mixed with Grimmia apocarpa, Pogonatum urnigerum, Pohlia grandiflora, 
Anthelia nivalis and Gymnomitrium concinnatum. On Vestmannaey the 
plant grew abundantly on blocks of lava lying on the slope of Helga- 
fell. It occurred there everywhere in small scattered tufts associated 
with Grimmia apocarpa, Rhacomitrium sudeticum, R. fasciculare and An- 
dreæa petrophila. The fruit was ripe on July dth; a few had thrown 
olfetheirslids 


THE BRYOPHYTA OF ICELAND 441 


In Iceland Dicranum Andersonii is a decidedly lowland plant, which 
is probably widely distributed in the southern and south-eastern parts 
of the country. It occurs especially near the coast; in the interior of 
the country it has been found only near Thingvellir, where it grew very 
sparingly on a lava block in company with D. fulvellum, which it 
resembles in the fact that the peristome teeth are wide-spreading when dry. 
Dicranum Andersonii is however easily distinguished by its shorter seta 
and by the extremely thick-walled cells of the capsule-walls and by the 
absence of stomata. 


23. Dicranum falcatum Hedw. 


E. Iceland: Lonsheidi!, at an altitude of about 400 metres; Beru- 
fjordur!. N.Iceland: Ljosavatnsberg!. NW. Iceland: Dyrafjérdur!; Isa- 
fjordur!. W. Iceland: Snæfellsnes: Saxarhol (H.J.)!; Frodarheidi (H. J.)!; 
Reynivellir (Grl.)!; Svinaskard (Grl.)!; Kolvidarhol!; Esja!. S.Iceland: Holt!; 
Seljaland!. The plant is recorded by Mårch from Iceland and figured 
in Flora Danica, tab. 2003, but Méårch's specimens in the Botanical 
Museum in Copenhagen belong to Dicranoweisia crispula. 

Dicranum falcatum is a plant characteristic of the damp gravelly 
flats of mountain heights, where it often occurs abundantly in large, 
compact, rounded cushions in the neighbourhood of patches of snow. 
Usually it is not met with until an altitude of about 600 metres; near 
Kolvidarhol, however, it was common even at an altitude of about 400 
metres. This species is very common in NW. and in S. Iceland and 
occurred also abundantly on Ljosavatnsberg. It is probably common 
everywhere on mountain heights, but, as for instance in E. Iceland, it 
has only been collected scantily, as its habitats are snow-covered in 
June when that part was explored. 

The fruit, which is always present abundantly, ripens about the 
middle of July. 


24. Dicranum Blyttii Schimp. 


Dicranum Schisti (Gunn.) Lindb. 


E. Iceland: Hamarfjérdur!. UN. Iceland: Reykjaheide!; Vidvik (P.S.)!; 
Hof (0. D.)!. NW. Iceland: Patreksfjårdur (Wiinstedt)!; Glåma (St.)!, com- 
mon on mountain heights!. OW. Iceland: Øxarrhyggur!; Svinahraun!; 
Hafnarfjérdur!. S. Iceland: Seljaland!, from an altitude of 350 metres 
upwards; Holt!, common on mountain heights; Thingvallahraun!. 

This species is common, at any rate in NW. and SW. Iceland, while 
it has been found but scantily in E. Iceland and only in a few, localities 
in N. Iceland. But the same is probably the case here as mentioned 
for Dicranum falcatum, that its habitats, at the time when E. Iceland 
was explored (June 1909), were covered with snow. 

Dicranum Blyttii belongs to the Alpine region and is not met with 
until an altitude of 300—400 metres. Near Hamarfjordur, however, it 
grew in the lowlands on rocks near the coast. It grows partly on 
stones and partly on gravel, and is especially frequent at a height of 
600—800 metres, both on gravelly flats soaked by the melting snow, 

The Botany of Iceland, Vol. I, part Il. 929 


449 A. HESSELBO 


and on scattered blocks of rock. It often forms there extensive, blackish- 
green or almost black cushions, 2—3 cm. deep. 

In lava-fields it occurs associated with several other Alpine species 
at the bottom of ravines, as on Reykjaheide (at an altitude of about 
300 metres), in Thingvallahraun, where it was rather common on fallen 
blocks in ravines (Gjå), and near Hafnarfjordur, where it was also rather 
frequent in lava-clefts. In Svinahraun it grew abundantly on damp 
ground in a large depression at the edge of the lava-field. 

The leaves usually spread erectly, more rarely, for instance in plants 
from Thingvellir, they are secund. The fruit ripens almost at the same 
time on mountain heights and in the lava-fields, viz. at the end of July. 
Near Thingvellir it was found on July 29th with fully ripe capsules, 
some of which had thrown off their lids. 


25. Dicranum Starckei Web. et Mohr. 


"In Islandia” (determined by Mårch as Dicranum polycarpon). 

Widely distributed all over Iceland. 

This species is most widely distributed on damp gravelly flats from 
an altitude of about 400 metres upwards to the snow-line and is often 
there, especially from about 400—600 metres, the most abundant constituent 
of the vegetation, growing in extensive, continuous mats, 2—3 cm. deep. 
It oceurs however also on stones and damp rocks. It is found more 
scantily in the more low-lying parts of the country, although, especially 
in NW. and N. Iceland, it is frequent, mixed with other møosses, on 
gravelly ground by streams. In the southern part of the country it 
does not appear to descend to a lower level than 300—400 metres. 

Like Dicranum Blyttii this species grows also in lava-clefts, and it 
was common on fallen blocks both near Hafnarfjérdur and Thingvellir. 
The plants from these localities were often 5—6 cm. high, with longer 
leaves, in which feature they approached very closely to D. molle. 

The direction of the leaves varies considerably, orthophyllous forms 
are the most frequent, especially on the ground; highlv drepanophyllous 
forms occur more particularly in drier localities, for instance on rocks. 

Fruit is almost always present. The majority of the specimens 
collected (June—July) had old, empty capsules, and also young green 
ones or half-ripe ones which probably ripen late in the summer. 


26. Dicranum molle Wils. 
D. arcticum Schimp. D. glaciale Berggr. 


E. Iceland: Seydisfjordur!, at an altitude of about 200 metres. N. 
lceland: Akureyri!. NW. Iceland: Dyrafjérdur!; Sugandafjérdur!; Isa- 
fjordur!; Kaldalon!; Dynjandi!. W.Iceland: Snæfellsnes, Mafahlid (H. J.)!, 
at an altitude of about 300 metres; Budahraun (H. J.)!; Dalasysla, Vig- 
hålsstadir (H. J.)!, 550 metres alt.; Olafsvig (Mérch)!; Svinaskard (F.)!; 
Esja, Mådruvellir!. at an altitude of about 450 metres; Hafnarfjérdur!. 
S. Iceland: Thingvallahraun !. 

This species is also recorded from several localities by Gronlund, 


THE BRYOPHYTA OF ICELAND 443 


but the specimens in the Botanical Museum are wrongly determined; 
most af them must be referred to D. Starckei or D. angustum. 

Dicranum molle usually grows on damp gravelly ground on moun- 
tain heights. In NW. Iceland it is one of the most commonly occurring 
mosses on rocky flats, and is common down to about 200 metres above 
sea-level; but also frequently descends as far down as the bottom of 
the valley. It grows both on gravelly flats soaked by melting snow 
and on damp slopes; also on lower levels on heathy soil, with other 
Dicranum spp. (D. scoparium and D. fuscescens), among Vaccinium, Em- 
petrum, etc., in large tufts which are usually closely interwoven with 
Lophozia lycopodioides, L. Kunzeana, L. quinquedentata, L. Floerckei, L. 
ventricosa and other Hepaticæ. Near Mådruvellir it grew partly on damp 
gravelly ground, partly intermixed in åbundance in Rhacomitrium hyp- 
noides-heaths; in this latter locality also it was mixed with Lophozia spp. 

In lava-fields it usually grows in company with D. Starckei, commonly, 
as for instance in the clefts near Thingvellir, in cushions as much as 
10 cm. deep. The plants from the latter locality and from Hafnar- 
fjordurhraun are sometimes not quite typical and approach so closely 
to D. Starckei, that they can hardly be distinguished. The leaves are 
sometimes orthophyllous, sometimes drepanophyllous. The fruit, which 
is always present abundantly, ripens during July. In the district of 
Isafjordur, however, ripe fruit was frequently found as early as in the 
latter days of June. 


27. Dicranum Bonjeani de Not. 
D. palustre Br. eur. 


SE. Iceland: Hornafjérdur!; Lon!.  N. Iceland: near Eyjafjérdur 
(O. D.)!. W. Iceland: Dalasysla; Ketilstadr GEL )NS Reykhboltdalur (Grl. 1); 
Reykjavik!. 

The plants from all the above-mentioned localities agree exactly 
with D. Bonjeani, but being quite sterile they cannot be determined with 
perfect certainty. Bog-forms of Dicranum scoparium have often erect 
leaves, slightly undulate at the apex, and åre then extremely difficult to 
distinguish from D. Bonjeani. This species grows in bogs, usually on 
knolls, and is common around Reykjavik and near Hornafjordur. 

Note. In several older lists Dicranum undulatum is enumerated as 
found in Iceland by Mårch and is also recorded by Gronlund from 
Reykholt and Reykjavik, but all the specimens in the herbaria must 
either be referred to D. scoparum or D. palustre. 


28. Dicranum angustum Lindb. 


E. Iceland: Hornafjérdur!; Berufjérdur!; Lon!; Hof!. N. Iceland: 
Myvatn (Grl.; determined as D. arcticum)!; Stadr near Hrutafjérdur!. NW. 
Iceland: Dynjandi!. W.Iceland: Stykkisholmur!; Reykjavik!. S. Iceland: 
Asolfskålaheidi near Holt!, at an altitude of about 400 metres. 

This species is no doubt widely distributed over the whole of Ice- 
land; it is, at any rate, rather common in E. and SW. Iceland, where 


29" 


444 A. HESSELBO 


it often occurs abundantly. It grows on wet boggy ground in loose 
tufts as much as 6 cm. high, either pure or mixed with Sphagna and 
Hypnum stramineum and interwoven with Hepaticæ such as Lophozia 
quinquedentata, Scapania irrigua, Cephalozia pleniceps, etc. Only sterile 
specimens have been found. 

This species is most widely distributed in the lowlands and has 
only once been collected at an altitude of 400 metres on Asolfskålaheidi, 
where it grew plentifully in a bog. 

Hagen (Musci Norv. Borealis, p. 22) states that D. angustum differs 
from all nearly allied species, and especially from D. scoparium, in having 
thin-walled cortical cells; but this distinction does not suffice as regards 
bog-forms of D. scoparium as in the latter species all transitional stages 
are found from the thin-walled, hyaline cortical cells of plants from 
wet ground to the thick-walled, brown cells of xerophilous forms. 


29. Dicranum majus Smith. 


NW. Iceland: Dyrafjérdur!; Sugandafjéordur!; Isafjordur!. W. Iceland: 
Neshraun (H. J.)!: Budahraun (H. J.)!. 

In the two habitats in W. Iceland, both of them on Snæfellsnes, 
the plants were collected in depressions in the lava-field. In NW. Ice- 
land, where it is commonly distributed, it grows on the ground on 
slopes covered with birch coppice, Vaccinium, etc., as far upwards 
as about 300 metres above sea-level. 

The majority of the plants collected have slightly secund leaves 
and entirely resemble D. scoparium in habit. A vigorous form with 
strongly falcate leaves was found in a birch coppice near Dyrafjordur 
at an altitude of about 150 metres. Only sterile specimens have been 
found. 


30. Dicranum scoparium (L.) Hedw. 


Very common. 

D. scoparium is most widely distributed in the more low-lying parts 
of the country up to a height of about 400—500 metres, but also ascends 
frequently to about 700 metres above sea-level. In S. Iceland it appears 
to be more rare and, as for instance on Fljåtshlid and south of Eyja- 
fjall, was only found in a few localities and very scantily. It grows 
both on rather dry and on damp ground, and also on rocks and on 
knolls in bogs, and varies considerably. Orthophyllous forms are the 
most common, especially on damp ground, while more or less distinctly 
drepanophyllous forms grow by preference in dry localities, for instance 
in lava-fields and on rocks. In bog-forms the leaves are slightly undulate 
at the apex, in which feature it approaches D. Bonjeani. The leaf-apex, 
especially in orthophyllous forms, is usually short and broad, and 
faintly toothed, or even quite entire, at the margin, at the same time 
the lamellæ on the back of the leaves develop more slightly or dis- 
appear almost entirely. Such forms agree well with var. integrifolium 
Lindb. 

Fruit is found rather rarely; in Nordredalur it was fully developed 
on August 2nd, but was still green. 


THE BRYOPHYTA OF ICELAND 445 


31. Dicranum fuscescens Turn. 


Iceland (Wiinstedt, determined by Berggren as D. Scottianum). NW. 
Iceland: very common!. W. Iceland: Nordredalur'!. 

In NW. Iceland D. fuscescens is the most frequently occurring species 
of this genus and is very common from the bottom of the valley up to 
a height of 400—500 metres. It is most widely distributed from 
about 100 to 300 metres on somewhat damp slopes covered with Vac- 
cinium-Empetrum heath, where it often constitutes the bulk of the 
Bryophyte vegetation in association with D. molle, D. scoparium and 
occasionally, D..majus. It is also common on the top of knolls in bogs. 
The densely matted tufts, which are as much as 10 cm. high, are al- 
most always interwoven with Hepaticæ, especially Lophozia spp., Ce- 
phalozia pleniceps, C. bicuspidata and Ptilidium ciliare. Im Nordredal it 
was growing in birch coppices in company with Hylocomium spp. 

The fruit, which occurs rather frequently, was quite green even in 
the beginning of July. 

All the Iceland specimens differ from the typical form and, more 
or less, especially approach Dicranum congestum without, however, form- 
ing any real transitional form to the latter. As a rule the leaf-nerve 
occupies about 7/4 the width of the leaf-base, but occasionally "/3—/6 
only. The cells of the leaf-apex are usually quadrate or roundish, 
mixed with numerous ones that are irregular in form; rarely, in the 
main, shortly rectangular, in which feature it approaches var. angusti- 
folium Arn. et Jensen; most frequently arranged more or less distinctly 
in rows; papillose at back, more rarely almost smooth, but in the same 
tuft or even on the same stem rather considerable variations occur as 
regards the development of the cell-tissue and the papillæ of the leaves. 


32. Dicranum congestum Brid. 


Very common in E. and N. Iceland and probably also in W. Iceland, 
somewhat rarer in S. Iceland. It appears to be quite absent from 
NW. Iceland and to be replaced there by D. fuscescens. 

It grows both on a dry and on a somewhat damp substratum, on 
earth and humus-covered stony slopes and on the top of knolls in bogs, 
and extends as far upwards as to the snow-line, for instance near 
Akureyri up to about 900 metres above sea-level. In S. Iceland it has 
not been found in the low land, but plentifully in several places on 
the rocky flat, for instance near Barkastadr, 500 metres above sea-level. 

Var. subspadiceum Arn. et Jensen occasionally occurs in localities 
similar to those of the type, but transitional forms are of more frequent 
occurrence. 

Var. flexicaule (Brid.) Br. eur. N. Iceland: Ljosavatn!, in Betula-nana 
heath; Hof (O. D.)!. 

Only sterile specimens have been found. 

Dicranum congestum is so variable a species that it is hardly pos- 
sible to find two plants which are exactly alike. The nerve occupies 
1/—"/10 the width of the leaf at its base, usually 7/s—"/9. The leaf-cells 


446 A. HESSELBO 


are irregular in form more or less far down the leaf, sometimes far 
below the middle, sometimes only at the leaf-apex itself; slightly or 
hardly porose, occasionally strongly porose in the lower half of the 
leaf, more rarely to near the apex, in the last case the walls are strongly 
incrassate. 

Dicranum brevifolium Lindb. is recorded from Vatnsdalsfjall (Grl., 
Tillæg til Islands Kryptogamflora, Bot. Tids., 20), but the plant from the 
latter locality is only a form of D. congestum in which the irregularly 
shaped cells are strongly incrassate and extend down below the middle 
of the leaf. Such forms, which, as a rule, are also distinguished by very 
strong nerves, have been found in several localities, for instance near 
Berufjérdur and about Eyjafjordur. The true D. brevifolium, which is dis- 
tinguished by its shorter leaves, crisped when dry, and its thin-walled 
cells, which are in a great measure quadrate and in rows, has not 
been found in Iceland. 


33. Dicranum spadiceum Zett. 


W. Iceland: Hvitidalur (H. J.)!. E. Iceland: Hornafjérdur!; Seydis- 
fjordur!. N. Iceland: Hof (0. D.)!; Vatnsdalsfjall (St.)!. 

Grows on a damp substratum, especially on knolis in bogs. Only 
sterile specimens have been found. 


34. Dicranum elongatum Schleich. 


«In Islandia” Mérch! (fr.). N. Iceland: By Hrutafjordur!. NW. Iceland: 
Dynjandi!; Hesteyri!. OW. Iceland: Stykkisholmur!. This species is also 
recorded from some localities by Gréånlund, but the specimens in 
the collections are wrongly determined, being D. Starckei, and others. 

D. elongatum grows everywhere on wet, boggy ground. By Hruta- 
fjordur it was common- on knolls in bogs along the eastern side of 
the fjord from Stadrbakki to Stadr, and it was also frequent in bogs 
near Stykkisholmur. 

All the specimens collected were quite typical. The nerve was 
about "/4 the width of the leaf at its base. With the exception of the 
specimen found by Mårch only sterile specimens have been collected. 

Note. According to Gliemann, Dicranum montanum was found 
in Iceland by Mårch, but no specimens of it are to be found in the 
collections, and the record of this species is undoubtedly due to an er- 
roneous determination. 


35. Campylopus Schimperi Milde. 


N. Iceland: Geitaskard!. E. Iceland: Hornafjérdur!; Lon!; Beru- 
fjordur!. W. Iceland: Reykjavik!; Esja (Grl.)!. S. Iceland: Breidabolstadr !. 

Commonly distributed in South-western and South-eastern Iceland 
on damp ground in bogs, where it forms compact cushions, 2—3 cm. 
deep. It has only been found in the low land. 


THE BRYOPHYTA OF ICELAND. 447 


36. Campylopus flexuosus (L.) Brid. 


N. Iceland: Myvatn (Grl.)!; Reykir near Svinavatn (Grl.)!. W. Iceland: 
Near hot springs in Reykholtdalur: Deildatunguhver (Grl.;!); Skribla (Grl.)!; 
Hagindishver!, etc. 

This species has been found only on steaming clayey flats near 
hot springs in North and West Iceland. In Reykholtdalur it is frequent 
on a warm substratum, and was found, for instance near Deildatungu- 
hver, in large mats, about 2 cm. deep, on warm clayey flats with a tem- 
perature of about 40? just below the surface. 


37... Campylopus fragilis (Dicks.) Br. eur. 


S. Iceland: Laugaråshver near Skålholt. 
It grew there on warm clayey flats close to the outlet from the 
spring. 


38. Trematodon ambiguus (Hedw.) Hornsch. 


W. Iceland: near Borgarnes!. 

It grew there rather scantily on damp peaty soil by the road, in 
company with Bryum inclinatum, Pogonatum nanum, Scapania curta, 
etc. The fruit had just ripened on Aug. 2nd, but the lids had not 
been thrown off. 

Note. Leucobryum glaucum is recorded by Vahl and Horne- 
mann as found in Iceland, but neither the name of the finder nor 
the habitat is given. No specimens of this species are to be found in 
the collections. 


Fam. FISSIDENTACEÆ. 
39. Fissidens bryoides (L.) Hedw. 


N. Iceland: Hof near Eyjafjérdur (O. D.)!. S. Iceland: Krokur (H., J.)! 
Almannagjå !. 

In Almannagjå it grew very scantily among other mosses on soil- 
covered stones at the bottom of a lava-cleft. Near Hof it grew on 
damp ground by a waterfall. The fruit, on the specimens from both 
localities, was ripe when collected (June 12). 


40. Fissidens osmundoides (Sw.) Hedw. 


Common on damp ground in bogs, at the base of rocks, in rock- 
clefts, etc., sometimes in low, dense tufts, sometimes mixed with other 
mosses. The plants are almost always found sterile. Only in Reyk- 
holtdalur, where it grows very abundantly on warm, boggy ground, has 
it been collected in fruit. It is most widely distributed in the lowlands 
up to about 300 metres above sea-level, but it may occur, although 
scantily, as high as about 500—600 metres. 


448 A. HESSELBO 


41. Fissidens adianthoides (L.) Hedw. 


N. Iceland: Hof near Eyjafjordur, fr. (0. D.)!. "W. Iceland: Klepp- 
jarnsreykir (Grl.!); Laugarnir near Reykjavik (Grl.)!; bog near Reykjavik, 
fr.!; Kollafjérdur! 

Rare and scanty on boggy or warm ground. ' 

Note. Fissidens taxifolius is recorded by Zoéga and Hornemann 
from Iceland (without finder, habitat or specimens in the collections) 
and by Gronlund from Myvatn; but Gronlund's specimen belongs to 
F. osmundoides. 


42. Fissidens decipiens De Not. 


W. Iceland: Sandur (H. J.)!; Hafnarfjérdur!. 
This plant grew in both localities in lava-clefts, associated with 
Lejeunea 'serpyllifolia. 


Fam. SELIGERIACEÆ. 
43. Blindia acuta (Huds.) Br. eur. 


Very common on wet rocks in or by rivers and waterfalls up to 
about 400—500 metres above sea-level, occasionally also on the ground 
and often in great abundance. The fruit, which is almost always pre- 
sent, ripens in the beginning of July. 

At a greater altitude it occurs more scantily, but may nevertheless 
be met with as far upwards as 600—700 metres. 


Fam. DITRICHACEÆ. 
44. Ceratodon purpureus (L.) Brid. 


Common on rocks, lava-flats, sandy soil, peat, etc. especially in 
N. and NW. Iceland, and on lava-flats in the interior of the country. 
It is less frequent in E. and S. Iceland. The fruit, which is usually 
present, ripens in the first half of July. 

It varies considerably, especially in leaf-form. Forms with shortly 
pointed or almost obtuse leaves (var. brevifolius) are the most frequent. 


45. Ditrichum tortile (Schrad.) Lindb. 


E. Iceland: Seydisfjordur!, sterile. 

It was growing there on damp gravelly soil along a river, inter- 
spersed in a Hepaticæ-mat formed by Alicularia scalaris, A. geoscypha, 
Anthelia Juratzkana, Lophozia ventricosa, etc. 


46. Ditrichum nivale (C. M.) Limpr. 


E Iceland: Berufjardarskard!, at an altitude of about 500 metres, fr. 
The plant was growing very scantily in a cushion of Anthelia 
Juratzkana. 


THE BRYOPHYTA OF ICELAND 449 


The entire height of the plant was about 6 mm. The seta was 
about 5 mm. high. The capsules were, without the lid, about 0.5 mm. long. 


47. Ditrichum homomallum (Hedw.) Hampe. 


Fl. Dan. tab. 2688, ex Islandia leg. Mårch. E. Iceland: "Vestdalur 
near Seydisfjordur!, sterile. N. Iceland: Hof near Eyjafjordur (0. D.)!, 
sterile. W. Iceland: Reykjavik (Grl.;!), fr.; hot spring near Grafarbakki 
(F.)!, sterile; Hafnarfjårdur, at the bottom of a lava cave!, sterile. S. Ice- 
land: Geysir! 2. 

Around Reykjavik it was rather common along the side of ditches. 
There the fruit was ripe on August 7th. Near Geysir it occurred in 
extensive mats, about 1—2 cm. deep, on warm clayey flats with a tem- 
perature of 35—40". 


48. Ditrichum flexicaule (Schleich.) Hampe. 


One of the most commonly occurring mosses; it is met with on 
almost every substratum and at all altitudes. . It grows both on damp 
or soil-covered rocks and in bogs, where it especially occurs woven 
into the tufts found there, and on: damp or on more dry gravelly soil, 
in lava caves, in grass fields, in heaths, etc. 

This species varies considerably, especially as regards the length 
and direction of the leaves. Short-leaved forms (f. brevifolia) are fre- 
quent on more dry ground, where they form dense tufts with stiffly 
erect or slightly secund leaves. On damp ground and especially in 
sheltered localities, for instance among stones, or in clefts, the leaves 
become longer, often strongly secund, and the tufts higher and looser. 
Fruit it very rare. Near Seydisfjordur, in the beginning of July, the 
setæ had not developed to their full length and the capsules were 
barely formed. 


49. Sælania cæsia (Vill.) Lindb. 
Ditrichum glaucescens (Hedw.) Hampe. 


E. Iceland: Breiddalur!; Hallormstadr!. N. Iceland: Vidimyri (Grl.)!; 
several places near Eyjafjordur (0. D.;!). W. Iceland: Husafell (GrlL)!; 
Kalmanstunga (Grl.)!; Gilsbakki!; Hafnarfjordur!; Esja!. S. Iceland: Skål- 
holt!; common near Thingvellir'!. 

This species is rather common, especially in W. and SW. Iceland, 
in soil-filled rock and lava clefts, as a rule associated with Mnium 
orthorrhynchum, Plagiothecium Roeseanum, P. pulchellum, Pohlia cruda, 
Distichium montanum, etc. 

In the wood by Hallormstadr it was growing on the ground in 
company with Bartramia ityphylla, Pohlia cruda, Distichium inclinatum 
and Ditrichum flexicaule. The fruit, which is almost always present, 
ripens in the middle or at the end of June. The plant has not been 
observed at a higher altitude than about 200—300 metres. 


450 A. HESSELBO 


50. Distichium montanum (Lam.) Hagen. 
Distichium capillaceum (Sw.) Br. eur.; Swarlzia montana (Lam.) Lindb. 


Very common all over Iceland. 

Distichium montanum is one of the most widely distributed mosses 
in Iceland. It is found everywhere on almost every kind of substra- 
tum, with the exception of that which is very wet, from the bottom 
of the valley up to a height of about 500—600 metres above sea-level. 
At greater altitudes it is more rare, but may nevertheless be met with 
as far upwards as to the snow-limit, in sheltered localities among stones. 

This species has its main distribution in the more low-lying regions 
where, either intermixed with other species or in unmixed tufts, it is 
hardly absent from any cleft or soil-filled rock-crevice, but it also 
occurs everywhere intermixed in the moss-covering at the bottom of 
grass-fields, in bogs, on rocks, etc. 

The fruit, which is almost always present, ripens in the latter 
half of July. 


51. Distichium inclinatum (Ehrh.) Br. eur. 
Swartzia inclinata Ehrh. 


E. Iceland: Vallanes!. N. Iceland: Near Dettifoss!; Husavik!; Hof 
near Eyjafjordur (O. D.)!; Oxnadalur!; Vidimyri!. NW. Iceland: Gnups- 
dalur in Dyrafjérdur!. 

It was also collected by Steenstrup, but no habitat is given. 
Gronlund records this species from Hafnarfjordur, but the specimens 
in the Botanieal Museum belong to Distichium montanum. 

This plant was only found in the northern half of Iceland, where 
it occurred in many places from Vallanes in the east to Dyrafjordur in 
the north-west, and especially in North Iceland proper. It grows as a 
rule on damp ground, although not very abundantly, for instance 
along streams, associated with Dichodontium, Didymodon rubellus, Hepa- 
ticæ, etc. Near Vidimyri it was growing along the banks of streams, 
mixed with Scapania curta, Lophozia Miilleri, Haplozia atrovirens, Di- 
chodontium and Bryum pallens; near Dettifoss on glacier-sand, in com- 
pany with Ceratodon purpureus and Aongstræmia longipes and near 
Husavik on damp sandy soil, with Åongstræmia, Dichodontium and 
Philonotis tomentella. Im several places it was collected on peaty soil 
in association with Meesia trichoides, Catoscopium nigritum, Barbula ru- 
bella, Aongstræmia, Lophozia Wenzelii, etc. 

In North Iceland proper the fruit was almost ripe at the end 
of July. ; 


52. Bryoxiphium norvegicum (Brid.) Mitten. 
Eustichium norvegicum Br. eur. 


S. Iceland: Krisuvik (Mérch)!; Thingvellir (Mérch)!; glacier origi- 
nating from the Jåkul (Stp.)!; Kolvidarhol!; Flokastadagil!; Barkarstadr!; 
Drångshlid!. 

This species was only found in the south-western part of the 


THE BRYOPHYTA OF ICELAND 451 


country, where it is common in many places and often occurs in 
enormous quantities. Mørch records that it grows in holes in the 
lava. Near Kolvidarhol it was found everywhere on tuff, from the 
farm (at about 250 metres) up to about 450 metres above sea-level; it 
was found in extensive mats, especially on the vertical or overhanging 
sides of the tumbled-down blocks, and in. the interior of holes and 
clefts. It was most frequent on Fljåtshlid and south of Eyjafjall. 
There it was growing in many places on the vertical sides of tuff-rocks 
in the ravines and on the fallen blocks, for instance in Flokastadagil 
near Breidabolstadr it covered long stretches of the perpendicular sides 
of a ravine, from a distance of a few cm. above the water-level of the 
river upwards, with a dark-green closely adhering mat in which almost 
no other mosses were intermixed. In this part of the country it has 
only been met with up to about 100 metres above sea-level. Only ste- 
rile specimens have been found. 

Bryoxiphium norvegicum is the only Iceland moss which does not 
occur on the European continent or in the British Isles. Outside Ice- 
land it has only been known to occur in a few localities in North 
America (Ohio, Kentucky and Wisconsin). 


Fam POTTIAGEÆ.: 


53. Pottia Heimii (Hedw.) Br. eur. 


Iceland (Morch)! 

E. Iceland: . Hornafjérdur!. SN. Iceland: near Eyjafjordur (F.)!; 
Saudanes. (St.)!:"Grimsey (0. D:.)!. . NW. Iceland: Årmuli!. W. Iceland: 
Reykjavik (Grl.;!); Borgarnes!. S. Iceland: Vestmannaey'!. 

This species grows on sandy or muddy soil near the sea, and is 
probably commonly distributed along the whole coast in such localities. 
Around Reykjavik, on Vestmannaey and near Hornafjérdur it was com- 
mon, and was frequently found also on soil-covered rocks and in rock- 
cleftskeloseltolther sea 

The fruit ripens about July ist; on Grimsey the fruit was fully 
ripe on July 14th, but the lids had not been thrown off. Near Årmuli 
it was hardly ripe during the first days of July. 


54. Pottia latifolia (Schwågr.) C. M. 


N. Iceland: Hof near Eyjafjordur (0. D.)!: Ås (0. D.)!, with ripe 
capsules on June 17th. ; 

Note.  Pottia truncatula and P., lanceolata are enumerated in older 
lists as found in Iceland, but no specimens of these species are to be 
found in the collections, and their occurrence in Iceland is very im- 
probable. 

55. Didomodon rubellus (Hoffm). Br. eur. 


Barbula rubella (Hoffm.) Mitten. 
Very common everywhere on a more or a less damp substratum, 
in rock-clefts, on the ground, on rocks and gravelly soil by rivers, in 
bogs, etc. It grows sometimes in unmixed tufts and sometimes inter- 


452 A. HESSELBO 


mixed with other mosses, and as a rule sets fruit, which ripens late in 
August or early in September. It is most frequent in the low land up 
to about 300 metres above sea-level, but is also met with, although 
scantily, and often sterile, as far upwards as to the mountain heights, 
for instance on Isafjardarheidi, about 600 metres above sea-level. 

Forms which must be referred to var. brevifolia Lindb. et Arnell 
occur commonly together with transitional forms to the type. Near 
Seydisfjordur the variety was found abundantly in reddish brown, dense 
tufts, about 5 cm. high, in rock-fissures filled with soil. 

Note. Didymodon tophaceus is recorded by Lindsay from Ice- 
land, but no specimens are in the collections and the record is pro- 
bably due to an erroneous determination. 


56. Didymodon rufus Lorentz. 


E. Iceland: Stafafell!:; Geithellir!. N. Iceland: Stadr near Hrutafjér- 
dur!; Akureyri, at an altitude of 900 metres!. W. Iceland: Lundur! 
Breidabolstadr in Reykholtdalur'!. 

The plant, which was only found sterile, grew in all the above 
localities in loose tufts, a few cm. high, on damp gravelly ground, as 
a rule rather scantily. Near Akureyri it grew scantily among Sphæro- 
cephalus turgidus and Rhacomitrium hypnoides. 


57. Leptodontium flexifolium (Dicks.) Hampe. 


N. Iceland: Grimsey (0. D.)! (f. compacta). 
Enumerated in Lindsay's list, but neither the name of the finder 
nor the habitat is given. 


58... Trichostomum littorale Mitten. 
Mollia littoralis (Mitten) Braithw. 

S. Iceland: Helgafell on Vestmannaey, on blocks of lava!; Hafnar- 
fjordur, in a lava cave!; on warm clayey flats near Reykjanes light- 
house (Ostf.)!, associated with Preissia, Riccia and Archidium. 

59. Tortella inclinata (Hedw.) Limpr. 
Mollia inclinata (Hedw.) Lindb. 
E. Iceland: Vallanes (H. J.)!. 


60. Tortella tortuosa (L.) Limpr. 
Mollia tortuosa (L.) Schranck. 


Very common all over Iceland. 

This species has its main distribution in the low land up to about 
400 metres above sea-level. It appears to be rare everywhere at higher 
levels, and hardly ascends higher than about 600 metres. " It grows 


THE BRYOPHYTA OF ICELAND 453 


both on rocks and on earth. In lava-fields it is one of the most com- 
monly occurring species, and grows there in enormous cushions on 
rock-sides in caves and clefts, in association with Amphidium Mougottii, 
Ånæctangium compactum and Grimmia torqualta. On rocks it grows 
on perpendicular faces, on soil-covered ledges, and in clefts. On the 
ground it is found especially where it is not too damp, for instance in 
grass-fields and on the top of knolls in bogs, usually intermixed in tufts 
of other mosses. At higher altitudes it rarely grows on rocks, but as 
a rule on gravelly ground or in bogs. Only sterile specimens have been 
found. 

This species varies considerably in the size and length of the 
leaves... Forms from shady and protected localities, especially from 
lava-clefts, form very tall and loose tufts with exceedingly long and 
strongly crisped leaves. 


61. Tortella fragilis (Drumm.) Limpr. 
Mollia fragilis (Drumm.) Lindb. 


Commonly distributed all over Iceland. 

This species grows in localities similar to those of T. fortuosa, and 
often in company with the latter, but as a rule rather scantily. It 
usually occurs in unmixed tufts on soil-covered rocks and on some- 
what damp ground, but only sterile specimens have been found. Åround 
Reykjavik, for instance, it was common on knolls in boggy tracts which 
had been partially drained. In NW. Iceland it is rather rare, and has 
only been found in a few localities near Kaldalon. 


62. Barbula unguiculata (Huds.) Hedw. 
var. cuspidata (Schultz) Braithw. 


S. Iceland: Vestmannaey!, on damp gravelly soil near the town. It 
has previously been collected here by an unknown finder (H. Jonsson 
believes by G. Brynjolfsen). W. Iceland: Saurbær near Hvalfjérdur, 
at an altitude of about 100 metres, on soil-covered rocks. In both 
places only sterile specimens have been found. 


63. Barbula fallax (Hedw.). 


S. Iceland: Reykir g. W. Iceland: Saudlauksdalur (H. J.). 

Var. lævifolia n. var. 

In wide dense cushions, 2—3 inches deep, rusty brown in the in- 
terior, brownish green at the top. The lower leaves as in the type, the 
uppermost shortly pointed with rounded, broad and flat apex, into 
which the strong nerve runs out, widely revolute to near apex, with 
two deep plicæ at the base. Leaf-cells in form and size as in B. fallaæ, 
in the lower half of the uppermost leaves, however, more thin-walled, 
shortly rectangular; all quite smooth or, more rarely, indistinctly papil- 
lose. The innermost perichætial leaves ovate or widely lanceolate, with 
faint or indistinct nerves. 


454 A. HESSELBO 


S. Iceland: Thorlåkshver, on warm clayey ground. 

Although this form differs somewhat from B. fallax in habit, in the 
smooth leaves with rounded apex, and in the divergent form of the 
perichætial leaves, yet there is hardly any doubt that it must be re- 
ferred to this species. Setting apart the absence of papillæ the leaf- 
tissue is quite typical, and the form and the plication of the lower 
leaves are exactly as in B.fallax. It should be remembered that the 
occurrence on a warm substratum produces very divergent forms also " 
in other mosses, e. g. Philonotis, Catharinea and Oligotrichum. Near 
Reykir, B.fallax was collected both on stones by the river, in sand- 


Ni 


a a b b c c c 





Fig. 1. Barbula fallax Hedw. a, Leaves of the plant from Reykir (X 20). b, Leaves of 
var. lævifolia; c, perichætial leaves of same. 


filled tufts about 2 cm. high, in a somewhat typical form, and on a warm, 
damp substratum. Plants from the latter locality form a transitional 
stage to var. lævifolia. It grew in somewhat loose tufts, 3—4 cm. high, 
which in colour and appearance agreed with var. /ævifolia. The lower- 
most leaves are pointed, the uppermost rounded; the perichætial leaves 
are lanceolate, rounded, with rather faint nerve. The leaf-cells are di- 
stinctly papillose, sometimes as highly as in the typical form, some- 
times much more faintly. 


64. Barbula cylindrica (Tayl.) Schimp. 


N. Iceland: Hof near Eyjafjordur (O. D.)!. S. Iceland: Vestmannaey!; 
Ytri Skogur (H. J.)!; Fljétshlid!; Holt!; Drångshlid!; Hornafjérdur!. 
W.-Iceland: Esja near Kollafjérdur!. 

This species occurred commonly, and often abundantly, on tuffrocks 
in S. Iceland and on Vestmannaey. There it also grew abundantly on 
sandy soil and on lava blocks near the sea. In the other localities it 
was found but scantily. Fruit was collected only near Holt, where, at 
the end of July, both old and yet green capsules were collected from 
the same tuft. 


THE BRYOPHYTA OF ICELAND 455 


65. Barbula icmadophila Schimp. 


SE. Iceland: Hornafjérdur!, in several places in small quantities. 
N. Iceland: Skagafjordur (Grl.)!; Hofsfjall near Eyjafjordur (0. D.)!. 
W. Iceland: Esja, at an altitude of 400 metres!. 

This species has been collected everywhere only in small quantities, 
in cushions 1—2 cm. deep, on humus-covered rocks. 


66. Desmatodon latifolius (Hedw.) Br. eur. 
Tortula latifolia (Hedw.) Lindb. 


E. Iceland: Djupivogur!, common near the sea; Vallanes!; Hamar- 
fjordur!; Berufjérdur!. N. Iceland: SS at an altitude of about 
400 .metres; Reykjahlid!; Grimsey (0. D.)!; Stadr near Hrutafjérdur !. 
NW. Iceland: Isafjordur!; Grunnavik!; Hesteyri!: Årngerdaeyri!; W. Ice- 
land : Stykkisholmur!. 

Var. muticus Brid. 

Grimsey (O. D.)!; Reykjahlid!; Isafjérdur!, 450 metres above sea- 
level. Found in all the localities together with the type. 

In NW. Iceland D. latifolius is one of the most frequently occur- 
ring species, and the same appears to be the case in E. Iceland. It 
occurs especially near the sea on sandy soil or, also, on humus-covered 
rocks, and often abundantly. Near Vallanes it grew in masses on peat 
walls and on dikes around the farm, associated with Encalypta rhab- 
docarpa, Tortula subulata, Ceratodon purpureus, Bryum argenteum, Bar- 
tramia ityphylla, etc. 

In N. Iceland this species occurs more sporadically. Near Husavik 
only a single little tuft was found on the top of Husavikurfjall; near 
Reykjahlid it was common in lava-fields, in places covered by a thin 
humus-layer, associated with Ceralodon purpureus, Tortula ruralis, T. su- 
bulata, etc., and near Stadr it grew on the peat walls of the farm ex- 
actly as it grew near Vallanes. From SW. and S. Iceland this species 
appears to be quite absent, and in W. Iceland it was only found near 
Stykkisholmur, where it was found abundantly on earth and rocks along 
the shore. 

It is only by exception that Desmatodon latifolius has been found 
at higher altitudes. Near Isafjordur it occurred here and there on the 
rocky flat up to a height of about 500 metres, for instance on a stony 
slope 450 metres above sea-level, associated with Bryum elegans, Bar- 
tramia ityphylla, Pohlia commutata, Scapania curta and Encalypta rhab- 
docarpa. The fruit, which was present everywhere in abundance, ri- 
pens at the end of June. 


67. Desmatodon cernuus (Huibn.) Br. eur. 
Tortula cernua (Hubn.) Lindb. 


N: Iceland :. Hof. (0. D:)!: Hjalteyri. (0. D.:)!. 
On the plant from HJjalteyri the fruit had just ripened on Sep- 
tember 5711897: 


456 A. HESSELBO 


68. Tortula obtusifolia Schleieh. 


S. Iceland: Flokastagil on Fljåtshlid!; Drångshlid!. 

This very rare species was found in both places in very small 
cushions, only 5—6 mm. deep, with numerous capsules which were al- 
most fully ripe in the latter half of June. In Flokastadagil it grew 
here and there on fallen blocks, especially in small fissures on the 
vertical rock-faces.. Near Drångshlid it was found in small quantity on 
the dry faces of tuffrocks in company with Tortula muralis, Grimmia 
Doniana and Orthotrichum anomalum. 


69. Tortula muralis (L.) Hedw. 


S. Iceland: Vestmannaey!; Drångshlid !. 

The plant was found very scantily in both places on dry faces of 
tuffrocks. On Vestmannaey it grew on a rock-face with a southern ex- 
posure, associated with Grimmia marilima and Barbula cylindrica; near 
Drångskhlid it grew in association with Grimmia Doniana and Tortula 
obtusifolia. 


70. Tortula subulata (L.) Hedw. 


Commonly distributed over the whole of Iceland. 

The plant grows almost exclusively in the low land up to about 
300 metres above sea-level, and only in quite a few localities in S. Ice- 
land has it been found at a higher level, for instance near Barkastadr 
at an altitude of 410 metres. It is most frequent in S. and E. Iceland; 
in N. and NW. Iceland it is somewhat less common, but has never- 
theless been collected from a great number of places. It usually grows 
on a rather dry substratum, e. g. humus-covered rocks, in rock-clefts, on 
dikes and on peat walls of houses, mostly associated with Bartramia 
ityphylla, Pohlia cruda, Encalypta rhabdocarpa, Brachythecium albicans 
and other species. Fruit, which is always present, ripens in the end of 
June or in the beginning of July. 


71. Tortula mucronifolia Schwgr. 


Vestmannaey !. 

It occurs here in localities quite similar to those of Tortula subu- 
lata, for instance on dikes, in sandy soil and on humus-covered rocks, 
and appears to be as frequent as this. The fruit was not yet quite 
ripe towards the middle of June. 


72. Tortula ruralis (L.) Ehrh. 


Commonly distributed over the whole of Iceland. 

Tortula ruralis is a decidedly xerophilous species, and has, there- 
fore, its widest distribution in the driest parts of the country. In the 
lava-fields in the interior, for instance around Myvatn, it occurs abun- 
dantly; in NW. Iceland it is also more frequent than in the other parts 


THE BRYOPHYTA OF ICELAND 457 


of the country, especially E. Iceland, where it occurs but scantily in 
many districts. 

It grows in sandy soil, upon soil-covered rocks and at the base of 
vertical rock-faces, and is most frequent in the low land. Near Akur- 
eyri it was found with ripe fruit (20. 7. 1909). Near Grunnavik. in 
Jokulsfjordur it was found in fruit at an altitude of 270 metres. In 
Esja it has been collected up to about 420 metres above sea-level. Near 
Vallanes the fruit was ripe about July ist. At the coast, near Reykja- 
vik, it grew abundantly in large tufts which were quite filled with sand. 


73. Tortula aciphylla (Br. eur.) Hartm. 
T. norvegica (Web.) Wahlb. 


NW. Iceland: Dsrafjérdur!, at an altitude of 150 metres. 

It grew there in cushions, about 2 cm. deep, on humus-covered 
rocks. The capsules were quite green on June 17th. 

The leaf-form, nerve and leaf-hair in these specimens were quite 
typical, but the cells of the leaf-base were occasionally chlorophyllose 
towards the margin, forming a more or less distinct border, in which 
feature it approached Tortula ruralis. 


Fam. GRIMMIACEÆ. 
74. Schistidium maritimum (Turn.) Br. eur. 


E. Iceland: Hornafjordur!; Hof!; Hamarfjordur!; Djupivogur!. N. Ice- 
land: Husavik!; Grimsey (0. D.)!. NW. Iceland: Common everywhere 
along the coasts!. W. Iceland: Stykkisholmur; Hafnarfjérdur!; Reykjavik 
(Grl.;!); by Hvalfjordur (Grl.)!. S. Iceland: Vestmannaey!. 

This plant doubtless occurs everywhere along rocky shores. It is, 
however, absent from, or only occurs scantily in, the interior of deep, 
narrow fjords. Thus, around Berufjérdur it is very common towards 
the entrance of the fjord, near Djupivogur, while it is entirely absent 
from the head of the fjord. 

Ås a rule it grows only on rocks quite close to the sea, and up 
to a height of some 10 metres. More rarely, for instance near Djupi- 
vogur and on Vestmannaey, it descends to the sandy ground at the 
foot of cliffs. 

Fruit is almost always present abundantly. 


75. Schistidium apocarpum Br. eur. 
Grimmia apocarpa Hedw. 


Subsp. vwlgare (Chal.) Loeske. 


Common over the whole of Iceland. 
It grows, as a rule, on rather dry rocks, occasionally also in gra- 
velly soil, but then rarely in any great abundance. It is most frequent 
The Botany of Iceland. Vol. I, part II. 30 


458 A. HESSELBO 


in the more low-lying parts of the country, but may, however, occasion- 
ally be met with upwards of 600 metres above sea-level. 


76. Schistidium apocarpum Br. eur. 
Subsp. gracile (Schwågr.) Loeske. 


Very common. 

One of the most frequent mosses, and met with from the lowlands 
up to mountain heights. It occurs, in numerous forms, both on dry 
and on damp rocks, and on more or less damp gravelly soil. On gra- 
velly flats, especially above an altitude of about 300 metres, it often 
forms extensive growths. It- occurs also very abundantly on tuff-rocks 
(for instance in Esja and below Eyjafjall) in a low, reddish-brown 
form. Fruit is always present. 


77. Schistidium apocarpum Br. eur. 


Subsp. confertum (Funck) Dixon. 


E. Iceland: Hof!; Seydisfjérdur!. N. Iceland: Husavik, at an alti- 
tude of about 400 metres!; Reykjahlid!; between Hnausar and Lækjar- 
mot!; Vidvik (P. Såféniasson)!, several places near Eyjafjordur (0. D.)!. 

This form has its main distribution in N. Iceland, where it is 
rather common; thus, near Reykjahlid it was found abundantly on lava- 
blocks. It grows by preference on rather dry rock-faces, or upon 
stones, and has been gathered everywhere in fruit. 


78. Schistidium alpicola (Sw.) Limpr. 
Var. rivularis (Brid.) Wahlb. 


Very common in E., N. and W. Iceland, more rare in S. Iceland, 
where it is absent from many localities. 

This species grows on stones and rock surfaces in rivers, often in 
great abundance, covering the banks and the stones in the water for 
long distances. It does not appear to ascend much higher than about 
300—400 metres above sea-level. 

The chiéf reason for its occurring more scantily in South Iceland 
must be sought in the fact that the water in many rivers is turbid, 
owing to the glacier-clay carried in them. In this part of Iceland it is 
met with especially in rivers with clear water, at an altitude of 200— 
400 metres, more rarely in the low land. 

Var. eualpicola Loeske. Grimmia alpicola Swartz. 

NæÆLceland:EDetifoss ef 

This form, which in the above locality grew on blocks of basalt 
at some distance from the waterfall, but nevertheless exposed to the 
spray, appears to be as rare in Iceland .as in most of the rest of 
Europe. Forms, intermediate between this form and var. rivularis, have 
not been collected. 


THE BRYOPHYTA OF ICELAND 459 


79. Grimmia commutata Hihbener. 


Grimmia ovalis (Hedw.) Lindb. 


N. Iceland: Myvatnshraun (St.)!.  W. Iceland: Budir (H.J.)!. In the 
supplement to "Isl. Kryptogamflora” (B. T., vol. 20, p. 105) Grønlund 
has recorded G. ovata from Myvatnshraun and G. commutala from Val- 
lanes and Kolfreyjustadr. But according to the specimens in the Bota- 
nical Museum in Copenhagen, determined by C. Jensen, the plant from 
Myvatn is Grimmia commutata and those from the two other localities 
are G. ovata. Grønlund has, therefore, probably confused the two 
names (G. ovata and G. ovalis) used by C. Jensen. 


80. Grimmia Doniana Smith. 


E. Iceland: Common from Hornafjéordur to Seydisfjordur!. N. Iceland: 
Hålar (Grl.)!; Vidimyri (Grl.)!.. NW. Iceland: Dyrafjérdur!; Gnupsdalur!; 
Dynjandi!. Common in SW. and $S. Iceland. 

With the exception of N. Iceland, where it appears to be rare, this 
species is widely distributed in the more low-lying districts of the 
country, as it is only by exception that it ascends higher than 200— 
300 metres. Near Holt in S. Iceland it has, however, been collected 
plentifully at about 500 metres above sea-level. It usually grows in 
small, round tufts on loose blocks, more rarely in wide cushions, 
irregular in form. In South Iceland proper — for instance near Drångs- 
hlid — it occurs also on faces of tuff-rock. The fruit, which occurs 
everywhere, was hardly ripe even at the end of July. 


81. Grimmia alpestris Nees. 


E. Iceland: Arnkelsgerdi; Eidar (H. J.)!, sterile. S. Iceland: Hafurs- 
holt!, fr. 


82. Grimmia ovata W. et M. 


Grimmia ovalis Lindb. 


Iceland leg Mårch!. 'E. Iceland: Kolfreyjustadur (H. J.)!; Vallanes 
(H. J.)!; Seydisfjordur!; Berufjérdur!. N. Iceland: Hnausar in Vatnsdalur'!: 
Reykjaheidi!; Herjardalsheidi (Grl.)!. Only sterile specimens have been 
found. 

This species grows on dry stones and rocks; it is rather frequent 
in East and North Iceland, but has not been found in the other parts 
of the country. 


83. Grimmia incurva Schwgr. 


E. Iceland: Vallanes!. 
It was growing in the above locality on a slightly inclined, partly 
inundated rock-surface in the river Lågarfljot. 
30% 


460 A. HESSELBO 


84. Grimmia patens (Dicks.) Br. eur. 
Dryptodon patens Brid., Limpr. 


E. Iceland: Hornafjérdur!. W. Iceland: Budahraun (H. J.)!: Hafnar- 
fjordur!. 

In Budahraun it was growing in the clefts in the lava-field; near 
Hafnarfjordur it was found in abundance associated with Orthotrichum 
rupestre and O. Sturmii on the, face of a dolerite rock with a nor- 
thern exposure. 


855. Grimmia funalis (Schwgr.) Sch. 


Widely distributed over the whole of Iceland. 

G. funalis is by far the most frequent species of this genus. It 
grows everywhere on dry rocks, especially on the vertical sides of ra- 
vines and lava-clefts, where it forms immense cushions, often 5—6 cm. 
deep, which very easily break up into pieces when loosened from the 
substratum. The fruit, which occurs rather frequently, was not ripe 
even in August. This species is most frequent up to about 300—400 
metres above sea-level, but often ascends, however, to somewhat above 
500 metres, for instance near Akureyri. 


86. Grimmia torquata Hornsch. 


E. Iceland: Hornafjérdur!, very common; Seydisfjordur!.  N. Ice- 
land: near Hrutafjordur (H. J.)!. NW. Iceland: Dynjandi!; Grunnavik!; 
Arngerdareyri!. W. Iceland: Melar (Grl.)!; Hafnarfjordur (Grl.;!); Buda- 
hraun (HJ) "Very common"in "SW Iceland! 

Fhistspeciestinkvery common kink SEES sFandtSsSyværcelanderather 
rare in N. and NW. Iceland. It grows on dry rock-faces in ravines 
and lava-clefts, but only sterile specimens have been found. 

Note. Grimmia pulvinala is enumerated in several older lists, but 
these records are probably due to a confusion with other species. 


87. Rhacomitrium sudeticum (Funck) Br. eur. 
Grimmia microcarpa (Gmel.) Lindb. 


Common over the whole of Iceland. 

In many parts of Iceland R. sudeticum is one of the most frequent 
species, being found from the sea-level up to the gravelly flats of 
mountain heights. It is most widely distributed in NW. Iceland and 
along several fjords in E. Iceland, for instance Seydisfjordur, while it 
appears to occur less abundantly in N. Iceland. It grows on rather dry 
rocks and blocks of lava, in clefts, and on wet stones in and by streams. 
In mountain heights, especially in NW. Iceland, it forms the bulk of the 
vegetation on gravelly flats from about 300 metres upwards. 

It varies considerably in size, habit, colour and length of hair 
point. On wet rocks it becomes very large, and. almost black, with 
very short hair point, while the forms growing on dry rocks are as a 
rule very low in growth and slender, often occurring in small round, 
compact tufts, blackish green or brownish red in colour, with longer 


THE BRYOPHYTA OF ICELAND 461 


hair point. In the birch coppices of NW. Iceland a form frequently 
occurs on larger, scattered stones which, in habit, reminds one of 
R. fasciculare. The tufts are flat towards the edge, dark-green; the hair 
point is very short and often wanting. The forms from mountain 
heights are often jet-black (var. atrata) or — on damp gravelly flats — 
yellowish green at the top, and grow in extensive, discontinuous mats, 
a few cm. thick. The fruit, which is very common, ripens in the be- 
ginning of June. 


88. Rhacomitrium heterostichum (Hedw.) Brid. 


E. Iceland: Hornafjérdur!; Hof!; Geithellir!; Berufjérdur!. UN. Ice- 
land: Akureyri!, at an altitude of 350 metres. NW. Iceland: Patreks- 
fjordur!; Arngerdareyri!. Common in S. and W. Iceland!. 

Common throughout E. Iceland, from Berufjérdur, and southwards 
through the whole of S. and W. Iceland. 

In W. Iceland it occurs abundantly at the head of Isafjéårdur, near 
Arngerdareyri, but has not been collected about the fjord further out- 
wards towards its entrance; it was also common by Patreksfjordur. It 
appears to be more rare in N. Iceland. 

The plant grows most frequently on dry rocks, and more rarely 
on those that are damp; it occurs in masses, for instance, in the lava 
districts of SW. Iceland. It is most common in the low land, but is, 
however, also met with at about 600 metres above sea-level. Fruit 
is rather common. 


89. Rhacomitrium microcarpum Brid. 
Grimmia ramulosa Lindb. 


N. Iceland: Hof by Eyjafjordur (O. D.)!, sterile. 

Recorded by Grånlund from Reykjavik and Husafell, but the 
specimens in the Botanical Museum in Copenhagen must be referred to 
R. canescens. 


90. Rhacomitrium canescens (Weis) Brid. 


Very common. 

Next to R. hypnoides the most widely distributed species, and met 
with as frequently on mountain heights as in the low land, and in 
numerous forms. It grows both on rocks and on earth, especially 
where it is somewhat damp. In the ”"Grimmia heath” it often covers 
the more low-lying tracts with its light yellowish-green to greyish-green 
mats, while R. hypnoides grows in the more dry parts, but both species 
often occur mixed. It is also the most abundant constituent of the vege- 
tation on damp sandy or gravelly soil by streams, in grass-fields with 
a poor soil, etc. Forms with the hair-point entirely or almost wanting 
(f. epilosa) are frequent in wet localities. 

Fruit is only rarely ånd scantily met with. Near Hof in E. Ice- 
land plants were found in the middle of June with old capsules, and 
ncar Sugandafjoérdur about July i1st with yet green capsules. 


462 A. HESSELBO 


91. Rhacomitrium hypnoides (L.) Lindb. 
Grimmia hypnoides Lindb.; Rhacomitrium lanuginosum (Ehrh.) Brid. 


Very common over the whole of Iceland. 

Rhacomitrium hypnoides is by far the most common plant in Ice- 
land. In all parts of the country, both along the coasts and on the 
mountain heights, it covers, almost without any mixture of other plants, 


vast tracts of land — the Grimmia heaths so well-known also from 
other Arctic countries — with its thick greyish-white mats. It also 


occurs in almost every locality where it is not too damp, and very 
often intermixed in tufts of other mosses. Thus, it is very common on 
rocks and stones, and often covers large tracts of lava-fields with 
cushions one foot deep. In bogs it grows, as a rule, on the top of the 
knolls, in company with species of Hylocomium, and in birch cop- 
pices it forms, in association with Hylocomium spp, Hypnum uncinatum 
and several other species, the moss carpet of the floor. Only sterile 
specimens have been found. 


92. Rhacomitrium fasciculare (Schrad.) Brid. 


Very common in the Southern, Western and Eastern parts of Ice- 
land. In N. and NW. Iceland it appears to be somewhat less plentiful, 
but nevertheless it must also there he regarded as one of the more 
commonly occurring species. 

It occurs at all altitudes up to the limit of vegetation on mountain 
heights. It forms, there, like R. sudeticum, black Alpine forms on 
dry rocks. 

This species usually grows on stones and rocks in dry situations, 
but it also ocecurs frequently in gravelly soil.  Occasionally it may 
even be found in the lowermost portion of birch-trunks. The fruit, 
which is rather common, ripens in the lowlands in the first half of June. 


93. Rhacomitrium aciculare (L.) Brid. 


Very common on wet rocks in and by rivers and waterfalls every- 
where in the low-lying parts of the country. It does not appear to 
ascend much higher than 300—400 metres. The fruit, which occurs 
everywhere, ripens, as a rule, in the beginning of June. 


Note. Grimmia (Rhacomitrium) elliptica is recorded in older lists 
as found in Iceland by Mérch. The specimens in the collections belong 
to R. fasciculare. 

Cinclidotus fontinaloides, also, is recorded by Hornemann as 
found in Iceland, but it was undoubtedly confused with another specics. 
Specimens are not to be found in the collections. 


94. Hedwigia albicans (Web.) Lindb. 
Hedwigia ciliata Ehrh. 


SW. Iceland: Hafnarfjérdur (Grl.)!, sterile. 


THE BRYOPHYTA OF ICELAND 463 


Fam. ORTHOTRICHACEÆ. 
95. Amphidium lapponicum (Hedw.) Schimp. 
Ånæctangium lapponicum Hedw. 


Very common. 

This species usually grows on damp rocks, especially in clefts and 
crevices, but may also occur on a more dry substratum, for instance in 
lava caves and among stones on heaps of debris at foot of cliffs (Urd). 
It grows both on bare rocks and on humus in rock-clefts,. occasionally 
also in gravelly soil or on earth, and even upon the sides of knolls in 
bogs. It does not appear to ascend to any great height on the moun- 
tains; it is most widely distributed up to 200—300 metres above sea- 
level, but it has, however, been found, although scantily, up to about 
500 metres above sea-level. 

The fruit was usually ripe in the first half of June. 


96. Amphidium Mougeotii (Br. eur.) Schimp. 
Anæctangium Mougeotii Lindb. 


Iceland (Mørch; Stp.)!. E. Iceland: Seydisfjordur!; Stéd (H. I)!; 
Hof near Eyjafjordur (O. D.!); Hrutafjérdur (St.)!; Myvatn (Grl.)!. NW. Ice- 
land: Kaldalon!; Bæir!; Arngerdareyri!; Dynjandi!. West, South-west and 
South Iceland: common!. 

Common all over the southern and western part of the country 
and probably also in NW. Iceland, certainly everywhere north of Isa- 
fjordur. In N. Iceland it appears to be rarer. but occurs abundantly 
near Eyjafjordur. In E Iceland it is common everywhere around Sey- 
disfjordur and is probably also frequent elsewhere. It usually grows 
in large, thick cushions on damp rock-sides in clefts, and by waterfalls, 
hut is also one of the species most commonly found in lava-caves. 

This species rarely ascends to any great height on the mountains. 
In Esja it occurred 400 metres above sea-levei, and has nowhere been 
collected at a greater altitude. Only sterile specimens have been found. 


97. Ulota phyllantha Brid. 
Ulota maritima C. M. et Kindb. 
Weissia maritima Britt. 


E. Iceland: Everywhere along the coast from Hornafjøårdur to. Be- 
rufjordur!. NW. Iceland: Frequent around Isafjérdur!. W. Iceland: Snæ- 
fellsnes (H. J.)!; Stykkisholmur!; common near Reykjavik, Hafnarfjéordur 
and everywhere around Hvalfjérdur!. S. Iceland: Vestmannaey!. 

This species is probably common everywhere along the coasts, 
where it grows in company with Grimmia maritima quite close to the 
sea-side. It is only rarely met with at a distance from the sea, as, for 
instance, around Reykjavik, and in a few places in the lava-fields near 
Hafnarfjordur. The reason why it has not been collected in North 


464 A. HESSELBO 


Iceland is probably due to the fact that the coast there has been but 
little investigated. At Husavik the coast cliffs consist of easily weathered 
tuff which is very poor in mosses, the surface being too much in- 
clined to crumble away. Grimmia maritima occurred also very sparingly 
there, and Ulota was not found at all. Like Grimmia maritima it keeps 
chiefly to the shores of broad fjords; therefore it was not found, or was 
very rare, in the interior of the deep fjords of East and North-west 
Iceland, nor did these two species occur at Akureyri. On Vestmannaey 
Ulota phyllantha was the most commonly distributed moss; it occurred 
abundantly everywhere, reaching up the face of the cliffs and being 
also found on that side of the latter which faced the interior of the island. 
It often descended to the ground, it was thus found on the knolls in a 
wind-torn grass-field on the slopes of Helgafell associated with Rhaco- 
mitrium hypnoides and Hylocomium spp. 


98. Orthotrichum anomalum Hedw. 


S. Iceland: Vestmannaey!; Drångshlid (H. J.;!). 

On Vestmannaey it was rather frequent on tuffrocks at the foot of 
Stora Klit and in Heljusdal. The fruit was ripe in the first days of 
June. Near Drångshlid it was found on dry tuff-rocks, associated with 
O. rupestre, Grimmia Doniana and Homalothecium. The plants from this 
habitat form a transition to O. saxatile, as slender cilia, which easily 
fall off, are often found on them. The capsule is however 8-striate, 
and the diameter of the spores is 0,012—0,018 mm. as in O. anomalum. 


99. Orthotrichum saxatile Schimp. 


SE. Iceland: Hornafjérdur!, on rocks in company with O. Stlurmii. 
The fruit was ripe in the middle of June. 


100. Orthotrichum cupulatum Hoffm. 


S. Iceland: Vestmannaey'!, fr. 

It was growing in the above locality on fallen blocks of tuff at the 
foot of Stora Klit; in the first days of June it had capsules which had 
just ripened. 

In older lists this species as well as O. affine and O. lejocarpum is 
recorded from Iceland as collected by Mårch; the habitats are not 
given more particularly, and no-specimens of these species are to be 
found in the collections, so all the records are probably due to con- 
fusion with O. Blyttii. 


101. Crthotrichum rupestre Schleich. 


Common all over Iceland except in the north-west, where it has 
been observed only at the head of Isafjérdur near Arngerdareyri. 

It grows on dry rocks and detached blocks in the lowlands, and 
probably does not ascend higher than about 400 metres above sea- 
levels trhe fruitripenssinfthe first halfkor July: 


THE BRYOPHYTA OF ICELAND 465 


102. Orthotrichum Sturmii Hornsch. 


E. Iceland: Hof!; Hornafjérdur!. W. Iceland: Hafnarfjordur!; near 
Hvalfjordur'!. 

Growth similar to that of O: rupestre, and usually found in com- 
pany with it. Near Hafnarfjordur it was found abundantly on the dry 
northern face of a dolerite-rock. 

The plant from Hornafjérdur forms in regard to the eight coarse 
cilia a transition to 0. rupestre, but the upper half of the leaf consists 
of two layers of cells instead of only one in O. rupestre. 


103. Orthotrichum Killiasii C.M. 


E. Iceland: Seydisfjårdur!. N. Iceland: Åsbyrgi!. W. Iceland: Dala- 
sysla, Melar on a cliff inhabited by sea-fowl (?) (H. J.)!. 

The plant from Melar, which was quite sterile, was determined by 
C.Jensen, but owing to the absence of fruit the determination is 
doubtful. 

Near Seydisfjordur, where it grew on rocks close to the sea shore, 
the fruit was ripe about the ist of July. 


104. Orthotrichum Blyttii Schimp. 


Iceland (Stp.)!. E. Iceland: Djupivogur!; Seydisfjårdur!. N. Iceland: 
Geitaskard!; Vidimyri (Grl.;!). NW. Iceland: Arngerdareyri!. W. Iceland: 
Stykkisholmur!; Esja (Grl.)!. 

Grows on rocks, especially near the coast. Thus near Djupivogur 
Seydisfjordur, Arngerdareyri and Stykkisholmur it was found abundantly 
on the rocks of the coast. But it occurs also in the interior of the 
country; for instance, abundantly in several localities in Blånddalur and 
eastwards to Vidimyri. 

Gronlund's specimens are determined by Lindberg as Orthotri- 
chum arcticum, but on comparing numerous specimens af O. arcticum and 
O. Blyttii it has not been possible for me to find any real difference 
between these two forms. Hagen (Musci Norvegiae Borealis. p. 83) 
also regards O. arcticum as a variety of O. Blyttii, from which it is said 
to differ both by its shorter and broader leaves and by the shorter 
striæ of its capsule; but both these features vary so greatly in the 
Iceland plants, that there is hardly any basis for establishing variety, 
still less for separating them into two species. The perichætial leaves 
are usually about 2—5 times as long as they are broad, with margins 
which are revolute from a short distance below the apex down to the 
base. The capsule is usually striated along its whole length, rarely (as 
in the plant from Stykkisholmur) in the upper half only. The peristome- 
teeth are finely papillose, rarely marked with faint sinuose lines in 
their lower half. 


105. Orthotrichum lævigatum Zett. 


N. Iceland: Grimsey (0. D.)!: Akureyri, 350 metres above sea-level!; 
Vidimyri!. NW. Iceland: Arngerdareyri!. W. Iceland: Kalmanstunga (Grl.)!. 


466 A. HESSELBO 


Like O. Blyttii it grows on dry rocks and in many places in 
company with this. It has been everywhere collected in fruit. 


Fam. ENCALYPTACEÆ. 
106. Encalypta ciliata (Hedw.) Hoffm. 


Leersia laciniata Hedw. 


Iceland (Morch)!. E. Iceland: Hornafjérdur!; Djupivogur!; Hof!: Beru- 
fjérdur!.  N. Iceland: Ljosavatn!; Tverå in Oxnadalur!. —W. Iceland: 
Stykkisholmur!. Common in South and West Iceland (Grl. ;!). Vestmannaey !. 

This species has its widest distribution in the southern part of the 
country, where it is common from Berufjordur in East Iceland and 
thence southwards and westwards as far as Borgarfjordur. It has not 
been found in NW. Iceland and it is less frequent in North Iceland. 

It grows sometimes in small colonies on humus-covered rocks, 
sometimes in rock-fissures filled with soil, usually associated with Mnium 
orthorrhynchum, Anoectangium lapponicum, Bartramia ityphylla, Pohlia 
cruda, Distichium, Myurella spp. etc. It has its main distribution in 
the lowlands. In Esja it ascends on exception to about 400 metres 
above sea-level. . 

In E: Iceland the fruit. was quite unripe at the end of June; at 
Ljosavatn it had just ripened on July 19, but the calyptra still per- 
sisted. In S. Iceland, also, it ripens in the latter half of July. 


107. Encalypta rhabdocarpa Schwgr. 
Leersia rhabdocarpa Lindb. 


Common in localities similar to those of the preceding species; it 
often occurs abundantly also on the peat-walls of houses and on walls. 
It is met with both near the coast and in the interior of the country 
— for instance, abundantly at Reykjahlid — and is frequent as far up- 
wards as almost 500 metres above sea-level. 

BhesfruiteripenstastakruletatstherendkoRJuueebutkborbkoyerripe 
and just ripe capsules may frequently be found in the same tuft much 
later in the year. 

Var. pilifera (Funck) Br. germ. frequently occurs in company with 
the type. 


108. Encalypta contorta (Wulf.) Lindb. 
Leersia contorta Lindb. 


W. Iceland: Budahraun, on lava (H. J.)!, sterile. 

Note. Encalypta vulgaris is recorded by Zoéga and Morch, and 
E. commutata by Lindsay, but no specimens of either species occur in 
the collections. Georgia pellucida is recorded by Zoega and Vahl as 
found in Iceland, but of this species, also, no specimens are found in 
the collections, and all the above records are no doubt due to erroneous 
determinations. 


THE BRYOPHYTA OF ICELAND 467 


Fam. SPEACHNACEÆ: 
109. Dissodon splachnoides (Thunb.) Grev. 
Tayloria lingulata (Dicks.) Lindb. 


Iceland (Mérch)!. E. Iceland: Sevdisfjårdur. N. Iceland: Common 
everywhere!. NW. Iceland: Sugandafjérdur!; Isafjordur!; Armuli!; Arn- 
gerdareyri!. W. Iceland: Brattabrekka!. S. Iceland: Barkarstadr!, 550 
metres above sea-level. 

Grows on damp boggy ground, almost always mixed with other 
mosses such as Hypnaceæ, Cinclidium, Meesia trichoides, Dichodontium 
and Bryum ventricosum. In N. Iceland, where it is most widely distri- 
buted, it extends from the lowlands to a height of upwards of 600 
metres above sea-level. Thus, near Husavik, it was very common in 
Hypnum bogs, and near Akureyri it was found especially in bogs at an 
altitude of 500—600 metres. Near Seydisfjordur it was common from 
about 100 metres upwards to the mountain heights. From the south- 
western and southern part of the country it appears to be absent, or 
is at any rate very rare; it was found in this part of the country only 
in a Hypnum bog near Barkarstadr, at an altitude of 530 metres. 


110. Tetraplodon bryoides (Zoéga) Lindb. 
Tetraplodon mnioides (L. fil.) Br. eur. 


E. Iceland: Djupivogur!; Berufjérdur!. N. Iceland: Akureyri (Car- 
rington); Myvatn (Grl.)!; Reykir near Svinavatn (Grl.)!; Flateyardals- 
heidi (St.)!; near Eyjafjérdur (0. D.)!. NW. Iceland: Dynjandi!. W. Iceland: 
Husafell (Grl.)!; Brunnar (Grl.)!; Reynivellir (Grl.)!; Kolvidarhol!; Hafnar- 
fjordur!. S. Iceland: Draupahlid (Wiinsted)'!. 

Occurs here and there all over the country, but as a rule only in 
single tufts on the bodies of dead animals. Near Djupivogur a large 
tuft was found on a rotten fish. ' 

The fruit, which occurs abundantly everywhere, ripens at the end 
of June. 


111. Splachnum sphæricum (L. fil.) Swartz. 
Splachnum pedunculatum (Huds.) Lindb. 


Iceland (Morch)!; commonly distributed !. 

Grows on cow-dung in wet, marshy ground, often mixed with $. vas- 
culosum, but is less frequent than the latter. It occurs most frequently 
in North Iceland and also, to a certain extent, in East Iceland (Seydis- 
fjordur and Vallanes), where it is met with abundantly in many localities, 
it is somewhat rarer in South and South-west Iceland and probably 
does not ascend higher than 300—400 metres above sea-level. 

The fruit ripens rather irregularly, so that in the same tuft both 
ripe and also quite young capsules may he found. The majority of the 


468 A. HESSELBO 


fruits ripen during July; near Seydisfjordur many were found to be ripe 
on July ist, and in South Iceland ripe capsules were collected between 
10-15 June: 

Splachnum tenue is recorded by Gliemann to have been collected 
by Morch, but the specimens in the herbarium collected by Mårch 
belong partly to Splachnum sphæricum and partly to S$. vasculosum. 


112. Splachnum vasculosum L. 


Commonly distributed. 

Like the preceding species it grows, often abundantly, on cow-dung 
in wet marshy ground. It is most frequent in the lowlands, but ascends 
also as far upwards as to the mountain heights. Thus at Seydisfjardar- 
heidi it was common up to about 600 metres above sea-level, and in 
NW. Iceland also it was found as far upwards as about 500 metres. 
Such forms from mountain heights are as a rule low in growth and 
have short setæ. 

As is the case with Splachnum sphæricum the fruit ripens at difte- 
rent times during the. course of the summer, from the first half of June 
into August. 

Splachnum ampullaceum, as recorded to have been found by Koenig, 
is undoubtedly nothing else but $£. vasculosum. 


FUNARIACEÆ. 
113. Funaria hygrometrica L. 


Commonly distributed all over the country. 

It usually grows on peaty ground, associated with Dicranella. crispa, 
Polytrichum gracile, P. juniperinum, Bryum spp., etc., but is also found 
on humus-covered rocks and on tuff. It does not occur abundantly, as 
a rule, and it does not ascend to any great height, doubtless not higher 
than to an altitude of 300—400 metres. The fruit ripens in the be- 
ginning of August. The spores vary considerably in size in this species, 
from 0.011 to 0.021 mm., in plants which otherwise show no deviation 
from the type. 


114. Entosthodon ericetorum (Bals. et De Not.) Br. eur. 
Funaria obtusa (Dicks.) Lindb. 


W. Iceland: Kleppjarnsreykir in Reykholtdalur!; Englandshver!. In 
both places with ripe fruit (25—30 July). 

This species grew in the above localities on warm ground. Near 
Englandshver it grew along the outlet of a spring. Near Kleppjarns- 
reykir it was found both on a stone standing in water of a temperature 
of about 50”, mixed with Blindia acuta, AÅneura multifida, Anthoceros 
and Scapania irrigua, and also on warm, damp ground mixed with 
Fissidens osmundoides. 


THE BRYOPHYTA OF ICELAND 469 


Note. Entosthodon fascicularis is said to have been found by 
Hornemann near Reykjavik, but no specimens of it can be found in 
the collections, and the occurrence of this species in Iceland is not pro- 
bable. 


Fam. BRYACEÆ. 
115. Leptobryum pyriforme (L.) Schimp. 


Very common in the majority of the districts of Ieeland. This 
species, especially in N. and NE. Iceland, is common on peat or damp 
ground, but also on rocks, especially in soil-filled fissures. In S. Iceland 
it occurs abundantly on damp rock-sides, in the clefts of tuff rocks, and 
by rivers. It is only in NW. Iceland that it appears to be rarer. There 
it is common along the sea-side around Nauteyri and Arngerdareyri, 
but is otherwise rare in this part of the country and has only been 
found close to the coast. 

The fruit ripens late in summer. In N. Iceland only some of the 
fruits were ripe at the end of July. In specimens collected near Eyja- 
fjordur by 0. Davidsson in Sept.—Qct. the capsules were ripe, but the 
lids were not yet thrown off. 


116. Anomobryum filiforme (Dicks., Lindb.) Husnot. 


E. Iceland: Hornafjérdur!; Berufjérdur!; Stafafell!. "W. Iceland: Pa- 
treksfjårdur!, at an altitude of about 150 metres; Botnsdalur!; Esja, 
Mådruvellir', at an altitude of about 200 metres; Ålafoss!. S. Iceland: 
Fljøtshlid, common!; Eyjafjall, common!. 

Rather common in E. Iceland as far as Berufjordur, and in W. Ice- 
land: common in S. Iceland; not found in the northern part of the country. 
It grows, as a rule, scantily on damp rocks, mixed with other mosses, 
such as Blindia acuta, Ånæctangium compactum, Amphidium lapponicum, 
Aneura pinguis, etc., occasionally, however, also in large, pure cushions. 
In S. Iceland it occurred everywhere on the faces of damp tuff rocks, 
mixed with other mosses. Fruit was found only near Holt in S. Iceland. 


117. Anomobryum concinnum (Spr.) Lindb. 


S. Iceland: Hrutafell!. 

It was growing here rather scantily on the face of a damp tuff 
rock, mixed with Blindia acuta, Metzgeria furcata, Barbula rubella, B. cy- 
lindrica and Myurella julacea. 


118. Plagiobryum Zierii (Dicks.) Lindb. 


E. Iceland: Stafafell!; Berufjérdur!.  N. Iceland: Myvatn (Grl.)!. 
NW. Iceland: Arngerdareyri!; Patreksfjordur!. OW. Iceland: Nordredalur 
by Borgarfjérdur!; Esja, in several places!; Lundur!; Bardshellir (Grl.)!; 


470 A. HESS ELBO 


Hafnarfjordur (Grl.)!; Geithåls!.  S. Iceland: Skålholt!; Thingvellir!; 
Breidabolstadr!; Vestmannaey !. 

Frequent in the western and south-western parts of the country 
from Borgarfjordur southwards, rarer in the other parts. It chiefly 
occurs in humus-filled rock-fissures and in lava caves, but also upon damp 
rocks by waterfalls, in small reddish tufts, or mixed with other mosses 
such as Plagiothecium pulchellum, Mnium orthorrhynchum, Pohlia cruda, 
Myurella apiculata, M. julacea, Blepharostoma trichophyllum, etc. It does 
not appear to ascend higher than about 300 metres above sea-level. 
Fruit has been found only in Almannagjå and very scarcely; $ plants 
are rather common. 


119. Plagiobryum demissum (H. et H.) Lindb. 


W. Iceland: Modruvellir in Esja!. 

It was growing here in company with Anomobryum filiforme on a 
stony slope at an altitude of about 200 metres. The fruit was ripe on 
July 17th, but .the lids had not yet been thrown off. 


120. Pohlia acuminata Hornsch. 


W. Iceland: Geitabergsvatn!. S. Iceland: Seljaland!, 250 metres above 
sea-level; Thingvallahraun near Tintron!; Almannagjå!. 

This species grows on dry stony ground in birch coppices or on 
rock-ledges. Thus on Thingvallahraun it grew on small slopes in the 
birch coppice in company with Pohlia nutans and Heterocladium squar- 
rosulum. Near Geitabergsvatn, also, it grew on the ground in the coppice 
in unmixed tufts, 5—10 mm. high, with setæ, 10—15 mm. long. On 
July 20th the capsules were ripe, dark brown, with lids low and conical, 
muticous. 


121. Pohlia polymorpha H. et H. 


W. Iceland: Modruvellir in Esja!. 
It was growing in the above locality on a tuff-slope at an altitude 
of about 400 metres. 


122. Pohlia cruda (L.) Lindb. 


Very common. 

This is one of the most commonly occurring mosses, and is of 
almost equal frequency at all altitudes. It usually grows in rock-clefts 
filled with soil and on soil-covered ledges, but is also met with both 
on rather dry and on damp ground, as for instance on knolls in bogs. 

The fruit, which is very frequent, ripens in the first half of August. 


THE BRYOPHYTA OF ICELAND ' 471 


123. Pohlia nutans (L.) Lindb. 


E. Iceland: Seydisfjordur!; Vallanes!. N. Iceland: Myvatn (Grl.)!; 
Hålsskogur!; Heljardalsheidi (Grl.)!. NW. Iceland: Dyrafjérdur!; Isa- 
fjordur!; Sugandafjordur!; Jokulsfjordur!. W. Iceland: Breidabolstadr'!; 
Reykjavik (Grl.). S. Iceland: Frequent near Thingvellir (Grl.;!); Draupahlid 
(Wiinsted)!. 

P. nutans was first found in Iceland by Steenstrup, who did not, 
however, give the habitat. It is also recorded by Mårch, but all the 
specimens collected by him have been wrongly determined. It usually 
grows on a rather dry substratum. In the majority of the localities it 
occurred on the ground in birch coppices, as for instance in NW. Iceland, 
where it is common everywhere in the coppices at the head of the 
fjords. In that part of the country it also grows both frequently and 
abundantly on peat, and has been collected up to a height of about 
350 metres above sea-level. 

In the other parts of the country it is rather rarc, and was as a 
rule found in small quantities only. Only around Thingvellir was it 
rather common in the birch coppices. In NW, Iceland the fruit was ripe 
about July ist. 


124. Pohlia cucullata (Schwågr.) Bruch. 


Iceland (Mérch)!; Holtevad heath (Grl.)! (determined by Zetterstedt 
as Webera Ludwigii). N. Iceland: Akureyri, at an altitude of about 800 
metres. NW. Iceland: Isafjérdur!, at an altitude of about 350 metres; 
Dynjandi!; Arngerdareyri !. 

Near Akureyri it was growing in gravelly soil saturated with melting 
snow, associated with Polytrichum sexangulare, Dicranum Starckei, Pleuro- 
clada albescens, etc. The fruit was almost ripe on July 23rd. In NW. Ice- 
land it was common in several places north of Isafjordur on damp ground 
(peat) at the bottom of the valley. Here the fruit was not quite ripe 
in the beginning of July. 


125. Pohlia Ludwigii (Sprengel) Schimp. 
Pohlia Weigelii (Schimp.) Lindb. 


N. Iceland: Ljosavatnsberg!. 
In the above locality it grew abundantly on damp gravelly soil at 
an altitude of about 500 metres. Only sterile specimens were found. 


126. Pohlia commutata (Schimp.) Lindb. 


Very common. 
Pohlia commutata is one of the most frequent species, especially in 
the Alpine region; at elevations from about 300 to 600 metres it is met 


472 A. HESSELBO 


with everywhere on damp ground, often forming the bulk of the 
vegetation, for instance in several places in NW. Iceland. It is, however, 
also common in the more low-lying districts, where it grows especially 
on damp soil along streams. 

Pohlia commutata is a most variable species, and occurs in Iceland 
in numerous forms. The tufts are, as a rule, loose, and as much as- 
6 cm. high. In more particularly sheltered localities, for instance in 
lava-clefts near Thingvellir, the plants are very vigorous, and form deep, 
not-coherent tufts which are hardly tomentose. But in damp depressions 
on the mountain heights they often form extensive, deep and dense 
cushions which are brownish grey in the interior and vellowish green 
only at the top; compact forms like this occurred abundantly on damp 
gravelly flats, 400—500 metres above Isafjordur. The single plants are 
slender, with a thin fragile stem, and with leaves appressed, shortly 
pointed, slightly toothed towards the apex and longly decurrent. The 
leaf-cells are shorter, sometimes thin-walled, sometimes rather highly 
thickened and porous, but both the form of the leaves and of the cells, 
and the thickness of the walls may vary considerably in the same tuft 
or even on the same plant. ; 

The fruit, which occurs rather frequently, was usually ripe at the 
end of July. 


127. Pohlia gracilis (Schleich.) Lindb. 


Very common. 

In the lowlands this species grows, as a rule, on damp ground 
along streams, either in unmixed tufts or, more often, associated with 
Alicularia scalaris, Lophozia quinquedentata, Cephalozia bicuspidata, Pohlia 
commutata, etc. On cold, wet gravelly ground it is sometimes the most 
abundant constituent of the vegetation; thus it was growing in enormous 
quantity below the "Jéåkull” in Kaldalon (NW. Iceland). In the Alpine 
region from about 300—700 metres, Pohlia gracilis is one of the most 
frequently occurring species, and grows everywhere on damp gravelly 
flats, chiefly scattered among other mosses such as Anthelia Juratzkana, 
Alicularia spp., Polytrichum sexangulare and Oligotrichum hercynicum, 
more rarely in larger, unmixed growths. 

The fruit, which occurs very frequently, ripens on the mountain 
heights usually at the end of July, in the lowlands a month earlier 
(Kaldalon, June 22). 


128. Pohlia Rothii (Correns) Broth. 
Pohlia annotina (L.) Lindb. 


N. Iceland: Husavik g!: Stadr near Hrutafjordur!. 

It was growing in both places on damp sandy soil, associated with 
Dicranella crispa, Aongstræmia longipes and Hepaticeæ. The gemmæ, 
which occurred only few in number, were quite typical. 

Webera annotina Hedw. is recorded from the peninsulas of NW. 
Iceland (Stp.) and from Hafnarfjérdur (Grl.), but as the collections con- 


THE BRYOPHYTA OF ICELAND 473 


tain no specimens from these localities it cannot be decided which 
species is meant. 


129. Pohlia tenuifolia (Schimp.). 
Webera bulbifera Warnst. 


E. Iceland: Hamarfjérdur!; Berufjérdur!; Seydisfjårdur!. N. Iceland: 
Geitaskard!; Hnausar!. VW. Iceland: Reykjavik! gg." 

P. tenuifolia is the most frequent of the small gemmiferous species 
of Pohlia. It usually grows on damp sandy soil, and often very abun- 
dantly, always bearing the characteristic gemmæ. Near Geitaskard it 
covered in company with species of Bryum the bottom of a dried up lake. 

The leaves in all the Iceland plants are very longly decurrent. 


130. Pohlia grandiflora H. Lindberg. 


S. Iceland: Reykir!; Laugarvatn! 3; Barkarstadr!; Midskålagil near 
Holt!; cave near Steinafjall!. 

Near Reykir it was growing on the ground along a stream, asso- 
ciated with Dicranella crispa, Pogonatum urnigerum, Scapania curta and 
Lophocolea minor. Near Laugarvatn it was found abundantly on a warm 
clayey flat, associated with Aongstræmia longipes, Gymnocolea inflata and 
a small sterile Philonotis. 3 flowers were very numerous, but gemmæ 
were absent. In the other places it was found on faces of tuff rocks, 
associated with Bartramia ityphylla, Pohlia cruda, Didymodon rubellus 
and Distichium montanum. The gemmæ were quite typical in all these 
species. 

Pohlia tenuifolia is an excellent species, easily known, even when 
gemmæ are absent, by the strong sheen and the narrow, longly decurrent 
leaves, in the upper half of which the cells are very narrow and often 
wavy. 


131. Pohlia proligera Lindb. 


N. Iceland: Vidimyri!. 
It grew here rather scantily on damp sandy ground along a stream, 
bearing typical gemmæ. 


132. Mniobryum albicans (Wahblb.) Limpr. 


Pohlia albicans Lindb. 
Very common. 


Var. glacialis (Schleich.) Limpr. 


Very common. 
Both the type and the variety are the most common mosses every- 
where on damp ground. The type grows on the ground along streams, 
The Botany of Iceland. Vol. I, part II 31 


474 A. HESSELBO 


in bogs and on damp rocks. Var. glacialis grows always in cold water, 
especially in moss bogs, where in company with Philonotis it usually 
forms the bulk of the vegetation. In the Alpine region Pohlia albicans 
occurs abundantly, even at a height of about 600 metres. There it 
grows especially in and along watercourses, often filling up their — 
during summer — dried up beds by growing in large cushions both on 
the ground and on stones. As in P. commulala, the Alpine forms be- 
come more compact, with shorter, concave, appressed leaves and looser 
cell-tissue. Such forms have been found, for instance, near Dyrafjordur 
at an altitude of 300—400 metres, Esja at an altitude of about 500— 
600 metres and near Seljaland at an altitude of about 500 metres. Fruit 
occurs rather rarely, and in the type, as a rule, in a small quantity 
only. .Fruiting specimens of the type were collected near Akureyri (ripe 
July 22) Hol (ripe July) Fand Breiddalk (hardy rip] une st) Var 
glacialis was collected in fruit near Eyjafjordur (O. D.) and near Kaldalon 
in NW. Iceland, where the plant grew abundantly on inundated ground 
along small streams, and was often covered wilh fruit which was not 
guitetripeatkthetendkorune: 


133. Bryum purpurascens (R. Br.) Br. eur. 


E. Iceland: "Seydisfjordur!, common; Vallanes! (HJ): NS Iceland: 
Myvatn (Grl.)!; Akureyri!, common; Husavik!, common. NW. Iceland: 
common about Isafjordur and Joåkulsfjordur!. W. Iceland: Reykholtdalur 
(Grl.)!; Nordredalur!; Grimstunga (Grl.)!; Bardshellir (Grl)! … S. Iceland: 
Traustholtsholm in the Thjorså (F.)!. 

Grows on damp peaty or sandy soil, especially near the coast. It 
occurs abundantly chiefly in places where the peat has been pared off. 
Near Seydisfjordur it was collected up to a height of about 400 metres. 
The fruit ripens in the beginning of August. 

Bryum purpurascens varies rather considerably in the form of the 
capsule and in the structure of the peristome. The capsule is usually 
symmetrical, sometimes slightly asymmetrical, and then reminds one of 
B. uliginosum, from which it is, however, easily distinguishable by the 
large lid. The hase of the peristome teeth (fundus) is often large and 
reddish yellow, and in the majority of the plants which have been in- 
vestigated the lamellæ are sinuose in the middle, sometimes as strongly 
so as in Bryum lapponicum. The dorsal transverse segments are more 
or less distinctly transversely striated. The spores vary somewhat in 
size, from 0.024—0.035 mm. 

It is a peculiarity with this species that the capsule, on being cut 
through and warmed in diluted ammoniacal water, imparts to the latter 
a substance which is intensely, but temporarily coloured red by the 
oxygen in the air. The same is often the case also with B. uliginosum. 
In the course of time the old capsules turn almost black in colour. 


134. Bryum lacustre Blandow. 


E. Iceland: Berufjérdur!; Vallanes (H. J.;W. N. Iceland: Common near 
Husavik!; common near Eyjafjordur (0. D.;!); Saudanes (St.)!; Svinadalur!;. 


THE BRYOPHYTA OF ICELAND 475 


NW. Iceland: Bildudalr (C. H.)!. W. Iceland: Lundur!. S. Iceland: Traust- 
holtsholm in the Thjorså (F.)!; Breidabolstadr!. 

Rather common, especially in N. and E. Iceland. It grows on damp 
sandy or peaty soil, especially in places where the peat has been pared 
off, and by the sides of ditches. Near Husavik it was growing plentifully 
on damp slopes stretching down towards the sea, in association with 
Leptobryum, Aongstræmia, Bryum inclinatum, etc. Near Lundur it was 
growing in company with Dichodontium, Aongstræmia, Dicranella Schreberi 
and Bryum inclinatum on peaty flats which had been pared off. The 
fruit was ripe near Berufjérdur on June 20th, near Breidabolstadr on 
July 16th. 


135. Bryum Kaurini Philib. 


E. Iceland: Berufjérdur (Grl.)!. 
The specimens, which are quite typical, were gathered with ripe 
fruit on July 6th, 1878, and determined by Berggren as B. inclinatum 


136. Bryum archangelicum Br. eur. 


NW. Iceland: Hesteyri!. W. Iceland: Esja (Grl.)!; Brjanslækur (H. J.)!. 
S. Iceland: Drångshlid (H. J.)!. 

Both rare and scanty on rather dry, sandy ground or on soil-covered 
cocks Elbe ruitkripensfatkthetendsorsJulyg 


137. Bryum Jårgensenii Kaurin. 


SW. Iceland: Thingvellir!. 

The Iceland plant agrees exactly with B. archangelicum in habit. 
The leaves are narrowly and indistinctly bordered; the perichætial leaves 
have their margins revolute to the apex. The plants are often purely 
2: usually, however, a few antheridia and many archegonia are present. 
In none of the plants investigated did the lamellæ show the slightest 
sinuosity. The spores, which were yellow and almost smooth, were of 
very much the same size as in B. archangelicum. 


138. Bryum inclinatum (Sw.) Br. eur. 


Very common everywhere on a more or less damp substratum, on 
peat, sand and gravel, on soil-covered rocks, and in rock-clefts. It does 
not appear to ascend higher than about 300—400 metres above sea-level. 

Bryum. inclinatum varies so considerably in all its parts, in habit 
and size, in the length of the setæ, in the form and degree of inclination 
of the capsule, in the structure of the peristome, the size of the spores 
and in the form and cell-tissue of the leaf, that it is hardly possible to 
collect two tufts which do not show diversities in one respect or another. 
Perhaps on a closer investigation many of the forms might possibly 
be referred to some of the numerous "small species” established by 


315 


476 A. HESSELBO 


Hasen BryhnifRyanstiarnelstEimprrehtandkolbertauthorstkor 
might with as great justice be described as new species. But without 
comparison with original specimens, and only according to the descriptions, 
it is impossible to form any opinion of the value of these species, which 
have very often been collected only in small quantity, and from a single 
locality, and which will therefore in many cases prove to be maintainable 
only so long as there does not exist more than the one specimen. It 
is very probable that it will not even be possible to maintain as varieties 
a great many of these forms which have been given specific rank, the 
characters used for separating the species being so variable that, on 
having abundant material for investigation, it is impossible to draw any 
boundary line between the forms. Here, only thorough investigations 
made at the habitat will be able to throw some light on the influence 
of the external conditions on the specific characters. 

The height of the tufts and the length of the stems vary according 
to the degree of dampness, so that plants from dry and exposed habitats 
are low in growth with close-set leaves, and from damp localities, as a 
rule, higher, with elongated stems and more distant leaves. Also in 
places, slightly illuminated, for instance among grass and Carex-tufts, 
do the plants become higher with elongated stems. The leaves in Bryum 
inclinatum, in contradistinction to those in several other species of the 
same group, are stated to be not decurrent. This feature appears, how- 
ever, to vary considerably, and to be closely connected with the habit 
of the plant, forms with short stems having close-set, not decurrent 
leaves, while the leaves on elongated stems are, as a rule, more or less 
decurrent. In such forms the uppermost close-set leaves have almost 
always a rounded, not decurrent base, while the leaves further down 
on the stem "become" more”strongly "decurrent the "further they occur 
apart from one another. The form of the cells of the basal angles of 
the leaf also vary greatly according to the position of the leaf on the 
stem. The not decurrent leaves have a rounded base with quadrate, 
somewhat widened and thin-walled cells, but as the leaves downwards 
become more and more decurrent, the cell-form becomes elongate rect- 
angular. The leaf-form varies from narrow-lanceolate' to almost ovate, 
with a shorter or longer point. The lower leaves are usually shorter 
and broader, and the nerve vanishes just below the apex. The leaf- 
margin is revolute to the apex, more rarely it is plane in the upper part. 
The form of the leaf-cells is partly dependent on the leaf-form, long, 
narrow leaves having elongated cells, while broad, short leaves have 
broader cells, and even in leaves of the same plant the cell-form may 
vary considerably. The cell-walls vary somewhat in thickness. Xero- 
philous forms have often thick cell-walls, while bog-forms with large, 
broad leaves have thin-walled cells. The capsule varies in form from 
obovate to elongate-ovate. The neck of the capsule is usually straight, 
more rarely slightly curved. 

The peristome teeth vary in colour from pale yellow to brownish 
yellow, being very strongly coloured where inserted. The "insertion” 
may be very differently developed, all transitions are met with, from a 
slightly indicated yellowish-brown or orange ring at the base of the 
peristome teeth to a highly developed compact thickening (fundus) 


THE BRYOPHYTA OF ICELAND FIT 


which may be almost as large as, for instance, that in B. retusum. It is, 
however, not usual for the thickening to be distinctly localized around 
the base of each individual peristome tooth; it has more the charac- 
ter of a continuous ring, 0.02—0.07 mm. broad. Hagen (Musci Norv. Bo- 
realis, p. 141) pronounces the opinion that ultimately all the species with 
a distinct fundus will be united together in a separate group in a na- 
tural system of species of Bryum (Brya hæmotostomata). The feature 
in question appears, however, to be too variable for that purpose. It 
is especially in the group of forms of inclinatum that this peculiar de- 
velopment of the peristome base occurs, but it is also met with in 
other groups, although less frequently, for instance in B. purpurascens 
(where it may be almost as large as in Brya hæmotostomata Hagen) 
and in B. uliginosum. B. curvatum Kaur. & Arnell, which Hagen refers 
to B. hæmatostomata, also belongs to the purpurascens pallens group. On 
the other hand, forms may also be met with in which the fundus is 
rather slightly developed or even quite wanting, for instance B. Kau- 
rini, B. retusum and B. Graefianum, referred by Hagen to B. hæmato- 
stomata. This diminishes the value of the character in question, and 
it does not appear to have greater systematic importance than has the 
peculiar structure of the lamellæ for the group Hemisynapsium, which 
structure is especially met with in the group of species in which the 
fundus formation occurs. Both the fundus formation and the indentation 
of the lamellæ, as also the occurrence of perforations in the median 
line of the peristome teeth, are probably connected with certain external 
influences, and are very often met with together. Here it is especially 
the degree of salinity of the substratum which is of importance, aula- 
codonte peristomes being found almost exclusively in Bryum forms 
collected from localities close to the coast, while forms with a distinct 
fundus-formation, although most frequent in the vicinity of the sea, yet 
have also been found far inland, or at a height of 400—500 metres 
above sea-level. In Iceland indented lamellæ and peristome teeth more 
or less perforated in the median line have been observed in forms of 
B. inclinalum, B. retusum, B. archangelicum, B.calophyllum and B. purpur- 
ascens, in the last-mentioned species, however, the peristome teeth are 
never perforated. 

The spores vary somewhat in size, most frequently 0.022—0.025 mm., 
more rarely as much as 0.029 mm. 

Bryum lapponicum Kaurin has not been found in Iceland, but forms 
which approach this species by having broader and longly decurrent 
leaves are rather frequent. 


139. Bryum retusum Hagen. 


N. Iceland: Hnausar in Vatnsdalur!; Vidimvri!; W. Iceland: Stykkis- 
holmur!; Borgarnes!. 

This species grows on peat or soil-covered rocks. The capsules 
ripen in the beginning of July. Near Stykkisholmur they were almost 
ripe on June 15th. In all the plants referred to this species the leaves 
were shorter and broader than in B. inclinatum, with the nerve vanishing 
in the apex and shortly excurrent in the perichætial leaves. The capsule 


478 A. HESSELBO 


is narrower than in Bryum inclinatum, not-gløssy, dark brown, finely 
pitted (in B. inclinatum smooth and glossy), with a high and conical lid. 
The peristome teeth are narrow, dark yellow, with a very large, red 
fundus which is very easily loosened from the mouth of the capsule 
and falls off together with the peristome tooth belonging to it, equally 
wide to the middle or gradually tapering from the base, and very 
differently formed in the same peristome. Lamellæ 12—18 in number, 
very differently developed even in the same peristome, straight or sinuous, 
frequently also porous in the median 
line"af the"teeth: <The”sporesvary 
somewhat in size, from 0.020 mm. 
(Borgarnes) to 0.024—30 mm. (Vidi- 
myri). 


140. Bryum islandicum n. sp. 


Cæspites densi ad 5—6 mm. alti. 

Caulis ruber, fragilis, valde to- 
mentosus, innovationibus brevibus, 
numerosis. 

Folia caulina inferiora decur- 
rentia, 1.5 mm. longa et 0.7—0.9 mm. 
lata, superiora non vel vix decur- 
rentia, ovata, longius cuspidata, api- 
cem versus parce denticulata, 2—2.2 
mmsllonsaker td 07 sata 
basin rubra; margine tota longitudine 
reflexa, seriebus 3—4  cellularum 
angustiorum limbata.  Cellulæ lep- 
todermes paulum porosæ, basilares 

HiE 55 SR BE JUL FCO TER ION GE: rectangulæ, 0.03—0.04 mm. longæ et 

0.02 mm. latæ, ceteræ elongato hexa- 

gonæ, 0.03—0.04 mm. longæ et 0.015—0.018 mm. lata. Costa ad inser- 
tionem rubra, in aristam longam excurrens. 

Folia perichætialia intima anguste lanceolata, vix limbata, costa 
excurrente longiuscule cuspidata. 

Inflorescentia synoica; antheridia et archegonia numerosa. 

Seta ad 15—20 mm. longa et 0.16 mm. crassa, rubra. 

Capsula pendula, deoperculata ca. 3 mm. longa et 1 mm. crassa, 
opaca fusca leptodermis, sicca leviter rugulosa.  Operculum humile 
conicum. Annulus latus. Cellulæ exothecii leptodermes, marginales in 
series 3—4 transverse rectangulæ-polygonæ quadratæ, ceteræ irregulares. 

Exostomii dentes e fundo fusco-aurantio 0.4 mm. longi et 0.065 
mm. lati, fusco-aurantii apice lutei, anguste limbati, subtiliter punctulati. 
Lamellæ 14—16, inter se liberæ, marginibus integris. Endostomium 
exostomio haud connatum, luteolum, parce papillulosum;. processibus 
lanceolatis, in carina ovaliter fenestratis, ciliis pro more rudimentariis. 

Sporæ 0.039—0.038 mm. diam. olivaceæ, punctulatæ. 

N. Iceland: Akureyri, on damp ground at an altitude of about 300 
metres, July 20th, 1909. 








THE BRYOPHYTA OF ICELAND 479 


141. Bryum calophyllum R. Brown. 


E. Iceland: Vallanes!, on wet .boggy ground. The capsules were 
quite green even on June 27th. 











DØNNN 
MAN DDO 
NON 


Fig, 3... Bryum islandicum, aa, Lecaf margin (X 120); b and c, perichætial leaves; d and e, leaves. 





142. Bryum uliginosum (Bruch) Br. eur. 


N. Iceland: Husavik!; Svinadalur!. W, Iceland: Haukadalsheidi! at 
an altitude of about 300 metres. 

In all the above localities it grew abundantly on damp, sandy ground. 

On Haukadalsheidi the fruit was 
ripe on Aug. ist, in the other localities it 
was unripe in the middle of July. 

Besides g and 2 flowers, a few 
hermaphrodite flowers also occur fre- 
quently. The papillæ of the peristome 
are in the lowermost transverse seg- 
ments often distinctly arranged in 
transverse rows, and sometimes there 
are also distinct transverse bars as in 
'BEpurpurascens KASSE the case in 
the latter species, the ripe capsule 
contains a colouring matter which on 
being treated with ammonia assumes 
alredtcolourtimethetarr: 


143. Bryum Grænlundii n. sp. 





Cæspites laxiusculi, ca. 1.2 cm. 
alti, Superne virides, subnitentes: SUT- Fig. 4. Bryum islandicum. Peristome 
culi innovationes tenues. (X 120; phot.). 


480 A. HESSELBO 


Folia caulina decurrentia, inferiora ovata, breviter cuspidata, 
superiora longius cuspidata, breviter vel non decurrentia, 2.3 mm. longa 
et 0.9—1 mm. lata, apice integra vel leviter dentata; margine ad apicem 
reflexa, seriebus 2—4 cellularum angustiorum limbata. Cellulæ foliares 
leptodermes, indistincte porosæ, basilares rectangulæ, 0.035—0.050 mm. 
longæ et 0.018—0.020 mm. latæ, ad insertionem rubræ; illæ folii medii 

elongato-hexagonæ, vel rhomboideæ 

| 0.05—0.08 mm. longæ et 0.015— 

0.018 mm. latæ, apicales angustiores. 

Costa fusco-lutea, ad insertionem 

0.08 mm. lata, in foliis inferioribus 

sub apice desinens, in superioribus 
in cuspidem integram excurrens. 

Folia perichætialia intima 
anguste triangula, margine limbata. 

Inflorescentia synoica; an- 
theridia perpauca; archegonia 
et paraphyses numerosæ, roseæ. 

Seta 3—4 cm. longa et 0.20 mm. 
crassa, rubra. 

Capsula nutans, arcuato-clavata 
deoperculata 3.35 mm. longa et 1 mm. 
| crassa, sub orificio obliquo rubra 
contracta; collum 1.5 mm. longa. 
Operculum humile conicum, apice 
rubro. Annulus 0.14 mm. latus, 
triplex. Exostomii dentes e fundo 
purpureo vel rubro-aurantio bene 
definito 0.40—0.45 mm. longi et 
0.10 mm. lati, fusco-lutei, anguste 
limbati, dense et subtiliter papil- 
lulosi; lamellæ ca. 24, inter se liberæ, 
margine integra. Endostomium 
exostomia vix adhærens, luteum, 
subtiliter papillulosum; processus 
dentibus æquilongi, fenestrati; Cilia 

FEE BE Gren OC 3) nulla. Sporæ olivaceæ, 0.025—0.033 
mm. diametro, punctulatæ. 

W. Iceland: Haukadalsheidi on damp gravelly ground at an altitude 

of about 350 metres, August ist, 1909. 














144. Bryum fallax Milde. 


N. Iceland: Geitaskard!, fr. 
It was growing here in a dried up pool among Pohlia tenuifolia. 


145. Bryum æneum Blytt. 
Bryum rutilans Brid. 


E. Iceland: Djupivogur!; Seydisfjordur!; at an altitude of 400 metres. 
N. Iceland: Akureyri!, at an altitude of 350 metres. NW. Iceland: Bildu- 


THE BRYOPHYTA OF ICELAND 481 


dalur (C. Hansen)!. NW. Iceland: Lundur!, fr. S. Iceland: Vestmannaey!; 
fr.; Skålholt!; Austerhlid near Geysir!; Breidabolstadr!; Barkarstadr!; 
Holt!, fr.; Drångshlid !. 

Common in the southern part of the country and on Vestmannaey; 
rarer in the other parts of the country. In S. Iceland it grew especially 











Fig. 6. Bryum Grænlundii. a, Capsules (X 20); b, leaves (X 20); c, perichætial leaves (X 20); d, leaf 
margin at base, and e, from the middle of the leaf (X 120). 


abundantly everywhere in clefts of tuff rocks, and on Vestmannaey it 
was also frequent on the faces of damp tuff rocks. It was found.in 
several localities in fruit, which was quite green even in the beginning 
of July. Near Lundur it was found on damp ground, associated with 
other Bryum spp. and Dichodontium, and had ripe fruit at the end of 
July. In several of the fruiting specimens from this locality a few 
hermaphrodite flowers were found besides the numerous 2 flowers. 2 
plants occur everywhere, but gg plants have not been found. 

In the leaf-angles of all the plants investigated hair-formations were 
found, consisting of readily falling, $—10 celled, brown, papillose hairs, 
which very easily break into pieces of 2—4 cells. The leaves vary from 
shortly pointed, with nerve vanishing just below the apex, and shortly 


482 A. HESSELBO 


rhomboid, equilateral-hexagonal or almost quadrate cells (forms from 
Akureyri), to more or less longly pointed, with excurrent nerve, and 
narrower, rhomboid or elongate-hexagonal cells. 





146.. Bryum bimum Schreb. 


E. Iceland: Vallanes. 

It has been collected only 
scantily on damp ground, with 
capsules which were not ripe 
at the end -of June. 

Two sterile Brya, collected 
by Helgi Jønsson near Ståd 
in E. Iceland and in Budahraun, 
are referred to this species by 
C.Jensen. The leaves in both 
the plants taper to a long point, 
are longly decurrent, have nar- 
row basal cells, and the nerve 
excurrent; therefore, the plants 
probably belong to Bryum affine, 
but owing to the absence of 
fruit the determination is quite 
uncertain. 





147. Bryum affine (Bruch) 
Lindb. 


Bryum cuspidatum Schimp. 





i K Ange " , 


Commonly distributed. 
Fig. 7. Bryum Grænlundii. Peristome (X 250; phot.). One of the most frequent 
species of Bryum, and occurs 
everywhere in the lowlands on damp sandy or peaty ground, and on 
humus-covered rocks. The fruit ripens in the first half of July. It 
yaries exceedingly in all its parts. It is said to differ from the very 
nearly allied B. cirratum in its sex, the smaller, smooth spores, and the 
decurrent leaves with the broader cells; but all these features are so 
variable that in reality it is hardly possible, at least in the Arctic 
countries, to separate these two species. 

The plant is usually synoicous, but & flowers are also almost 
always present; 2 flowers are more rare. The spores vary in size from 
0.008 to 0.020 mm., and often differ rather considerably in size in the 
same capsule, or in different capsules from the same tuft; the most 
frequent size is 0.012—0.017 mm.; they are sometimes smooth, sometimes 
fincly papillose. The leaves vary in form from about ovate to lanceolate, 
tapering more or less to a long point, and with the nerve longly excurrent. 
In form the leaf-cells correspond nearly with the leaf, so that short 
leaves have broader cells than have those which are long and narrow. 


THE BRYOPHYTA OF ICELAND ASE 


"The leaf-base, at any rate in the lower leaves, is decurrent, with 
elongated, narrow alar cells. The close-set uppermost leaves are usually 
not decurrent, and then. have the base rounded, with quadrate, some- 
what widened alar cells. 

The structure of the leaf-base evidently depends on the degree of 
dampness of the habitat. Bryum cuspidatum usually grows on boggy 
ground and forms here tufts, 1—3 cm. high, with rather long innovations. 
On such elongated shoots the leaves are always decurrent. On a more 
dry and exposed substratum, for instance on rocks, the tufts become 
lower in growth and denser.. and then the leaves are only slightly, or 
not at all, decurrent, and in connection with this the alar cells are 
short as in Bryum cirratum. 


148. Bryum cirratum Hoppe et Hornsch. 


Rather common. 

Grows in drier localities than does B. affine, for instance in gravelly 
soil, on slopes and among grass. It is,especially common in NW. Iceland, 
and occurred abundantly for instance on stony slopes near Dyvrafjordur. 

The forms which are here referred to B. cirratum have é, 2 and 


numerous gg flowers. The leaves are not decurrent, taper to a very long 
point, with narrow, almost linear cells in the upper portion. The spores 
are 0.014—0.020 mm., yellow, and finely papillose. Bryum affine and B. 
cirratum form a continuous series of forms, in which B. affine represents 
the hygrophilous and B. cirratum the xerophilous adaptation-form. 


149. Bryum intermedium (Ludw.) Brid. 


N. Iceland: Grimsey (0. D.)!: Vidimyri!; Stadr near Hrutafjordur!. 
S. Iceland: Merkjåfoss (F.)!. 

Rather rare and scanty on damp soil. Near Stadr the fruit was 
ripe on August ist, but the lid still persisted. 


150. Bryum pallescens Schleich. 


Widely distributed. 

One of the most frequent species everywhere up to a height of 
about 300 metres above sea-level. It usually grows on damp rocks, 
where it often forms very large and deep cushions, with numerous 
capsules; but it also occurs on damp soil. In South Iceland it is very 
common everywhere on faces of tuff rocks. The plant is, as a rule, 
monoicous, but very often hermaphrodite flowers are also met with, 
containing many antheridia and a few archegonia. , 


151. Bryum subrotundum Brid. 


S. Iceland: Uxavatn (F.)!. 


484 Å. HESSELBO 


152. Bryum capillare L. 


E.Iceland: Hamarfjérdur!. N.Iceland: Mådrufellshraun (St.)!; Grimsey 
(O. D.)!. W. Iceland: Hvammur (Grl.)!: Hafnarfjérdur!; Videy!, several 
places near Reykjavik!. S. Iceland: Breidabolstadr!; Holt!. Vestmannaey !. 

This species occurs, as a rule only sparingly, on humus-covered 
rocks and in lava-clefts. Only on Vestmannaey was it common, occur- 
ring especially on the tumbled-down blocks at the foot of cliffs inhabited 
by sea-fowl. 

The nerve usually vanishes just below the hair-shaped leaf-point. 
In the plant from Moådrufellshraun and in the majority of the specimens 
from Vestmannaey the nerve varied considerably in length, sometimes 
reaching scarcely beyond the middle of the leaf, sometimes almost to 
the apex. The plants from the former habitat, in addition, bore clusters 
of easily falling brown threads in the uppermost leaf-angles, quite similar 
in appearance to those in Bryum æneum. 


153. Bryum cæspiticium L. 


Iceland (Krabbe, 1863)!. E. Iceland: Seydisfjordur!; Vallanes (H. J.;!); 
Hallormstadaskogur!.  N. Iceland: Hålsskogur!; Stadartunga (0. D.)!; 
Vidvik (P. Soféniasson)!. S. Iceland: Almannagjå!. 

Gronlund records this species from Reykjavik and Laugardalur, 
but there are no specimens of it from these localities in the collections. 

Grows in dry sandy soil, especially in birch coppices, and appears 
to be rather frequent in North and East Iceland. It occurred abundantly 
in several localities in Hålsskogur. Near Vallanes it occurred sometimes 
on dry sandy soil, sometimes on the dikes and peat-walls of the farm. 
In Hålsskogur the fruit was ripe on July 19th, but some of the lids 
still persisted. 


154. Bryum comense Schimp. 


N. Iceland: Reistarårgil (0. D.)! 2; Hof (St.)!. 
Both the above localities are situated on the east side of Eyjafjordur. 


155. Bryum elegans N. v. Es. 


E.Iceland: Djupivogur! gg: Berufjérdur (Grl.)!. N.Iceland: Reykjahlid! 
d. NW, Iceland: Isafjardarheidi, at an altitude of 450 metres! 3 and ?. 
W. Iceland: Hafnarfjérdur!; Svinahraun!. Vestmannaey !. 

Not common, but sometimes abundant in humus-filled rock and 
lava clefts, etc. Near Reykjahlid it was common in the lava-field. 

Varies considerably in leaf-form, the length of the nerve, etc. The 
plant from Isafjérdur has distinctly bordered, erect spreading, and 
narrower leaves, and belongs most nearly to var. carinthiaca (Br. eur.) 
Breidl. In lava-clefts there usually occurs a low, compact form with 


THE BRYOPHYTA OF ICELAND 485 


concave, adpressed, and not bordered leaves, with loose cell-tissue and 
not excurrent nerve, which agrees most closely with var. Ferchelii (Funck) 
Breidl. 


156. Bryum argenteum L. 


Commonly distributed. 

This species has been found only in the lowlands, where it is 
especially frequent around inhabited localities. It grows there on the 
peat walls of the houses, on the dikes and on the ground in the neigh- 
bourhood of the dwelling houses, and prefers, on the whole, a wel)- 
manured substratum. It is also common near the sea-side, both on 
sandy soil and upon stones, and occurs especially abundantly below 
cliffs inhabited by sea-fowl, for instance on Vestmannaey. 


157. Bryum Neodamense ltizigs. 
Var. ovata (Jur.) Lindb. et Arnell. 


E. Iceland: Seydisfjordur!, sterile. 
It was growing here in cushions about 6 cm. high, in water in a 
stream, at an altitude of about 100 metres. 


158. Bryum Duvalii Voit. 


Very common. 

Grows everywhere on very wet ground or in water, especially in 
moss bogs or along the banks of streams, where it often forms large 
vinous red cushions in the light green patches of Philonotis, Bryum ven- 
tricosum and Mniobryum albicans. As a rule it does not ascend higher 
than 300—400 metres. Near Barkarstadr in S. Iceland it was found at 
an altitude of 500 metres. The fruit, which ripens in the middle of 
July, is rare, and has been found only rather scantily near Berufjérdur, 
Ljosavatn and Tverå in Øxnadalur. 


159. Bryum pallens Sw. 


Widely distributed. 

Grows on a damp substratum, both on soil and on rocks. In 
S. Iceland it is very common on faces of damp tuff rocks. It grows 
especially on the marginal strip of soil and on the slopes along streams, 
in small reddish tufts, associated with Dichodontium, Philonotis and 
Dicranella crispa, but it is very often sterile. On the promontory near 
Reykjavik it grew in enormous quantities on damp peaty ground, with 
masses of capsules which, in the first week of August, were ripe, but 
the lids were not yet thrown off. 

It is only by exception that Bryum pallens ascends higher than 
about 300 metres above sea-level. Near Berufjérdur it has been collected 
at an altitude of about 500 metres. 


486 A. HESSELBO 


Note... Bryum turbinatum is recorded from Iceland by Morch, 
Hornemann and Gronlund, but all the specimens in the collections 
are wrongly determined. 


- 160. Bryum ventricosum Dicks. 


Very common. 

One of the most frequent mosses, which occurs in numerous forms 
everywhere on a damp substratum, both on soil and on rocks. In 
water, for instance in moss bogs and along the banks of streams, it 
forms extensive, light green mats. The plants are only slightly tomentose, 
with large, spreading leaves, and thin-walled cells. In bogs, where it 
occurs everywhere in abundance, it is sometimes low in growth, some- 
times as much as 6—7 cm. high, and often richly fruiting. Forms from 
a more dry substratum, for instance from rocky clefts, are low in growth 
and densely tomentose. It is common upwards of 600 metres above 
sea-level. 


161. Bryum arcticum (R. Br.) Br. eur. 


This species is widely distributed over the whole of Iceland, but 
occurs, as a rule, rather sparingly. 

It grows on peat, on damp sandy soil and on humus-covered rocks, 
usually mixed with other Brya, and hardly ascends higher than 200— 
300 metres. The fruit ripens at the end of July. 

It varies rather considerably. The capsule is sometimes short and 
inflated, sometimes more slender. The spores vary from 0.020—0.035 mm. 
The lamellæ are usually connected by one or two transverse bars. 
Near Seydisfjordur a form has been collected with a partly aulacodont 
peristome and very few transverse bars. 


162. Bryum pendulum (Hornsch.) Schimp. 


N. Iceland: Hof (O.D.)!. W. Iceland: Reykjavik!; Reykholtdalur!. 
S. Iceland: Gilfoss (F.)!; Barkarstadr!; Breidabolstadr!. Vestmannaey!. In 
previous lists some more habitats have been enumerated for this 
species (Grl.; H. J.), but they are all due to confusion with other species, 
mostly with B. arcticum. 

Bryum pendulum appears to be rather rare and, as a rule, has been 
found only in small quantity mixed with other Bryum spp. on damp 
sandy soil or on damp tuff rocks. Fruit ripens in the beginning of 
August. 

As in Bryum inclinatum the base of the peristome may be rather 
differently developed. Thus, in a specimen from Breidabolstadr, a 
rather large reddish-yellow thickening had been developed which could 
be loosened from the mouth of the peristome together with the peristome 
tooth belonging to it, as in the species of Hagen's Brya hæmatostoma. 

One of the specimens from Hof, collected by 0. Davidson, was 
monoicous, but was otherwise quite typical. 


THE BRYOPHYTA OF ICELAND 487 


163. Bryum Brownii Br. eur. 


N. Iceland: Hof (0. D.)!. W. Iceland: Reykjavik '!. 

On the peninsula near Reykjavik it occurred abundantly on damp 
sandy ground in company with Bryum pallens and B. inclinatum. Fruit 
was ripe on August Sth. 

It is also recorded from several localities which are cnumerated in 
Gronlund's lists, but all the corresponding specimens in the collections 
are wrongly determined. 


164. Mnium hornum L. 
Åstrophyllum hornum Lindb. 


E. Iceland: Common from Hornafjérdur to Berufjérdur!; Seydis- 
fjordur!; Vallanes (H. J.)!." N. Iceland: Akurcyri!: Hof near Eyjafjordur 
(O. D.)!; Geitaskard!; Vatnsdalur!. NW. Iceland: Common everywhere 
about Isafjordur!; Bildudalr (C. H.)!: Dvrafjérdur!; near Jokulsfjordur!. 
W. Iceland: common everywhere (Grl.; H. J.;!). S. Iceland: Common!. 
Vestmannaey !. 

Common in East, South, West and North-west Iceland, rarer north- 
wards. It occurred abundantly even in Vatnsdal and Blånddal (Geita- 
skard), but further eastwards it was collected only in a few localities 
near Eyjafjordur. It appears to be quite absent, for instance, from the 
district around Husavik. It grows almost exclusively on somewhat damp, 
peaty ground, especially on knolls or along the sides of ditches, more 
rarely on soil-covered rocks. 2 plants occur rather frequently; on the 
other hand, it has not been found in fruit. 


165. Mnium orthorrhynchum Brid. 
AÅstrophyllum orthorrhynchum Lindb. 


Very common everywhere in soil-filled rock-clefts, and ravines, on 
faces of damp tuff rocks, etc., and occurs in wide-spread unmixed tufts 
or more often mixed with other mosses. gg flowers are rather common; 
on the other hand, it has not been collected in fruit. It does not appear 
to ascend higher than 400—500 metres above sea-level. 


166. Mnium serratum Schrad. 
Astrophyllum marginatum (Dicks.) Lindb. 


N. Iceland: Hof near Eyjafjérdur (O. D.)!. Vidvik (P. Soféniasson)'!. 
W. Iceland:  Giljårfoss and Kleppjarnsreykir in Reykholtsdalur (Grl.)!; 
Kalmanstunga (Grl.)!; Fell near Kollafjérdur (St.)!; Alafoss!. S. Iceland: 
Flokastadagil!; Barkarstadr!; Holt!; Austarhlid near Geysir!. Vestmannacy!. 
It grows on damp, shady rock-sides in caves and clefts in the 
lowlands, and is rather common in the southern part of the country, 


488 A. HESSELBO 


where it often occurs abundantly in the clefts of tuff rocks. In West 
Iceland it is met with here and there, while it is rare in North Iceland 
and has not yet been found in East Iceland. The fruit, which is found 
everywhere, ripens in the first half of June. 


167. Mnium spinosum (Voit) Schwægr. 
Astrophyllum spinosum Lindb. 


N. Iceland: Åsbyrgi (St.)!; Hof near Eyjafjordur (O. D.)!; Akureyri!. 

In all the above localities it was found on dry grass-covered ground 
growing among Hylocomium proliferum, H. squarrosum, H. triquetrum, 
Hypnum uncinatum and Lophozia lycopodioides. Near Akureyri it oc- 
curred, however, also on a somewhat damp slope (200 metres above 
sea-level), intermixed in a large tuft of Mnium stellare. Only 2 flowers 
have been found. 


168. Mnium undulatum (L.) Weiss. 


Astrophyllum undulatum Lindb. 


W. Iceland: Kollafjérdur!, at an altitude of 100 metres. S. Iceland: 
Vik; Hnappavellir; Hjorleifshåfdi (H. J.)!; Reykir!; Breidabolstadr!; 
Barkarstadr!; Seljaland!; Holt!; Drångshlid!. Vestmannaey!. 

This species has been found only in the south-western part of the 
country, where it is common on Flåtshlid and along Eyjafjall. It was 
found in several places on Vestmannaey, for instance abundantly in 
Heljusdalur. It grows as a rule in damp soil at the foot of rock-walls 
and in clefts, especially in localities which are well protected and have 
a luxuriant grass-vegetation. It is also met with in damp caves, for 
instance near Reykir. Only sterile specimens have been found. 


"169. Mnium cuspidatum (L.) Leyss. 


Astrophyllum silvaticum Lindb. 


N. Iceland: Hof near Eyjafjordur (0. D.)!. S. Iceland: Holt!.  Vest- 
mannaey '!. 

Near Holt it grew at the bottom of and up on the sides of a small 
tuff-cave. On Vestmannaey it was found rather abundantly among grass 
on a somewhat damp slope. 

Mnium cuspidatum is enumerated in Vahl's list, and has, moreover, 
been recorded from several localities by Groånlund, Stefansson and 
Helgi Jonsson. All these specimens are, however, wrongly determined, 
and the majority of them belong to M. affine. 


170. Mnium medium Br. eur. 


Astrophyllum medium Lindb. 


SE. Iceland: Hornafjordur. 
It grew here on boggy ground mixed with M. affine and M. cinclidioides. 


THE BRYOPHYTA OF ICELAND 4859 


171. Mnium affine Blandow. 


Astrophyllum cuspidatum (L.) Lindb. 


Very common. 

This species grows both in bogs, where it occurs everywhere and 
almost always intermixed with other bog mosses, especially Hypnaceæ, 
Cinclidium stygium and Mnium cinclidioides, and on the ground among 
grass, as also on rocks. It is most frequent in the lowlands, and hardly 
ascends higher than 400—500 metres above sea-level. 2 plants are very 
common; on the other hand it has not been collected in fruit. 

It varies considerably in habit, leaf-tissue and serrature of the leaf- 
margin. In very wet localities the annual shoots become erect and the 
stems often highly tomentose (var. elata Lindb.). On a more dry substratum, 
especially on humus-covered rocks, the annual shoots are bent down- 
wards in a curve, and are rooting at the apex, and the fertile shoots 
are quite short. The leaves are only very slightly or not at all decur- 
rent. In the forms from a dry substratum the leaf-cells are rather 
thick-walled, distinctly porous, and often collenchymatous; in bog-forms 
they are more thin-walled and slightly porous. The teeth of the leaf- 
margin are usually unicellular and obtuse, more rarely 2—4-cellular; 
forms are very frequently met with in which the teeth are very slightly 
developed or almost quite wanting, so that ail transitions to var. inte- 
grifolia Lindb. occur. The latter form is occasionally met with, especially 
on rocks. 


172. Mnium Seligeri Jur. 
Astrophyllum Seligeri (Jur.) Lindb. 


E. Iceland: Hornafjérdur!; Berufjérdur!; Seydisfjordur!; Vallanes 
(H. J.)!. N. Iceland: Husavik!; Akureyri!; Ljosavatn!: Helgavatnsfloi (St.)!. 
W. Iceland: Stykkisholmur!. Very common in SW. Iceland!. 

Grows on a very wet substratum, in marshes, moss bogs and along 
streams, and often very abundantly. 9? plants are almost always met 
with; on the other hand, gg plants or plants in fruit have not been 
collected. i 

M. Seligeri is one of the mosses of most common occurrence in the 
south-western part of the country, from Hvalfjéordur southwards, and 
also about Hornafjérdur; it often forms the bulk of the vegetation, for 
instance in the great bogs in Olfus. In E. Iceland it is rather common, 
and is also hardly rare in N. Iceland, but from NW. Iceland it appears 
to be quite absent. It is easily distinguished from M. affine by its broadly 
decurrent leaves, which are concave on the under surface. The leaf- 
cells are, as a rule, somewhat smaller than in M. affine, 0.04—0.07 mm., 
more thick-walled than in the latter species and decidedly collenchy- 
matous; forms are, however, often found with thin-walled and somewhat 
larger cells, which are then hardly to be distinguished from Mnium 
affine v. elatum except by the decurrent leaves. 

The Botany of Iceland. Vol. I, part II. 32 


490 A. HESSELBO 


173. Mnium stellare Reich. 
Astrophyllum stellare Lindb. 


ES Iceland: FBerujordur' REF ORSEEN Teen ds FA kur eye Er EVER 
in Oxnadalur!. W. Iceland: Waterfall in.Brynjudalur (Grl.)!. S. Iceland: 
Holt!; Vestmannaey!. 

This species grows on damp soil, in rock-clefts, and on tuff, and 
as a rule sparingly. Near Akureyri it was found mixed with M. spinosum 
on a damp slope, and in fruit which was ripe on July 20th, but the 
capsules had not then thrown off their lids. Near Berufjérdur it grew 
in the fissures in the basalt, mixed with Pohlia cruda, Distichium mon- 
tlanum and Plagiotheciaom pulchellum. In Heljusdalur on Vestmannaey it 
covered the vertical faces of large, tumbled-down blocks of tuff rock 
in company with M. serratum and M. orthorrhynchum. 


174. Mnium cinclidioides (Blytt) Huben. 
Astrophyllum cinclidioides (Blytt) Lindb. 


Very common over the whole of Iceland. 

The most common species of Mnium. It is hardly absent from 
any bog, where it often forms the bulk of the vegetation, and it is also 
found everywhere in moss bogs and along the banks of streams. It is 
common even at a height of above 400 metres above sea-level, and 
occurs, for instance near Seydisfjordur and Akureyri, in bogs at an 
altitude of about 500 metres. 2 plants are common, gg plants and fruit 
have not been found in Iceland. 


175. Mnium punctatum (L.) Hedw. 


Astrophyllum punctatum (L.) Lindb. 


Very common all over Iceland. 

Grows everywhere in damp localities, in marshes, where it often 
forms the bulk of the vegetation, in moss bogs, along the banks of 
streams, on wet rocks, by waterfalls, etc. It is most frequent in the 
lowlands up to 300—400 metres, but very often it ascends considerably 
higher, for instance in Svinaskard to about 500 metres above sea-level, 
and in Berufjardarskard to about 550 metres. 

? plants are common; the fruit, which is frequently found, was 
overripe in the middle of June, and the capsules had thrown off their 
lids (E. Iceland). 


176. Mnium subglobosum Br. eur. 


Astrophyllum pseudopunctatum (B. S.) Lindb. 


E. Iceland: Hornafjérdur!, sterile. N. Iceland: Hof near Eyjafjérdur 
O. D.)!, fr.; Stadr near Hrutafjérdur!, fr.; Méådruvellir (St.); Myvatn 
(Grl.)!, sterile. "W. Iceland: Reykjavik!. The other habitatg recorded 
by Gronlund belong to Cinclidium stygium. 


THE BRYOPHYTA OF ICELAND 491 


This species is no doubt rather widely distributed, but often over- 
looked on account of its great resemblance to the far more frequently 
occurring M. punctatum. Like the latter it grows on wet boggy ground. 
Fruit was unripe both near Hornafjéordur (June 11th) and near Stadr 
(July 29tb). 


177. Cinclidium stygium Sw. 

Very common. 

In East and North Iceland this species is one of the mosses of 
most common occurrence, and is found everywhere in bogs, where it 
is often the most abundant constituent of the vegetation. Here it grows 
by preference in the wettest parts, growing even in the shallow water 
on inundated ground or along streams, but it occurs also on more dry 
ground, and then usually mixed with other bog mosses, for instance, 
Hypna, Mnia and Sphagna. In the other parts of the country it is also 
common, although there it does not occur so abundantly; in South Ice- 
land I have only observed it growing intermixed — as a rule scantily 
— with other bøg mosses in, I think, every bog, but never in large 
growths. Near Akureyri it is frequent as far upwards as about 600 
metres above sea-level, and in East Iceland, for instance near Berufjordur, 
it was collected up to a height of 550 metres. The fruit, which is 
common especially in North Iceland, ripens at the end of July. 

Note. A small sterile plant, collected by Helgi Jonsson near 
Håfdi, was by C. Jensen referred with doubt to Cinclidium subrotun- 
dum, but a closer investigation has proved that it must be referred to 
C. stygium. 


Fam. MEESEACEÆ. 
178. Paludella squarrosa (L.) Brid. 


This species has a similar distribution to that of Cinclidium stygium. 
It grows in bogs and is exceedingly common especially in N. Iceland, 
where it sometimes occurs mixed with other bog mosses, sometimes in 
large, pure cushions. It is also common in E. Iceland, at any rate from 
Berufjordur northwards, in NW. Iceland and in W. Iceland. In S. Iceland 
it is far more rare, and has not been found, for instance, on Fljåtshlid 
or below Eyjafjall. It does not appear to ascend very much higher 
than about 400 metres above sea-level. Fruit was found only near 
Eyjafjordur. 


179. Amblyodon dealbatus (Dicks.) P. Beauv. 


N. Iceland: Hof near Eyjafjordur (0. D.)!, fr. 
It was growing here scantily, intermixed in a tuft of Distichium 
montanum, Ditrichum flexicaule, Myurella julacea, etc. 


180. Meesea trichoides (L.) Spruce. 


Meesea uliginosa Hedw. 
Very common. 
This species grows especially on boggy ground, but is also frequent 


32 


499 A. HESSELBO 


on wet sandy or gravelly ground and on damp rocks. In the greater 
part of the country it is very common in such localities; only in NW. 
Iceland and partly in S. Iceland does it occur more scantily, for instance 
on Fljétshlid and below Eyjafjall. It is most frequent in the low land, 
but nevertheless often ascends up to a height of 500—600 metres above 
sea-level. The fruit, which is always present plentifully, ripens at the 
end of July. 


181. Meesea triquetra (L.) Ångstr. 
Meesea tristicha Br. eur. 


Common in E. and N. Iceland!. NW. Iceland, Årngerdareyri!; W. 
Iceland: frequent about Reykjavik and Esja!; rarer and most scanty in 
S. Iceland. - 

It grows on boggy ground, sometimes in small unmixed tufts, some- 
times intermixed with other mosses such as Hypnaceæ, Paludella, Mnium 
affine, etc. Only sterile specimens have been found. 


182. Catoscopium nigritum (Hedw.) Brid. 


Very common in E., N. and W. Iceland, more råre and rather scanty 
in NW. and S. Iceland. It grows on wet boggy ground, where it some- 
times forms very dense tufts about 8$—10 cm. high, sometimes grows 
interspersed in tufts of other mosses. It occasionally occurs so abun- 
dantly, especially in N. Iceland, that it may form the main part of the 
vegetation in the bogs. 

Gronlund records that this species is one of the character-plants 
of the warm ground, and C. Jensen (Bot. Tid. XIV 1885) has described 
a var. Grønlundii: "filiformis, humilis, superne flavescens, inferne nigre- 
scens. Folia subremota imbricata, minima, ovata lanceolata, integra vel 
subintegra, costa crassa in apicem evanida, cellulis minoribus, incras- 
satis et papillosis,” which was collected by Gronlund near a hot spring 
in Reykholtdalur. In Gronlund's collections in the Botanical Museum 
in Copenhagen there are some mosses determined as Catoscopium nigri- 
tum or C. nigritum var. Grénlundii from a warm substratum. Of these 
only one specimen from Reykholtdalur belongs to var. Gronlundii, and 
I have collected similar forms rather scantily near other hot springs. 
This form differs only slightly from the type, chiefly only in being less 
tomentose and in having shorter leaves. Almost all the other specimens 
of Catoscopium collected by Gronlund were wrongly determined, as 
they belong to Archidium phascoides, and on the whole Catoscopium 
occurs only accidentally and scantily on a warm substratum. 

The fruit, which occurs very frequently and plentifully, was in 
June-July either old and emptied from the previous year or yet quite 
unripe. 


183. Conostomum boreale Swartz. 
Conostomum tetragonum (Vill.) Lindb. 


Widely distributed over the whole of Iceland. 
In W. and S. Iceland this species is found only by exception at a 


THE BRYOPHYTA OF ICELAND 493 


lower level than about 250 metres, and not until a height of about 400 
metres and upwards is it common. In N. and E. Iceland it is also most 
frequent in the Alpine region, but grows, however, abundantly down to 
the low land in many places, for instance near Hof in the south-east. 
In NW. Iceland it is very common from the sea-level up to the snow-line. 

It usually grows on dry and stony — more rarely damp — ground, 
most frequently on rocky flats, where it forms cexceedingly dense and 
compact cushions, as much as 10 cm. deep, which occur wedged very 
firmly into the substratum. The cushions are, as a rule, interwoven 
with hepatics such as Lophozia alpestris, L. ventricosa v. porphyroleuca, 
Pleuroclada albescens, Cephalozia pleniceps, etc. 

Like many other Alpine species Conostomum also occurs in the 
iava-fields, for instance abundantly in the clefts near Thingvalla; the 
tufts are here considerably looser and less tomentose than in the Alpine 
form. Fruit, which occurs everywhere, ripens in the first half of August. 


Fam. AULACOMNIACEÆ. 
184. Aulacomnium palustre (L.) Schwågr. 
Sphærocephalus palustris (L.) Lindb. 


Very common on damp boggy ground along the banks of streåms, 
and on damp, soil-covered rocks. It occurs most abundantly on knolls 
in bogs, in company with species of Sphagnum, and among (arices in 
damp tracts of meadow land. Only sterile specimens have been found. 


185. Aulacomnium turgidum (Wahlb.) Schwågr. 
Sphærocephalus turgidus (Wahlb.) Lindb. 


E. Iceland: Hof!; Seydisfjordur!; Skreiddal!; Berufjardarskard!, at 
an altitude of 540 metres; Djupivogur!. N. Iceland: Vatnsdalsfjall (St.)!; 
Lækjarmot!, at an altitude of 350—450 metres; Akureyri!, at an alti- 
tude of about 900 metres. NW. Iceland: Grunnavik!, at an altitude of 
260 metres; åÅrmuli!, at an altitude of 150 metres. W. Iceland: Esja!, 
at an altitude of 420 metres. S. Iceland: Holt near Eyjafjall!; in several 
places from the low land up to 400 metres above sea-level. 

This species is rather common especially in E. Iceland, in fact it 
may almost be called common there; for instance, near Hof in SE. Ice- 
land it was very frequent and, in several localities, plentiful. It appears 
to be of almost equal frequency in the lowlands and on the mountain 
heights. 

It grows especially on somewhat damp gravelly ground, more rarely 
on soil-covered rocks, associated with Hylocomium, Rhacomitrium cane- 
scens, Dicranum congestum, Hypnum uncinatum, etc. Near Hof it grew 
in great abundance along the river, on the gravelly flats covered with 
Grimmia canescens. Near Lækjarmot it was common as an intermixture 
in the Rhacomitrium hypnoides heath on the mountain slopes, about 
350—400 metres above sea-level, and near Grunnavik it grew on damp 
rocks among other møosses. Only sterile specimens have been found. 


494 A. HESSELBO 


Fam. BARTRAMIACEÆ. 
186. Bartramia ityphylla (Haller) Brid. 


Very common. 

One of the most frequent mosses, both on a dry and on a some- 
what damp substratum, from the lowlands to far up in the Alpine 
region. It grows in soil-filled rock-clefts, on dikes and house-walls, on 
peat and on the top of knolls in bogs. As a rule, it grows in small, 
unmixed tufts, but occasionally also mixed with other mosses. The 
fruit, which ripens during August, occurs plentifully everywhere. 


187. Plagiopus Æderi (Gunn.) Limpr. 
Bartramia (Ederi (Gunn.) Sw. 


E. Iceland: Berufjérdur!, fr. N. Iceland: Håf near Eyjafjordur (O. 
DJ)FEReistarar si (OD) FEE 

Near Berufjérdur it grew abundantly on a wet rock-face with a 
northern exposure, in cushions, about 3 cm. high, with numerous cap- 
sules which were ripe at the end of June. 


188. Philonotis fontana (L.) Brid. 


Very common over the whole of Iceland. 

One of the most frequent and abundantly occurring species which 
grows everywhere on a wet substratum, in marshes and moss bogs, 
along streams, and on wet rocks, but is also common on a more dry 
substratum, for instance in grass fields or on humus-covered rocks. It 
is of almost equal frequency at- all elevations upwards to the limit of 
vegetation. The fruit, which is found very frequently, ripens in the low 
land about August ist. 

Philonotis fontana is extremely variable, and conditions pertaining 
to dampness especially exercise great influence over the appearance of 
the plant. The usual bog forms are highly tomentose, with erect or 
more or less falcato-secund leaves (f. falcata Warnst.). On a more dry 
substratum the tufts usually become denser and more highly tomentose, 
and the plants more slender. 

In the cold water in moss bogs and on inundated ground along 
rivers where the temperature during summer is often only 49—69% this 
species, in association with Pohlia albicans var. glacialis, Brachythecium 
rivulare and species of Mnium, often forms the main part of the vege- 
tation. Here it forms very high and loose, in fact hardly cohering, 
tufts, which often resemble P. seriala in habit. The stems are flaccid 
and hardly tomentose; the leaves are short and appressed or slightly 
secund, and often obtuse or cucullate at apex. The nerve is very broad 
and widened at the base. Such cold-water forms agree exactly with 
Loeske's P. fontana-adpressa (L. Loeske, Kritische Bemerkungen uber 
einige Formen von Philonotis, Hedwigia, vol. 45, p. 100, and Kritische 
Ubersicht der eur. Philonoten, Hedwigia, vol. 45, p. 195). That it is the 
cold water which checks the growth of the radicles and causes the 


THE BRYOPHYTA OF ICELAND 495 


characteristic form of the leaves and the looser cell-tissue is seen from 
the fact that, in situations where the water-level is altered, forms may 
be found growing in tufts, in the lower part of which the stems are 
tomentose and” the leaves normal, while all the young shoots which 
have developed after the habitat has been inundated have the typical 
depressa appearance. Likewise are found all possible transitions between 
P. fontana and the depressa form. 

On mountain heights, where the snow lies long, the slopes are often 
covered with f. nigrescens Loeske, with black, prostrate and hardly to- 























Fig. 8. Philonotis fontana (L.) Brid. Slender form from warm ground 
(Thorlåkshver; nat. size). 


mentose stems and light yellowish-green annual shoots. This form has 
beyond doubt originated from the pressure of the snow-covering. The 
leaves are, as a rule, shortly pointed. Where the water from the melted 
snow remains and soaks the tufts the leaves become still more shortly 
pointed and the cell-tissue looser (f. borealis, Hagen, Loeske). By hot 
springs P. fontana often occurs abundantly both in the lukewarm water 
and on the warm clayey flats. Here it forms very soft, not cohering 
tufts. The stems are erect and very slender; the leaves are small, often 
strongly falcato-secund, longly pointed, highly papillose, and sharply 
toothed along the margin, with narrow (0.05—0.06 mm. broad) nerve. 
Note: "P. calcarea from Frodarheidi (leg. H. J.) is! P' seriata. 


189. Philonotis Arnellii Husnot. 


N. Iceland: Hof near Eyjafjérdur (0. D.)!. SW. Iceland: Thingvellir!; 
between Reykjavik and Hafnarfjérdur!. 


496 A. HESSELBO 


Near Thingvellir it grew on a soil-covered rock-ledge in a cleft, 
in association with Eucalyx subellipticus, Bartramia ityphylla and Pohlia 
cruda. All the specimens found were gg plants. 


190. Philonotis seriata (Mitt.) Lindb. 


E. Iceland: Berufjérdur!; Seydisfjordur!; Kirkjubål (H. J.)!. N. Ice- 
land: Husavik!, at an elevation of 200—300 metres; several places near 
Eyjafjérdur (0. D.;!): Myvatn (Grl.)!; common in Oxnadalur!. NW. Ice- 
land: Very common by all the fjords from Dyrafjéårdur northwards!. 
W. Iceland: Frodarheidi (H. J.)!; Esja, many places!. S. Iceland: Selja- 
land (Stp.)!. 

Grows on inundated ground, as a rule in the water itself, for in- 
stance along streams and in moss bogs. 

In NW. Iceland it is one of the most frequently occurring species, 
and is met with abundantly up to a height of above 400 metres above 
sea-level. It often forms in association with Pohlia albicans, Scapania 
uliginosa, Haplozia cordifolia and Chiloscyphus polyanthus v. fragilis the 
main portion of the moss-carpet along streams. In the other parts of 
the country — with the exception of S. Iceland where it has been found 
only near Seljaland —- it is rather common. It is usually met with from 
about 250 to 500 metres above sea-level, but frequently ascends up to 
about 600 metres, for instance by Berufjordur. The fruit, which was 
quite green even in the first half of July, has been found only in a few 
localities in the district of Isafjordur and near Eyjafjordur, and only 
scantily. 


191. Philonotis tomentella Mol. 
P. alpicola Jur. 


Widely distributed over the whole of Iceland. 

P. tomentella usually prefers drier localities than does the closely 
allied P. fontana. In N. Iceland, where it is very common, it grows 
abundantly everywhere on partially dry, grass-covered ground, and is 
easily distinguished from P. fontana by its more slender growth and by 
the extremely dense tufts, with stems covered with brown tomentum 
almost to their summit. It grows, also, both on rather dry and on 
damp rocks and on wct boggy ground in company with P. fontana, with- 
out, however, extending into the water itself. 

It varies very considerably in habit and size. The leaves are more 
or less falcato-secund, and forms especially from dry rocks have leaves 
strongly falcato-secund (f. falcata). In shady localities, for instance in 
rock-clefts and caves, slender to almost thread-like forms (var. capillaris) 
are frequently met with. Forms growing in bogs are more vigorous, 
than are those growing on a dry substratum, and are also less densely 
tomentose. 

On damp rocks and in humus-filled rock-clefts there frequently oc- 
curs a slender form with numerous slender, easily falling branches which 
probably serve for vegetative propagation (f. flagellifera). 

Limpricht (Kryptogamenflora, vol. II, p. 573) records that the 


THE BRYOPHYTA OF ICELAND 497 


cortex in this species consists of 3—4 layers of sharply defined, small 
stereids covered with a sphagnoid outer cortex, while P. fontana has not 
a sharply differentiated cortical layer. This feature appears, however, 
to be closely connected with the degree of dampness of the habitat, so 
that forms from a dry substratum, both of P. tomentella and of P. fon- 
tana, have several layers of thick-walled cortical cells, while bog-forms 
have thin-walled cortical cells, and (as in Dicranum scoparium and D. 
angustum) all possible transitions may be found.between thin-walled and 
thick-walled cortical cells. 

Fruit occurs very frequently and ripens at the end of July or in 
the beginning of August. 


Fam. TIMMIACEÆ. 
192. Timmia norvegica Zett. 


E: Iceland: Hof!: Vallanes (H. J.)!. N: Iceland: Pollar (St.)!: Akureyri!, 
at an altitude of 300 metres; Hof near Eyjafjordur (0. D.)!; Vidimvri 
(Grl.)!. 'S. Iceland: Vestmannaev!. 

It grows on earth and in rock-clefts, but has been collected every- 
where only scantily. Near Hof (E. Iceland) it grew on gravelly ground 
in association with Oncophorus virens, Ditrichum flexicaule, Myurella 
Julacea and M. apiculata at an altitude of about 150 metres. Near 
Akureyri it was found sometimes in rock-clefts, in company with Mnium 
orthorrhynchum, sometimes on rocks mixed with Philonotis tomentella. 
Only sterile specimens have been found. 


193. Timmia austriaca Hedw. 


Very common especially in N. and E. Iceland, but more rare in 
NW. Iceland (Dyrafjårdur, Isafjéordur, Grunnavik!). It grows on earth 
or on soil-covered rocks, and often in masses. Thus in several places 
in Hallormstadaskogur it was the most abundant member. of the moss- 
carpet under the birches, especially on knolls and soil-covered stones. 
In several places in N. Iceland. for instance in Oxnadalur, it covered 
the slopes along streams, often greatly mixed with other species. 

Timmia austriaca is most frequent in the lowlands up to a height 
of about 300—400 metres, but is also often found on mountain heights 
(for instance on Seljaland at an altitude of about 620 metres) although 
usually in small quantity. Fruit has been found only near Tverå and 
Vidimvyri in N. Iceland; it occurred plentifully in both places and had 
just ripened at the end of July. 


FxMn SPOLEYTRICHACEÆ: 
194. Catharinea undulata (L.) Web. 


E. Iceland: Hornafjérdur!; Lon!; Berufjordur!. N. Iceland: Hof near 
Eyjafjordur (0. D.)!: Reykir near Svinavatn (Grl.)!. W. Iceland: Reykja- 


498 A. HESSELBO 


vik (Mérch;!); near hot springs in Reykholtdalur (Grl.;!); Borgarfjérdur!. 
Common in SW. and $S. Iceland!. 

Commonly distributed in SE… S. and W. Iceland from Lon in the 
south-east to Borgarfjoérdur in the west. In N. Iceland it has been found 
only near Eyjafjårdur, and it has not been collected in NW. Iceland. 
It grows sometimes on peat, sometimes on damp clayey ground, for 
instance by sides of ditches, and is occasionally found in fruit, which 
about Reykjavik was quite undeveloped even in August. It is a de- 
cidedly low-land species, and has not been found at a higher elevation 

















Fig. 9. Catharinea undulata (L.) Web. var. thermophila (nat. size). 


ihan 50—100 metres above sea-level. The usual forms of this species 
are low in growth, 1—2 cm. high, with leaves which are as a rule 
4—6 mm. long, narrowly lanceolate, obtuse or shortly pointed, only 
slightly undulate with a few spines at the back, and the margin often 
with single teeth. 

Catharinea undulata is one of the species which is found most fre- 
quently and abundantly on warm ground, where it occurs in rather 
divergent forms. Usually it resembles the southern woodland forms - 
with long, strongly undulating leaves, very rough at the back. AÅ very 
peculiar form is 

Var. thermophila n. var. 

Loose cushions as much as 10 cm. high. Leaves evenly distributed 
along the whole length of the stem, or somewhat denser at the top, 
often falcato-secund, 6—7 mm. long, linear-lanceolate, sharply pointed, 
with 4—5 lamellæ at the back. Lateral shoots are often developed along 


THE BRYOPHYTA OF ICELAND 499 


the stems. This form grew abundantly near the hot springs about 
Skålholt, especially on the slopes stretching down towards the outlets 
of the springs. 


195. Oligotrichum hercynicum (Ehrh.) Lam. 
Oligotrichum incurvum (Huds.) Lindb. 


Iceland (Morch). E. Iceland: Seydisfjardarheidi!. N. Iceland: Ljosa- 
vatn!, common 350 metres above sea-level and upwards; Askja (Caroc)!. 
NW. Iceland: Very common!. W. Iceland: Stadarfell (Stp.); Esja!, at an 
altitude of 400—500 metres; Reykholtdalur (Grl.;!); Hafnarfjordur!. S. 
Iceland: Common from about 350 metres and upwards!; Krisuvik (Stp.)!; 
everywhere near hot springs!. 

This species has a very peculiar distribution in Iceland. It has 
its main area of distribution on the mountain heights, where it doubt- 
less occurs over the whole of Iceland, and often in masses. It grows 
here on damp gravelly flats, especially in the neighbourhood of the 
snowW-patches, in association with Anthelia Juratzkana, Pleuroclada albes- 
cens, Pohlia gracilis, Polytrichum sexangulare, etc, and as a rule sets 
fruit.  Scarcely anywhere has it been found until at a height of 350— 
400 metres, and was most abundant at about 500—700 metres above 
sea-level. Only in NW. Iceland, where it is exceedingly common, did it 
descend to a lower altitude, in many places as low as to the sea-level. 
In the greater part of the country it is quite absent from the low land; 
in SW. Iccland it has been found only scantily in a lava cave near 
Hafnarfjordur. 

This species has another area of distribution near the hot springs 
in SW. and W. Iceland. Here it has been collected near a great number 
of springs in Reykholt- and Reykirdalur, around Skålholt and in several 
other places, where it often occurs abundantly on the warm clayey flats, 
but only sterile. These warm-soil forms differ somewhat from the Alpine 
forms. As a rule they are quite low in growth, about 5—10 mm. high, 
with softer leaves, twisted or incurved when dry, with a few (5—6) 
lamellæ at the back. The leaf-cells are more thin-walled, chlorophyllous, 
larger and more regularly quadrate and transversely elongated, usually 
0.015—0.018 mm., but occasionally also 0.023 mm. in diameter, shortly 
rectangular at the base, 0.018 mm. broad. The nerve is often only half 
as broad as in the Alpine form. Such forms grow especially on the 
warm clayey flats, in company with Anthoceros, Fossombronia, Haplozia 
crenulata and other hepatics. Near Deildartunguhver in Reykholtdalur 
it grew on the warm ground among other mosses, in tufts about 3 cm. 
high. Forms exactly resembling the Alpine forms occur also frequently 
on the warm ground. 

In the majority of the localities in NW. Iceland the fruit was ripe 
in the latter half of June, on the mountain heights early in July. 


500 A. HESSELBO 


196. Psilopilum lævigatum (Wahilb.) Lindb. 


Oligotrichum glabratum Lindb. 


O. lævigatum Br. eur. Psilopilum arcticum Brid. 


E. Iceland: Hornafjérdur!; Berufjérdur!; Seydisfjårdur!. N. Iceland: 
Husavik!; Akureyri!; Hof near Eyjafjordur (0. D.)!:- Geitaskard!. NW. 
Iceland: Isafjordur!; Snæfellsstréond!. W. Iceland: Stykkisholmur!; Stadar- 
fell (Stp.)!; Hafnarfjordur!; Reykjavik (Morch; Ho.; Stp.;!). S. Iceland: 
Common in Olfus?. 

Commonly distributed everywhere near the coasts. It grows there 
on rather dry peaty ground, and often in great abundance. For instance 
near Reykjavik it covers in association with Dicranella crispa, Bryum 
pallens, Scapania curta, etc. large tracts of ground on the peninsula. It 
appears to thrive more particularly in drained bogs whence peat is being 
cut; thus around Husavik, Akureyri and Isafjordur it occurred in very 
great abundance, partly on the ground and partly on the piled up heaps 
of cut peat, which were often quite covered by it. Funaria hygrometrica 
and Pogonatum urnigerum often occurred abundantly in company with 
it. Fruit ripens in the beginning of (Reykjavik) or at the end of June 
(N. Iceland). 

Note. F. Hagen in his preliminary works on a Frondose-Moss Flora 
of Norway (XIX, Polytrichaceæ) supposes that Psilopilum tschuctschicum 
(Mulle Hak)EParshasibeenicolleeteds mæcen de by WÆRRER 0 oe TEA 
the specimens I have had an opportunity of investigating, both numerous 
older specimens collected by Morch, Krabbe, Groånlund and others 
and my own specimens collected in a considerable number of habitats 
from all parts of Iceland, belong to Psilopilum lævigatum, which is doubt- 
less the only species of this genus found in Iceland. 


197. Pogonatum polytrichoides (L.) Brockm. 
Pogonatum nanum (Schreb.) P. B. P. subrotundum Lindb. 


Iceland (Ho.; Morch)!. W. Iceland: Reykjavik (Grl.;!); "Stykkisholmur 
(Stpy)"FSletta (Sp) Finer district or Borsarfjordur FEST and RN earn 
the Thjorså (Stp.)!; frequent in Olfus!. 

This species was found only in SW. and W. Iceland, but was rather 
common in both places on somewhat damp and especially peaty soil, 
for instance by the sides of ditches and along roads. In the district 
of Borgarfjordur it was common along roads. Around Reykjavik it was 
found everywhere by the sides of ditches. Fruit was found everywhere, 
the capsules were partly old specimens from the previous year and 
partly such as were quite young even in August. 

Note. Pogonatum aloides (Hedw.) is recorded to have been collected 
by Steenstrup near Sletta, Stykkisholmur and the Thjorså, but all the 
specimens belong to P. polytrichoides. 


THE BRYOPHYTA OF ICELAND 501 


198. Pogonatum dentatum (Menz.) Brid. 
Var. minus (Wahlb.) Hagen. 
Reykjavik !. 
Here it grew in several places on peaty soil, and in one place very 
abundantly, with numerous capsules, some of which were almost ripe 
in the first half of August. 


199. Pogonatum urnigerum (L.) P. B. 
Polytrichum urnigerum L. 


Widely distributed over the whole of Iceland. 

Grows usually scattered as individual plants, or a few plants to- 
gether, among other mosses on dry, humus-covered rocks or on the 
ground, both dry and somewhat damp: for instance along the banks of 
streams. It occurs in large quantities on peat, often forming large light- 
green patches, interwoven with Alicularia scalaris and, occasionally, Sca- 
pånia curta. On somewhat damp, moss-grown slopes it often forms 
large tufts several centimetres high among Hylocomium spp., Hypnum 
uncinatum and Polytrichum alpinum; such vigorous forms are also fre- 
quently met with in South Iceland on somewhat damp ground, for in- 
stance by the sides of ditches. It is most frequent in the lowlands and 
does not appear to ascend above 300—400 metres. The fruit, which 
'only occurs on large plants growing in tufis, was ripe or overripe in 
the beginning of June. 


200. Polytrichum alpinum L. 


Very common everywhere both» on dry and on somewhat damp 
ground, and almost equally frequent from the sea-level up to the limit 
of vegetation on mountain heights. It grows both mixed with species 
of Hylocomium and interspersed in Rhacomitrium-mats on knolls in bogs, 
on peat, in grass-fields and on the stony slopes of mountain heights. 
The fruit, which occurs very frequently, ripens in the lowlands about 
August Ist. 

Varies considerably in habit, length of leaf, form of capsule, etc., 
but the majority of the forms, however, approach closest to the type. 
Forms agreeing more or less with var. septemtrionale (Sw.) Brid. are 
also common. 


201. Polytrichum formosum Hedw. 


W. Iceland: ”Sæluhusid” in Såkkålfdalur (F.)!, sterile. 


202. Polytrichum gracile Dicks. 


Rather common over the whole of Iceland. 
This species grows in peaty soil, often abundantly, and sets fruit 
almost always. It occurs only in the low land and has hardly been 


502 A. HESSELBO 


found at a higher level than 200—300 metres. Near Akureyri the fruit 
was ripe at the end of July. 


Var. anomalum (Milde) Hagen. 


Hof near Eyjafjordur (0. D.)!. NW. Iceland: Hesteyri!. 

In both these places it was collected scantily only. Near Hof it 
occurred as a few scattered individuals in a specimen of the vegetation 
taken from wet boggy ground. Near Hesteyri it grew on very wet boggy 
ground. The plants were about 3 cm. high and sterile. Some of them 
agreed with var. anomalum; in some the leaves at the base of the stem 
resembled in structure those of the variety in having thin-walled cells 
0.020—0.025 mm. in size, while the leaves in the upper part of the 
stem resembled in structure those of the type in having cells 0.015— 
0.020 mm. in size. 

Var. anomalum is undoubtedly a hygrophilous form of P. gracile, 
which develops when the habitat is inundated. 


203. Polytrichum sexangulare Floercke. 


Iceland (Mérch)!. N. Iceland: Ljosavatn!; Reykjaheidi!; common near 
Eyjafjordur (0. D.;!).. NW. Iceland: Very common!. W. Iceland: Esja!; 
Kolvidarhol!. S. Iceland: Seljaland!; Holt!; Thingvallahraun!. 


Var. vulcanica C. Jens. 


"Seta perbrevis usque ad 4 mm. longa; apophysis valde indistincta; 
capsula non angulata, vetusta nigrescens et irregulariter 4—5-gona ; 
Cetero typo simile.” 


S. Iceland: The volcano on the road to Holt (Stp.)!. 


Var. tenellum n. var. . 

Plants 1—2 cm. high, more slender than the type, with shorter, 
narrower, more slightly incurved and not secund leaves, when dry; the 
nerve excurrent in a short mucro; otherwise exactly like the type. 

S. Iceland: Barkarstadr!; Drångshlid!; Seljaland!. 


Polytrichum sexangulare has a very peculiar distribution in Iceland. 
The type is a decidedly Alpine plant which does not feel quite at home 
until near the snow line, on the gravelly flats soaked by the melting 
snow. In N. Iceland it has been collected only at elevations of above 
500 metres. In NW. Iceland, where the species is very common, it is 
met with everywhere from about 200—300 metres upwards. On northern 
slopes where the snow lies long it sometimes descends as far down as 
to the sea-level. Near Kolvidarhol it was common at about 400 metres, 
and on Esja everywhere at elevations of above 500 metres. 

P. sexangulare has not yet been collected in E. Iceland, but the 
reason can hardly be that.it does not occur there, it must rather be 
due to the fact that the mountain heights were almost everywhere snow- 
covered at the timc of the year (June, 1909) when there was an op- 
portunity of making collections there. This species has an area of dis- 
tribution not only on the mountain heights, but also in the lowlands. 


THE BRYOPHYTA OF ICELAND 503 








a a a a b b b 
Fig. 10. Polytrichum sexangulare Floercke var. tenellum. 


a, in a damp. and b, in a dry condition (X 6). 


In lava clefts near Thingvellir and in deep lava clefts on Reykjarheidi 
(at an altitude of about 250 metres) it was found abundantly in several 
places in association with several other Alpine species, for instance 
Gymnomitrium concinnalum, G. varians, Pleuroclada albescens and Pohlia 
commutata. The occurrence of this species in such localities is un- 
doubtedly connected with the fact that there the conditions pertaining 





Fig. 11. Polytrichum sexangulare Floereke var. tenellum. 


a, b and c, leaves (X 15); d, leaf-apex (X 100). 


504 A. HESSELBO 


to soil present certain points of resemblance to the mountain heights, 
as the snow in the deep clefts, the bottom of which is never reached 
by the rays of the sun, often remains till late into the summer. The 
occurrence of var. fenellum is more peculiar. It has been found only 
in S$. Iceland, where it grew exclusively on dry or slightly damp blocks 
of tuff in the lowlands. But only near Seljaland did a few small tufts 
occur on a block of tuff at an altitude of about 600 metres. Near 
Drångshlid it grew abundantly on the fallen blocks of rock at the foot 
of the rock-wall behind the farm. There in company with Hypnum 
cupressiforme, Ceratodon, Didymodon rubellus, Rhacomitrium fasciculare 
and Bryum argenteum it formed tufts of varying size on the top of the 
blocks in dry situations. Near Barkarstadr it was found on a some- 
what damp tuff-face, and creeping over fallen blocks of rock, in asso- 
ciation with Pohlia cruda, P. grandiflora, Anæctangium compactum, Bry- 
oxiphium norvegicum and Anthelia spp. The plants from these habitats 
were quite sterile, and exactly alike in all respects. 

The fruit, which is common in the Alpine form, ripens in August 
(Akureyri: ripe Aug. 13th; near Isafjérdur: green in the first half of July). 


204. Polytrichum piliferum Schreb. 
Polytrichum pilosum Neck. 


Iceland (Morch)!. E. Iceland: Vallanes (H. J.;!).  N. Iceland: Rey- 
kjahlid (Grl.;!).  NW. Iceland; Bildudalur (C. Hansen;!); Gnupr near Dvra- 
fjordur!; Årmuli!: Arngerdareyri!. VW. Iceland: Stykkisholmur (H. J.;/); 
Grund in Skorradalur!; common near Reykjavik!; Videy!; Hafnarfjordur 
(Wiinstedt)!. S. Iceland: Krisuvik (Stp.;)!; Drångshlid!. 

Occurs here and there, and is rather rare in the majority of the 
districts of Iceland. It grows on dry sandy ground, upon soil-covered 
rocks and on dikes, often singly among other mosses. It is absent from 
or rare in the most rainy parts of the country, for instance at the East 
coast and in South Iceland and is most frequent in those districts where 
the rainfall is least. Thus it occurs abundantly near Mvyvatn, where it 
is common in the lava-field. It is also rather widely distributed in 
NW. Iceland, here it is often found in abundance, for instance on sandy 
fields near Årmuli and on rocks near Arngerdareyri. Fruit is common 
and ripens in the latter half of June. 


205. Polytrichum juniperinum Willd. 


Commonly distributed. 

It grows on more or less dry ground, for instance on knolls in 
bogs, in the Rhacomitrium-canescens heath, on slopes among Hylocomium 
and Rhacomitrium etc., usually scattered among other mosses, more 
rarely in cohering tufts, and almost always sets fruit which ripens. at 
the end of June and in the beginning of July (Seydisfjordur, July 6th; 
Isafjordur, June 24th). 

It is most widely distributed in the lowlands and does not appear 
to ascend much higher than about 400 metres above sea-level. 


THE BRYOPHYTA OF ICELAND 505 


206. Polytrichum strictum Banks. 


E. Iceland: Hornafjérdur, common!. N. Iceland: Husavik!; Akureyri, 
common to about 600 metres above sea-level!; Hof near Eyjafjérdur (O.D.)!; 
common from Geitaskard westwards to Hrutafjérdur!. NW. Iceland: 
Common around Isafjérdur!; Grunnavik!; Kaldalon!. W. Iceland: Grund 
in Skorradalur!; Olafsdalur (H. J.)!; Reykholtdalur, common!; Reykjavik!. 
S. Iceland: Skålholt!, $; Geysir (Stp.)!. 

Occurs rather commonly and often abundantly on wet boggy ground, 
but always sterile. In Reykholtdalur it also grows on warm ground. 


207. Polytrichum commune L. 


Widely distributed on damp ground, both in bogs and along the 
banks of streams, but as a rule rather scantily. 

In the low land, where P. commune has its greatest distribution, 
the large coarse form is usually met with. In higher regions, where it 
is found up to a height of about 500 metres, it usually becomes slen- 
derer and lower in growth. Thus, near Akureyri, it occurred abundantly 
in a bog at an altitude of 500 metres as a low-growing, slender form. 
mixed with Hypnum stramineum. 

P. commune is one of the most frequent species near hot springs, 
where it grows sometimes mixed with Sphagnum and is then large and 
vigorous, sometimes in extensive mats interwoven with hepatics and is 
then usually low in growth with shorter leaves. On a warm substratum 
it often fruits freely, while it has or else been found in fruit only near 
Grunnavik in NW. Iceland. 


Var. fastigiatum (Lyl.) Wils. 
SW. Iceland: Svinahraun!, abundantly on damp soil at the edge of 
the lava-field. 


208. Polytrichum Swartzii Hartman. 


NW. Iceland: Bæir!, sterile. 


Var. nigrescens (Warnst.) Hagen. Syn. Polytrichum inconstans Hagen. 

NW. Iceland: Thoroddstadaengjar (St.)!; Dynjandi!. 

The type grew on a damp slope in not-coherent tufts, about 15 cm. 
high; the plants were quite typical with sharply toothed leaves, and 
with lamellæ, the marginal cells of which had the irregular form peculiar 
to P. Swartzii. The variety grew on very wet or inundated ground in 
company with Hypnum stramineum, H. giganteum and Mnium cinclidioides 
similarly to P. gracile var. anomalum, and may be regarded as a form 
similar to the latter; as in aquatic forms of other species, both forms 
have flaccid-soft leaves with slightly developed stereom in the nerve and 
broader, slightly toothed leaf-margin; in the plant from Dynjandi the 
leaf-margin was almost entire. 


The Botany of Iceland. Vol. I, part ll. : 33 


506 A. HESSELBO 


Fam. BUXBAUMIACEÆ. 
209. Diphyscium sessile (Schmid.) Lindb. 
Webera sessilis Lindb. 


Common in East, South and West Iceland, and rather frequent in 
North-west Iceland, where it is common for instance in the district of 
Dyrafjoérdur. In North Iceland it has not been found east of Blåndudalur. 

It grows on firm, bare ground, especially on the top of knolls 
or on small elevations of earth on slopes, where it forms low, dense 
cushions, dark brown in colour; it is common, at any rate in E. Iceland, 
as far upwards as 600—700 metres above sea-level. Fruit, which oc- 
curred only scantily in all places, was collected near Reykjavik, near 
Lundur in West Iceland and near Geitaskard in Bléåndudalur. 


Fam. FONTINALACEÆ. 
210. Fontinalis antipyretica L. 


Found in all parts of the country and common on stones in brooks 
and rivers, and sometimes also in stagnant water, up to about 300—400 
metres above sea-level. Near Ljosavatn in N. Iceland it was found 
abundantly in a river even at an altitude of 300—400 metres, and near 
Isafjordur it occurred frequently as far upwards as 300 metres. It 
varies considerably in size, leaf-form and colour. In stagnant water 
(thus in very wet bogs near Jokulsfjordur) it becomes very robust, light 
brownish-yellow, with broad leaves. 

In swiftly flowing water there often occur slender, dark-green or 
brownish-green forms with strong metallic sheen, which agree most 
closely with var. montana H. Muller. In such forms the basal angles 
of the leaves are often somewhat concave, and formed of a single layer 
of cells, in which respects it comes very near to F. gracilis. This species 
is recorded from several localities by Gronlund, H. Jonsson and 
Feddersen, but all the specimens which have been investigated belong 
either to F. antipyretica or to F. androgyna. Only sterile specimens have 
been found. 


211. Fontinalis islandica Cardot. 


E. Iceland: Faskrudsfjordur (Jardin, 1865) (specimens not inspected). 


212. Fontinalis longifolia C. Jensen (Bot. Not. 1885, p. 83). 


Gracilis, viridis et flavoviridis, elongata, mollis, rubricaulis, ramulis 
brevibus, patulis subremotis. Folia laxa, imbricata vel erecto-patentia 
mollia, carinata, annosiora, bifida, ovato-lanceolata, sensim longe acumi- 
nata, integra vel ad apicem lenissime serrulata, decurrentia, cellulis 


THE BRYOPHYTA OF ICELAND 507 


flexuoso-linearibus, apicalibus brevioribus, angulorum multo majoribus, 
rectangulis. Folia ramulorum angustiora. Flores et fructus ignoti.” 

This species, which was first found by Feddersen in the Helgå 
in S. Iceland, grew very abundantly on stones in the Reykirdalså in SW. 
Iceland. This river has copious influx from the hot springs, so that, in 
the first days of June, the temperature of the water was about 12", 
which was essentially higher than in the majority of the other rivers 
where the temperature was, as a rule, only about 47—6”, 

















RER RE 


Fig. 12. Fontinalis longifolia C. Jens. (Reykir; somewhat reduced). 


Fontinalis longifolia agrees most closely with F. hypnoides R. Hartm. 
The plant from the Reykirdalså is rather robust, as much as 25 cm. 
long, very strongly and irregularly branched, with rhizoid-cushions around 
all the points whence the branches issue, and with plane or somewhat 
concave, longly tapering, very soft leaves, which only exceptionally are 
folded- along the median line. The gg plant is more slender than is 
the 2 plant, with very numerous, narrowly ovate dg flowers, which often 
occur in clusters of 2—4 on the main axis and branches and contain 
each 2—4 antheridia. The fruit, which in the beginning of June was 
developed to almost full size, but was as yet quite green, so that the 
peristome and spores could not be investigated, was found scantily on 
the lowermost part of the stem. As in F. hypnoides the capsule was 
half-exserted. 


33% 


508 A. HESSELBO 


213. Fontinalis thulensis C. Jensen (Bot. Tids. 20, p. 110). 


<Laxe caespitosa, sat robusta, mollis, sordide luteola, inferne nigres- 
cens subnitida. Caulis usque ad 25 ctm. longus, inferne plerumque 
nudus, plus minusque ramosus, ramis brevibus vel elongatis, erecto- 
patentibus, vel subsecundo-arcuatis, interdum acutis, cauli primario sat 
similibus. Folia dimorpha, tristicha; folia caulina erecto-patentia vel 
laxe imbricata, plus minusque distincta et recte carinata, interdum tantum 
complicata, late ovata-lanceolata, longe decurrentia integra vel apice in- 
distincte denticulata, adultiora subobtusa, juniora acuta, nunquam bi- 
fida, superiora vulgo 5—6 mm. longa et 2—3 mm. lata, inferiora minora, 
latere uno versus basim reflexo; folia ramulina minora et subangustiora, 
vulgo 3—4 mm. longa et 1,5—2 mm. lata, inferiora multo minora omnia 
ecarinata, valde concava, interdum complicata, erecto patentia, inferiora 
ramorum arcuatorum arcte imbricata, superiora interdum subsecunda. 
Cellulae in media foliorum 0,007—0,013 mm. latae, 8$—12 plo longiores. 
Alae basilares, planae, unistratae, e cellulis quadratis et hexagonis, aeque 
ac ceteris cellulis basilaribus, luteis formatae. Cetera ignota. 

Speciei americana F. Kindbergii valde affinis. sed ramis erecto-pa- 
tentibus, non pennatis, foliis eorum brevioribus et latioribus, cellulis 
angustioribus, colore luteolo (F. Kindbergii ferrugineus est) et nitore 
debiliore.” 

W. Iceland: The Laxå (Hjardarholt), 21.7.1886 (Feddersen). 

Note.  Fontinalis squamosa L. is enumerated on older lists, but of 
the specimens from Iceland, contained in the collections, some belong 
to F. antipyretica, some to F. androgyna. 


214. Fontinalis androgyna R. Ruthe. 


WiTceland:-The”Ellidarå”near "Reykjavik (Morceh- Gr HS] 
In the Ellidarå it grew abundantly in company with F. antipyretica 
on large stones and rocks under water. There not only ej buds were 


found, but also a few ripe capsules on the lowermost part of the stem 
(98831909) 


Fam. CRYPHÆACEÆ,. 
215. Leucodon sciuroides (L.) Schwgr. 


S. Iceland: Foss (H. J.)!; Drångshlid (H. J.;!); Stjornarsandur (H. J.)!; 
Hrutafell!. SE. Iceland: Hornafjordur!: Hof!. SW. Iceland:  Hafnar- 
fjordur!. 

All the Iceland specimens belong to var. morensis (Schwågr.) de 
Not. It grows on dry rock-walls with a southern exposure, both on 
tuff and on basalt, and appears to be rather frequent from Hof in SE. 
Iceland throughout S. Iceland to Hafnarfjéordur in SW. Iceland. It occurred 
abundantly on Hrutafell and Drångshlid, and covered the vertical faces 
of tuf£frocks for long distances. Here it was also richly fruiting with 
capsules which were partly ripe at the end of June. 


THE BRYOPHYTA OF ICELAND 509 


216. Antitrichia curtipendula (Hedw.) Brid. 


E. Iceland: Hornafjérdur, common!; Djupivogur!; Hof!; Starmvri!; 
Berufjordur!. N. Iceland: Moådruvellir (Thåroddsen)!; Geitaskard!; com- 
mon in Vatnsdalur!. NW. Iceland: Patreksfjordur!; Kaldalon!. Common 
in W. and SW. Iceland!. 

Common in SE. Iceland as far as Berufjérdur (where it was 
found in one locality only), in S. and W. Iceland and the western part 
of N. Iceland from Bléndudalur. Rare and scanty in NW., N. and E. 
Iceland. It grows usually on a rather dry substratum, on stony slopes, 
and soil-covered rocks, but especially on moss-grown slopes in company 
with Hylocomium spp., and on the ground in coppices. In Esja it is 
common up to a height of 300—400 metres above sea-level. Only sterile 
specimens have been found. 


Fam. NECKERACEÆ. 
217. Neckera complanata (L.) Hub. 


W. Iceland: Gilsbakki!:; Bardshellir (Grl.)!. S. Iceland: Paradishellir 
(Stp.)!; Reykir!; Austarhlid near Geysir!; Breidabolstadr!; Barkarstadr!; 
Holt!; Drångshlid!; Hrutafell!. 

Rare in W. Iceland, and rather common in S$S. Iceland. It grows 
everywhere on the roof and sides of dry caves, which it often covers 
with its extensive, dark-green mats. Only sterile specimens have been 
found. 


Fam. LESKEACEÆ. 
218. Myurella julacea (Will.) Br. eur. 


Very common on rocks, especially in humus-filled clefts ; occasion- 
ally also on the ground, for instance on knolls in bogs. It often occurs 
in unmixed cushions, but far more frequently mixed with other mosses, 
and is met with both on a dry and on a somewhat damp substratum. 
On wet rocks it often grows in association wit1 Anæctangium Mougeottii, 
Philonotis, Blindia acuta, etc.; in dry clefts in company with Mnium or- 
thorrhynchum, Plagiothecium Roesei, Bartramia ityphylla, Pohlia cruda, 
etc.: or it may be met with on dry rocks woven into the tufts of Hyp- 
num revolutum and Grimmia spp. It does not appear to ascend higher 
than 300—400 metres. Only sterile specimens have been found. 


219. Myurella tenerrima (Brid.) Lindb. 
Myurella apiculata (Huben.) Br. eur. 
Grows quite similarly to the preceding species and, as a rule, in 


company with it, and in the majority of the districts of Iceland is 
almost as common. In S. Iceland it appears, however, to be somewhat 


510 A. HESSELBO 


rarer. It is most frequent in rather damp situations, for instance on 
rocks near waterfalls. Only sterile specimens have been found. 


220. Leskea nervosa (Schwågr.) Myrin. 


SE. Iceland: Hof!, on rocks in company with Metzgeria furcata, 
sterile. S. Iceland: Drångshlid!; Hrutafell!, sterile. In both the latter 
localities it occurred abundantly on dry tuff-faces or on large fallen 
blocks. 


221. Leskea catenulata (Brid.) Mitten. 


Nupsdal (Stp.)!. 
Here, Gnupsdalur near Dyrafjérdur in NW. Iceland is probably meant. 


222. Anomodon viticulosus (L.) Hook. et Tayl. 


S. Iceland: Hrutafell!; cave near Skågafoss!.  Vestmannaey!. 

On Vestmannaey it grew at the foot of a dry, stony slope below 
Stora Klit; near Hrutafell it grew sometimes on rocks, sometimes on 
soil-covered rock-ledges. Only sterile specimens have been found. 


223. Pterigynandrum filiforme (Timm) Hedw. 


Very common everywhere. 

The most frequent form is var. decipiens (W. et M.) Limpr., which 
occurs abundantly everywhere on dry rocks, especially on perpendicular 
hasalt-faces, where it often forms very large cushions 5—6 cm. deep. 
Here all transitional forms are also met with between the type and the 
variety. The type is developed in deeper shade and occurs especially 
in clefts and crevices, where it forms thin. adherent mats on the sur- 
faces of stones. Only sterile specimens have been found. 

Both forms are common up to about 300—400 metres above sea-level. 


224. Lescuræa decipiens (Limpr.) 
Plychodium decipiens Limpr. 


E. Iceland: Berufjordur!Y, at an altitude of 80 metres. 

var. crassirete n. var. 

Leaf-cells very thick-walled, highly porous and distinctly papillose 
in the upper half of the leaf. Nerve very strong, 0.040—0.045 mm. 
broad for almost its entire length. 

NW. Iceland: Gnupsdalur, at an altitude of 280 metres!. 

L. decipiens is closely allied to L. saæxicola and is often difficult to 
distinguish from the latter species. Peculiar to both are the narrow, 
highly thickened and porous leaf-cells, which in L. decipiens are more 
or less distinctly papillose with a papilla at each of the uppermost 
cell-angles, at the back towards the leaf-apex; while L. saæicola almost 
always has quite smooth leaves. In L. saæicola the upper half of the 


THE BRYOPHYTA OF ICELAND salg 


leaf-margin is almost straight in outline, while in L. decipiens it is dis- 


tinctly concave, as in L. filamentosa. 


The variety grew abundantly on somewhat damp rocks; it is some- |. 
what coarser and less branched than the type which it resembles ex- 
actly in leaf-form and size of the cells. The leaf-cells in this form are 
so highly thickened and porous that the walls almost appear like a 


string of beads and are often broader than the cell-space. 











e RA d d d 


Fig. 13. a, Leaves of Lescuræa Breidleri (X 20): b, leaves of L. radicosa (X 20); 
c, leaves of L. decipiens (X 20); d, leaves of L. decipiens v. crassirete (X 20); e, leaf- 
cells of L. decipiens (X 320); f. leaf-cells of L. decipiens v. crassirete (X 320). 


225. Lescuræa radicosa (Mitt.) Hagen. 


Lescuræa rigescens (Wils.) Br. eur.; Pseudoleskea radicosa (Mitt.) Lindb. 
Ptychodium Pfundtneri Limpr. 


N. Iceland: Oxnadalur, Tverå!; sterile. 
In the above locality it grew scantily on a stone by the river. 


226. Lescuræa filamentosa (Dicks.) Lindb. 
Pseudoleskea atrovirens (Dicks.) Br. eur. 
Widely distributed. 


5) KØ A. HESSELBO 


Grows on stones and dry or somewhat damp rocks, and appears 
to be frequent, although not abundant, in all parts of the country up 
to about 300 metres above sea-level. In NW. Iceland it is very common 
on stony slopes, and is often richly fruiting while it has or less been 
found sterile only. 


227. Lescuræa Breidleri (Kindb.) Arn. et Jensen. 
Ptychodium oligocladum Limpr. 


E. Iceland: Berufjardarskard!, at an altitude of 600 metres; Seydis- 
fjordur!, 500 metres. N. Iceland: Reykjahlid!; Akureyri, at an altitude 
of about 770 metres!. Grimsey (O. D.)!; Oxnadalsheidi (Grl.)!. NW. Ice- 
land: Dvrafjérdur!; Gnupsdalur!; Isafjérdur!; Sugandafjordur!; Grunna- 
vik!. S. Iceland: Barkarstadr!, at an altitude of 530 metres. 

Frequent on rocks and stones in the more highly situated parts of 
the country. It occurs in greatest abundance and in its most typical 
form on mountain heights at elevations of above 500—600 metres, where 
it is often the most abundant constituent of the vegetation. It has its 
main distribution in NW. Iceland, where, for instance around Isafjéordur 
and Sugandafjérdur, it occurs very abundantly on the rocky flats from 
about 200 metres upwards. 9 plants are common, dg plants and fruit 
have not been observed. Å 

Lescuræa filamentosa and L. Breidleri, judging from the Iceland spe- 
cimens, are well-defined forms which, as a rule, may easily be distin- 
guished from each other. 

L. filamentosa is the lowland form, which has its main distribution 
in the birch region, and does not ascend much higher than about 300 
metres above sea-level. Here it forms low, dense, dark-green or blackish- 
green tufts on stones, and prefers somewhat damp localities. g plants 
are very frequent, ? plants occur more rarely. 

L. Breidleri is the Alpine form, which descends only by exception 
to the upper limit of the birch region. It forms deep, loose, yellowish 
tufts of great extension upon the dry stony slopes of the mountain 
heights, or upon large blocks of rock. Only sterile 9 plants have been 
found. Both the habit and the colour easily distinguish the two species 
from each other. L. Breidleri is almost branchless, but with some long, 
ascending branches often hook-shaped at the apex; L. filamentosa is 
densely and irregularly branched. In its most lowly situated habitats, 
for instance near Reykjahlid at an altitude of about 300 metres, and 
in some localities in NW. Iceland, L. Breidleri shows, however, a tendency 
to develop forms which approach in habit the vigorous forms of L. 
filamentosa var. brachyclados. The leaf-cells in all the investigated spe- 
cimens of L. filamentosa were short, oval or roundish and thick-walled, 
more or less but always distinctly papillose, frequently as far -down as 
to the leaf-base, the nerve was very rough at the back, which features 
most certainly distinguish doubtful forms from L. Breidleri. In the latter 
species the leaf-apex is longer, the nerve is narrower towards the apex 
and usually smooth, rarely indistinctly toothed at the back. The leaf- 
tissue in L. Breidleri is much more translucent than in L. filamentosa, 


THE BRYOPHYTA OF ICELAND Dig 


and is formed by prosenchymatous cells 0.008—0.009 mm. broad and 
3—4 times, at the apex about 6 times, as long, which are, as a rule, 
quite smooth, more rarely slightly papillose towards the leaf-apex. The 
leaf-margin in L. Breidleri is revolute for almost its entire length, so 
that only the apex itself is plane. In L. filamentosa the leaf-margin is 
sometimes plane, sometimes more or less revolute, and on the same 
plant leaves may be found with margins either quite plane or revolute 
to the apex, which effaces the difference between the type and the 
variety brachyclados (Schwågr.) Br. eur. This form, which occurs rather 
frequently in somewhat damp habitats, forms in several respects a 
transition to L. Breidleri, and has, as the latter, a well-developed central 
strand, while the usual forms of L. filamentosa never have a central 
strand. 


228. Lescuræa patens Lindb. 
Pseudoleskea patens (Lindb.) Limpr. 


E. Iceland: Berufjérdur!, %, 500—600 metres above sea-level. N. 
Iceland: Grimsey (O.D.)! (fr.). NW.Iceland: Gnupsdalur near Dyrafjérdur! 
fr.; Dynjandi! gg. W. Iceland: Reykjavik!; Kollafjérdur!$; Mådruvellir!, 
in several places up to a height of about 200 metres, gg. S. Iceland: 
Barkarstadr, about 460 metres above sea-level. 

Lescuræa palens appears to be rather widely distributed in all parts 
of Iceland. It is met with from the low land to high up in the Alpine 
region on somewhat damp, shady rocks or damp stony slopes, often 
very abundantly. Thus, in Gnupsdalur it occurred in masses on stony 
slopes from about 250—400 metres, in association with L. filamentosa, 
L. Breidleri, Pohlia commutata, Brachythecium reflexum and Lophozia spp.; 
and with numerous fruit which had just ripened on June 19th. Near 
Reykjavik it grew on the ground among large stones in a damp meadow, 
in company with Schistidium gracile. 

Note. Ptychodium plicatum (Schleich.) Schimp. is recorded by Grå n- 
lund from Grimstunga and Reykjavik. The plant from Grimstunga is 
only a form of Brachythecium albicans, and that from Reykjavik of 
Camptothecium lutescens. 


229. Heterocladium squarrosulum (Voit) Lindb. 
Heterocladium dimorphum (Brid.) Br. eur. 


E. Iceland: Berufjérdur!; Hof!. N. Iceland: Myvatn (Grl.)!; Ljosa- 
vatn!; Hof near Eyjafjordur (O. D.)!. SW. Iceland: Thingvallahraun!; 
Kollafjordur!. S. Iceland: Kihlhraun!. 

Occurred here and there on dry ground, especially on moss-covered 
slopes or in birch coppices. Near Hof in E. Iceland it grew on a stony 
slope (about 100 metres above sea-level) covered with Hylocomium spp. 
and RQRhacomitrium. In Thingvallahraun it was found associated with 
Hypnum uncinatum, Pohlia nutans and P. acuminata on a small slope 
in the birch coppice. Only sterile specimens have been found. 


DIA A. HESSELBO 


230. Thuidium tamariscinum (Hedw.) Br. eur. 


Vestmannaey: Heljusdalur and Stora Klit!. S. Iceland: Barkarstadr!; 
Holt!; Seljaland!. 

Gliemann records that this species was found by Morch, but 
there are no specimens of it in the collections. 

This species was found only in the southernmost part of Iceland, 
where it is frequent in several places on Vestmannaey and along the 
whole stretch of country south of Eyjafjall. Here it grows especially 
in the clefts at the foot of rock-walls or among large stones. Near 
Seljaland it was found abundantly on slopes stretching down towards 
the Seljalandså, associated with Thuidium delicatulum, Camptothecium 
lutescens, Scleropodium purum and Hylocomium spp. 


231. Thuidium delicatulum (Dill., L.) Mitten. 


NW. Iceland: Dyrafjordur (Hartz)!. W. Iceland: Esja!. S. Iceland: 
Reykirdalur!:; Skålholt!; Breidabolstadr!; Barkarstadr!; Seljaland!; Holt!; 
Drångshlid!. SE. Iceland: Hornafjérdur!. 

Common from Hornafjérdur in SE. Iceland throughout the whole of 
S. and SW. Iceland to Hvalfjéordur. There it grew on grass-covered 
slopes, especially on somewhat damp ground, up to a height of about 
200—300 metres; occasionally also on more damp meadow land. 
Near Barkarstadr it was common even at an altitude of about 320 metres. 

T. delicatulam becomes especially luxuriant on warm ground. By 
the hot springs near Skålholt it grew abundantly in cushions almost 
10 cm. deep, associated with Hypnum stramineum, H. Lindbergii and 
Hylocomium squarrosum. Only sterile specimens were found. 


232. Thuidium Philiberti Limpr. 


SE. Iceland: Hornafjérdur!. SW. Iceland: Ålafoss!. S. Iceland: 
Reykirdalur!; Holt!; Vestmannaey!. 

This species has a quite similar distribution to, and often grows 
in company with, 7. delicatulum from which it cannot be distinguished 
in the field. Near Alafoss it grew on boggy ground along the river. 
In Reykirdalur it was common on rather dry, moss-grown slopes; it 
was also common around Holt, south of Eyjafjall. 

Thuidium Philiberti is a weak species which in a sterile condition 
can be distinguished from T. delicatulum only by the length of the leaf- 
apex; the length varies considerably, however, in the Iceland plants, 
leaves being sometimes found in which the apex is elongated hair-shaped 
with 4—5 cells, and sometimes such leaves in which the apex consists 
only of a single cell or of two. Only sterile specimens have been found. 


233. Thuidium abietinum (Dill., L.) Br. eur. 


E. Iceland: Hof!; -Vallanes (H. J.;!).. N. Iceland: Akureyri. (Grl.;!); 
Hof near Eyjafjordur (O. D.)!: Mvvatn!; Vidimvri (Grl.)!; Tverå in Oxna- 
dalur!. W. Iceland: Esja!. S. Iceland: Barkarstadr!; Vestmannaey!. 


THE BRYOPHYTA OF ICELAND 515 


Occurs here and there on dry stony or moss-grown slopes up to 
about 200 metres above sea-level, especially in N. Iceland, where its 
rather common in many districts (around Eyjafjordur, in Oxnadalur, etc.). 


234. Thuidium lanatum (Stromgr.) Hagen. 
Thuidium Blandowii (W. et M.) Br. eur. 


Very common from Håskuldstadr in E. Iceland throughout N. and W. 
Iceland to Borgarfjordur. In NW. Iceland it occurs here and there, for 
instance near Isafjordur, Kaldalon and Dynjandi near Jokulsfjordur, but 
everywhere scantily only. In E. Iceland it has not been observed south 
of Berufjordur, and in W. Iceland it appears to be absent from the 
south of Borgarfjordur. In S. Iceland it has been found rather plenti- 
fully near Holt and Barkarstadr. 

T. lanatum grows almost always in bogs, where, especially in N. 
Iceland, it is often the most abundant constituent of the vegetation. It 
is more rarely met with on wet, soil-covered rocks. It is most frequent 
in the lowlands up to about 300 metres; but near Akureyri it is fre- 
quent even at a height of 400—500 metres above sea-level, and near 
Barkarstadr up to about 400 metres. Fruit, which occurs everywhere, 
ripens in the first half of July. 


Fam. HYPNACEÆ. 
235... Orthothecium rufescens (Dicks.) Br. eur. 
Stereodon rufescens (Dicks.) Mitt. 


W. Iceland: Melar (Grl.)!, sterile. 


236. Orthothecium intricatum (Hartm.) Br. eur. 
Stereodon subrufus (Wils.) Lindb. 


N. Iceland: Skagafjérdur (Grl.)!; Thrastarholsårgil near Eyjafjordur 
(O. D.)!. S. Iceland: Breidabolstadr!; Skågafoss!; Vestmannaey!. SE. 
Iceland: Hornafjérdur'!. 

On Vestmannaey it grew rather plentifully, sometimes on blocks of 
rock in Heljusdal, sometimes on damp slopes intermixed with other 
mosses. In the other stations it usually occurred on damp rocks, 
especially on tuff; thus on the sides of a cave near Skågafoss, and near 
Hornafjoérdur on the damp rock-faces near a waterfall. 


237. Orthothecium chryseum Schwågr. (Br. eur.) 
Stereodon chryseus Mitten. 


Very common in E. and N. iceland, rather scanty in NW. Iceland 
(Dyrafjordur and Sugandafjordur). In W. Iceland it was only found 


516 A. HESSELBO 


rather scantily in Botnsdalur and in a few localities in Esja, and in S. 
Iceland it has not yet been observed. 

It grows on a damp substratum, both on rocks ånd on soil up to 
about 400 metres above sea-level. Thus in N. and E. Iceland it is one 
of the most frequent species on wet gravelly ground. Only sterile 
specimens have been found. 


238. Cylindrothecium concinnum (de Not.) Schimp. 
Entodon orthocarpus Lindb. 


N. Iceland: Vididalr (Grl.)!. 'S. Iceland: Ytri Skogar (H. J.)!. 


239. Climacium dendroides (L., Dill.) W. et M. 


Very common over the whole of Iceland. 

Grows on damp substrata of every kind, especially in bogs, scat- 
tered among other mosses, but also on soil-covered rocks and damp 
gravelly ground; frequent also on more dry ground among grass. It 
hardly ascends higher than about 300 metres. Fruit was found only 
in E. Iceland below Fjardarheidi. 


240. Isothecium myurum (Pollich) Brid. 


SW. Iceland: Hafnarfjérdur!. S. Iceland: Holt!: Hrutafell!; Drångs- 
hlid!; Vestmannaey !. 

This species was only found in the south-western part of the 
country, where it was, however, rather frequent; and in the district 
south of Eyjafjall it was abundant in several localities, especially near 
Drångshlid, where it grew abundantly at the foot of perpendicular tuft- 
rocks facing south. Only sterile specimens have been found. 


241. Isothecium tenuinerve Kindb. 
Isothecium myurum var. piliferum C. Jens. 


E. Iceland: Papey (St.)!. NW. Iceland: Dynjandi!; Arngerdareyri!. 
W. Iceland: Stykkisholmur!; Budahbraun (H. J.)!; the hot springs near 
Reykjavik (Grl.;!); Videy!; Esja!; Hafnarfjérdur!; the lava-fields around 
Reykjavik!. S. Iceland: Holt!: Thingvellir!: Kihlhraun!; Krisuvik (Stp.)!. 

Common in the southern and south-western parts of the country 
in lava caves and clefts, probably also common in W. Iceland. In NW. 
Iceland rarer; and only found in one loåcality in SE. Iceland  Occurs 
in greatest abundance in lava clefts, for instance abundantly near Thing- 
vellir. Only sterile specimens have been found. 


242. Homalothecium sericeum (L.) Br. eur. 


Very common over the whole of Iceland. 
Grows on dry, more rarely on damp rocks, and often abundantly 


THE BRYOPHYTA OF ICELAND 517 


in Very extensive mats. It ascends to a height of about 400 metres 
above sea-level, and, for instance in Esja, is found abundantly even at 
an elevation of between 300—400 metres. on dry, steep rock-ledges. 
Fruit was found only near Breidabolstadr in S. Iceland, and in July it 
was empty of its spores. 


243... Camptothecium lutescens (Huds.) Br. eur. 
Hypnum lutescens Huds. 


Iceland (Wiinstedt)!. N. Iceland: Hnukr in Vatnsdalur (St.)!: com- 
mon near Geitaskard in Blondudalur!.  NW. Iceland: Dyrafjérdur!. 
W. Iceland: Breidabolstadr!; Lundur!; Esja!; Reykjavik (Grl.;!). S. Ice- 
land: Common!; Vestmannaey!. E. Iceland: Djupivogur!; Hornafjérdur!. 

Common in the southern and western parts of the country from 
Hornafjordur in East Iceland to Bléåndudalur in North Iceland. It has 
not been found in the remaining part of North Iceland or in the greater 
part of East Iceland from Berufjérdur northwards, and it is rare in 
North-west Iceland. It usually grows on dry, stony or grass-covered 
slopes or in coppices, occasionally also on rocks, but hardly extends 
higher than 200—300 metres. 

The leaves in this species have occasionally in their upper part 
cells which are shorter, and the angles of which are more or less 
distinctly shaped like a papilla. This is most distinctly seen in the 
forms from Reykjavik and Djupivogur, both of which grew on rocks. 


244... Camptothecium nitens (Schreb.) Schimp. 
Hypnum trichoides Neck., Lindb. 


Very common. 

One of the most frequent mosses on wet rocks. It occurs also 
frequently on soil-covered rocks or on more dry ground mixed with 
other mosses. In bogs it often forms the bulk of the vegetation and 
very frequently sets fruit, which ripens in the beginning of July. It 
occurs most frequently and abundantly in the low land, but ascends, 
however, to 400—500 metres above sea-level. 

The tomentose form is by far the most frequent. The smooth 
form, var. atrichum Kindb., appears to grow especially on rocks. It was 
found abundantly on soil-covered rocks close to the coast below Geit- 
hellir in E. Iceland. 


245... Brachythecium Mildeanum (Schimp.) Schimp. 


E. Iceland: Seydisfjordur (H. J.)!; Hamarfjordur!. SW. Iceland: Ala- 


foss!. å 
On wet boggy ground; near Hamarfjordur it grew abundantly on 
inundated ground along the river. 


518 A. HESSELBO 


246... Brachythecium salebrosum (Hoffm.) Br. eur. 
Hypnum plumosum Huds. 


Commonly distributed in all parts of the country. 

This species grows, as a rule, on rather dry ground, for instance 
on grass-covered slopes and on stones; occasionally also on damp ground, 
but everywhere scantily only and usually mixed with other mosses. It 
is most frequent in the low land; near Akureyri it has, however, been 
found at an elevation of about 500 metres, and near Seydisfjordur about 
400 metres above sea-level. Fruit has been found only near Reykir 
in S. Iceland. 


247... Brachythecium collinum (Schleich.) Br. eur. 


W. Iceland: Esja, Modruvellir!, on damp rocks at an altitude of 
about 100 metres. 

The form collected here differs in its somewhat more longly pointed 
stem-leaves and narrower, more slightly toothed, branch-leaves. 

Note. Brachythecium plumosum (Sw.) Br. eur. is recorded by Gron- 
lund from several localities, but all the specimens of it in the col- 
lections have been wrongly determined and the majority of them belong 
to Hypnum palustre. 











ig. 14. Brachythecium longipilum. Habit (X 3). 


248... Brachythecium longipilum n. sp. 


Monoicum. Flores masculini in cauli primario numerosa, antheri- 
diis c. 10 sordide flavis et paraphysibus paucis. Cæspites humiles, densi 


THE BRYOPHYTA OF ICELAND 519 


saturate virides, habitu fere formæ gracilis Braåchythecii velutini. Caulis 
repens, dense tomentosus, 2—3 cm. longus, 0.25 mm. crassus, ramos 
erectos numerosos, 4—5 mm. longos emittens. Folia caulina non de- 
currentia e basi late ovata vel triangulo-cordata in cuspidem longissimum 





N 
N 
NERE rg. 
Har 
a b bL b c c c 


Fig. 15. Brachythecium longipilum. <a, Capsule (X 20); b, branch-leaves (X 20); 
c, stem-leaves (X 20). 


et tenuissimum recurvatum producta, per totam marginem denticulata, 
1—1.2 mm. longa, 0.4—0.6 mm. lata, costa brevi et indistincta. Folia 
ramorum erecto-patentia aut subsecunda, lanceolata, longius acuminata, 
1.4—1.6 mm. longa, 0.4—0.6 mm. lata, per totam marginem denticulata, 











Fig. 16.  Brachythecium longipilum.  Leaf-base (X 175; phot.). 


costa mediam partem folii paulo excedente. Cellulæ angustæ, sinuatæ, 
0.004 mm. latæ, 0.060—0.075 mm. longæ, basilares in uno vel duo se- 
riebus, ovales, porosæ, pachydermiæ, 0.010 mm. latæ, in angulis quadratæ 
vel ovales, paucæ. i 

Perichætium e cauli primario egressum. Folia perichætialia exteriora 
apice brevi, interiora apice longo reflexo, ecostata vel costa perbrevi et 


520 A. HESSELBO 


indistincta, margine leniter dentato. Seta 8 mm. longa, c. 14 mm. crassa, 
purpurea, in dimidia parte superiori rugosa, inferne glabra. Capsula 
fusca, obliqua, longe ovata, operculata 1.7 mm. longa, 0.5 mm. crassa, 
deoperculata sub orificio contracta. Operculum conicum 0.6 mm. altum. 
Dentes peristomii 0.46 mm. longi, 0.067 mm. lati, pallide lutei, 
apicibus hyalinis, basi aurantiaci et leniter transverse striati. Sporæ 
0.021— 0.024 mm., papillulosæ. 

S. Iceland: Flokastadagil, on stones at the margin of the river. The 
capsules were empty in the middle of July. 


249. Brachythecium populeum (Hedw.) Br. eur. 


Hypnum populeum Hedw. 


S. Iceland: Vik (H. J.)!, fr.: Drångshlid!,. fr. 
Grew in both the above localities on rather dry tuff rocks. 


250... Brachythecium velutinum (L.), Br. eur. 


SFTcelan ds hine (GEN VIE HE) FRE Rn ak eavekakkan 
altitude of about 300 metres, fr. 

Near Thingvellir it grew in a lava-cleft, associated with Blepharo- 
stoma trichophyllum, Plagiochila asplenioides and Plagiothecium silvaticum. 


251... Brachythecium glaciale Br. eur. 
Hypnum glaciale C. Hartm. 


N. Iceland: Akureyri!, at an altitude of 770 metres. NW. Iceland: 
Sugandafjérdur!, at an altitude of 450 metres. W. Iceland: Dalasysla, 
Melår, on a cliff inhabited by sea-fowl (H. J.)!. 

On damp gravelly flats.. Near Sugandafjordur it grew abundantly, 
intermixed with Hypnum stramineum, Mniobryum albicans and Philonotis 
fontana, in cushions 3—4 cm. deep, on ground saturated by melting 
snow, and on stones. The plants were about 5 cm. long and irregularly 
branched with unequally long — mostly short — branches. The leaves 
were erectly spreading, not secund, and very longly decurrent with 
numerous quadrate, thin-walled cells at the basal angles of the leaves. 


252. Brachythecium reflexum (Starcke) Br. eur. 


Commonly distributed. i 

This species has its main distribution in gravelly spots on the 
rocky flat, especially on slopes and dry snowless patches on the moun- 
tain heights. It is especially abundant in NW. Iceland, where it is 
everywhere the most frequently occurring species on stony slopes. Here 
it is usually found associated with Hypnum uncinatum, Lescuræa spp., 
Lophozia lycopodioides, L. Flærckei, etc., and is met with as high as about 
500 metres above sea-level. In Esja, in SW. Iceland, it was common 
from about 200 metres upwards. In the low land it is rarer and appears 


THE BRYOPHYTA OF ICELAND BØJ 


to occur more particularly in the lava-fields, for instance near Hafnar- 
fjordur. Thingvallahraun and Kihlhraun. It is found rather frequently 
in fruit. 


253. Brachythecium glareosum (Bruch) Br. eur. 
Hypnum glareosum Bruch. 


E. Iceland: Seydisfjordur, on stones!. N. Iceland: Tverå in Oxna- 
dalur, on damp gravelly ground!.  W. Iceland: Botnsdalur, lamp 
rocks!. 


254. Brachythecium albicans (Neck.) Br. eur. 
Hypnum albicans Neck. 


Commonly distributed over the whole of Iceland. 

This species grows on a dry substratum, for instance on sandy or 
grass-covered ground, on soil-covered stones, on dikes and on the peat- 
walls of houses. It is most frequent in the low land up to a height 
of about 300 metres, but occasionally, for instance near Akureyri, it 
ascends as high as about 600 metres above sea-level. Only sterile speci- 
mens have been found. 


255. Brachythecium erythrorrhizon Br. eur. 
Hypnum erythrorrhizon Hartm. 


N. Iceland: Hof near Eyjafjordur (0. D.)!: Ljosavatn!; Tyverå in 
Oxnadalur!. W. Iceland: Esja!. 

Found in all the localities only scantily intermixed in the tufts of 
other mosses on rather dry ground. Near Ljosavatn it grew in the 
Betula nana heath in association with Hylocomium proliferum, Hypnum 
uncinatum, Heterocladium squarrosulum and Lophozia lycopodioides; in 
Esja it grew on a slope with a southern exposure, in company with 
Hylocomium spp. and Hypnum imponens. 


256. Brachythecium rivulare (Bruch) Br. eur. 
Hypnum rivulare Bruch. 


Very common over the whole of Iceland. 

One of the most frequent mosses, which occurs everywhere on 
damp ground up to about 400—500 metres above sea-level. It occurs 
both in large and small streams, where it covers the stones or the 
gravelly soil along the banks with its extensive mats, and in bogs, and 
is also one of the species which is most frequently met with in moss 
bogs. Occasionally, however, it grows also on a more dry substratum, 
for instance on soil-covered rocks and among grass. Only sterile speci- 
mens have been found. 


The Botany of Iceland. Vol. I, part Il. 34 


BD A. HESSELBO 


257. Brachythecium latifolium (Lindb.) Philib. 
Hypnum latifolium Lindb. 


Gjå near Alptagerdi (St.)!, in company with Mnium punctatum. 


258. Scleropodium purum (L.) Limpr. 
Hypnum purum L. 


SW. Iceland: Ålafoss!. S. Iceland: Breidabolstadr!; Barkardstadr!:; 
Holt!; Drångshlid!; Skågafoss!. 

This species has been found only in the southern part of Iceland, 
where it is frequent on Fljétshlid and below Eyjafjall. Near Ålafoss 
it-grew on the slope stretching down towards the river in which the 
temperature of the water was about 25”. Usually it grows at the foot 
of cliffs with a southern exposure, or in clefts, and, as a rule, in asso- 
ciation with Hylocomium spp., Eurhynchium piliferum, Mnium undulatum, 
Thuidium delicatulum and T. tamariscinum. 


259. Eurhynchium strigosum (Hoffm.) Br. eur. 
Hypnum strigosum Hoffm. 


Var. præcoæwæ (Hedw.) Lindb. 

N. Iceland: Hof (0. D.)!; Vidimvri (Grl.)!. W Iceland: Kalmanstunga 
Grl.)!; Gilsbakki!; Reykjavik!. S. Iceland: Barkarstadr!. 

Grows everywhere rather scantily on dry soil-covered rocks or 
on tuff. 


260. Eurhynchium diversifolium (Schleich.) Br. eur. 
Hypnum strigosum var. diversifolium Lindb. 


N. Iceland: Reykjahlid!; Hofsfjall near Eyjafjérdur (0. D.)!; Akureyri, 
at an altitude of about 900 metres!. 

Near Akureyri it grew in gravelly soil at the summit of the moun- 
fain, interspersed in tufts of Søhærocephalus turgidus, Dicranum congestum 
and Hylocomium proliferum. Near Reykjahlid it grew intermixed in a 
tuft of Timmia austriaca. 

Eurhynchium præcox and E. diversifolium are so closely allied that, 
according to my opinion, it is doubtful whether the last species can be 
regarded as anything more than a decidedly Alpine form of Eurhynchium 
strigosum. The leaves in the plants determined as E. diversifolium are 
longly decurrent. The cells of the basal angles of the leaves vary 
considerably in number, even on the same plant, usually they are very 
numerous; but specimens of E. strigosum var. præcox from Central Europe 
also vary considerably in this respect, and often have many more alar 
cells than has the type. 


THE BRYOPHYTA OF ICELAND 523 


261. Eurhynchium cirrosum (Schwågr.) Limpr. 
Brachythecium cirrosum (Schwågr.) Schimp. 


SW. Iceland: Esja!; Kolvidarhol!, at an altitude of about 300 metres. 

In Esja it grew rather scantily on a damp slope, in company with 
Hypnum chrysophyllum. Near Kolvidarhol it grew abundantly on the 
ground among grass and species of Carex. 


262. Eurhynchium piliferum (Schreb.) Br. eur. 
Hypnum piliferum Schreb. 


W. Iceland: Budahraun (H. J.)!: Ålafoss near Reykjavik!; Hafnar- 
fjordur!. SE. Iceland: Hornafjérdur!. S. Iceland: Reykirdalur!, up to 
about 150 metres above sea-level; Breidabolstadr!; Holt!, in several 
places up to about 300 metres above sea-level; between Thingvellir and 
Geysir (Grl.)!; common around Thingvellir!; Drångshlid!; Vestmannaey !. 

Rather common in the South-western and Southern parts of Iceland. 
It usually grows in protected localities, for instance in lava-clefts, but 
occasionally, for instance near Reykir, it occurs on moss-grown slopes 
in company with Hylocomium spp. Near Thingvellir it was found abun- 
dantly in Almannagjå, at the grass-covered bottom of the ravine. Only 
sterile specimens have been found. 


263. Eurhynchium Swartzii (Turn.) Curnow. 
Hypnum Swartzii Turn. 


S. Iceland: Drångshlid (H. J.;!); Merkjåfoss (F.)!; common on Fljéts- 
hlid and below Eyrjafjall!; Reykirfoss!. SW. Iceland: Alafoss. 

This species was found only in S. Iceland, but was common there 
on damp rocks, especially at the base of the sides of tuff rocks in clefts 
and in damp caves. Only stcrile specimens have been found. 

Note. Eurhynchium.- hians from Merkjåfoss (leg Feddersen) belongs 
to E. Sivarizil. 

Hypnum prælongum is recorded from Iceland by Zoéga. There 
are no specimens of it in the collections, but perhaps it has been con- 
fused with E. Swartzii 


264. Eurhynchium Stockesii (Turn.) Br. eur. 
Hypnum Stockesii Turn. 


S. Iceland: Drångshblid!; Hrutafell!; Holt!; Seljaland!; Vestmannaey 
(EJE: 

Rather common south of Eyjafjall and on Vestmannaey. It usually 
grows rather scantily on the ground, between blocks of rock or in clefts. 
On Vestmannaey it occurred in several places; among other localities 
at the foot of cliffs, in company with Mnium undulatum and Lophocolea 
cuspidata. Only sterile specimens have been found. 


347 


BD24 A. HESSELBO 


265. Rhynchostegium murale (Neck.) Br. eur. 
Hypnum murale Neck. 


S. Iceland: Drångshlid!; Hrutafell!. 

In the above localities it grew rather abundantly at the foot of 
dry tuff rocks and on fallen blocks. The fruit was ripe in the latter 
half of July, but some of the lids still persisted. 


266. Rhynchostegium rusciforme (Neck.) Br. eur. 
Hypnum rusciforme Neck. 


N. Iceland: Near Eyjafjordur (0. D.)!. NW. Iceland: Isafjérdur!. 
W. Iceland: Dalsmynne in Nordredalur!; Lundurreykjadalur!; Esja!; 
Ellidarå near Reykjavik!.  S. Iceland: Tungufoss. (E.)!: Vik (H-JJY: 
Reykirdalur!, fr.; frequent on Fljåtshlid and near Eyjafjall!. 

Var. atlanticum Brid. 

N. Iceland: Between Svinadalur and Dettifoss!; Husavik!. W. Iceland: 
Esja!tin several places FS SIceland: FReykird alu Ho AVE KRERIDN 

The type is rather frequent in the Southern and Western parts of 
Iceland, rarer in North Iceland, and has not been found in East Iceland. 
It usually grows on wet rocks near waterfalls, or on stones in the rivers, 
as a rule rather scantily, and mixed with Brachythecium rivulare, Hypnum 
ochraceum, etc. Only in Reykirdalur did it occur abundantly in the river 
and on the rock-sides by the waterfall, also in fruit. 

Var. atlanticum is found in swiftly flowing rivers, where it often 
covers the bottom and stones for long distances. 


267. Thamnium alopecurum (L.) Brid. 


W. Iceland: Dalasysla, Vogur (H. J.)!; Jærngerdarstadr (Sæmundson)!. 
S. Iceland: Reykirdalur!; Paradishellir (Stp.)!; common on Fljåthlid and 
below Eyjafjall!. i 

On wet shady rock-faces and in caves. In S. Iceland it is common 
in the tuff-clefts, especially on the inwardly sloping surfaces, and in the 
numerous dark, damp caves, where it often covers the roof and sides 
through which water is percolating. Only sterile specimens have been 
found. 


268. Plagiothecium silvaticum (Huds.) Br. eur. 


N. Iceland: Grimsey (0. D.)!. S. Iceland: Thingvellir (Grl.;!); AL 
mannagjå !. 

In Almannagjå it was found scantily at the bottom of a lava-cleft. 
Only sterile specimens were found. , 


269. Plagiothecium Roeseanum (Hampe) Br. eur. 


Syn. Plagiothecium silvaticum var. Roesei (Hampe) Lindb. 


Commonly. distributed, but as a rule rather scantily, in soil-filled 
rock-crevices, in company with Mnium orthorrhynchum,  Plagiochila 


25 


(ey | 


THE BRYOPHYTA OF ICELAND 


asplenioides, Pohlia cruda, etc., up to about 300 metres above sea-level. 
Near Kolvidarhol and Seljaland it was found at an altitude of about 
400 metres. Only sterile specimens have been found. 


270. Plagiothecium denticulatum (L.) Br. eur. 


N. Iceland: Myvatn (Grl.;); Grimsey (0. D.)!; Hof (0. D.)!; Vidvik 
(P. Såféniasson)'!; Asbyrgi!. NW. Iceland: Grunnavik!: Kaldalon!. W. Ice- 
land: Gilsbakki!; Hafnarfjérdur!; Reykjavik!. S. Iceland: Thingvellir!; 
Holt!: Vestmannaey !. 

Found here and there on humus-covered rocks or in lava-clefts, 
but everywhere scantily only, and sterile. 

Recorded by Kånig and Steenstrup from Iceland, but there are 
no specimens of it in the collections. 

Plagiothecium undulatum is recorded by Lindsay and P. silesiacum 
by Hornemann (from Geysir), but no specimens of either of these 
two species are to be found in the collections, so the determinations 
are undoubtedly erroneous. 


271. Plagiothecium pulchellum (Dicks.) Br. eur. 
Isopterygium nitidum v. pulchellum Lindb. 


Iceland (Mørch). Commonly distributed over the whole of Iceland. 

Grows in rock-crevices, caves, lava-clefts, etc.; either in small un- 
mixed tufts or, more often, intermixed with other mosses. It does not 
appear to ascend higher than about 400 metres above sea-level and, as 
a rule, sets fruit. 

The majority of the plants which have been investigated belong to 
the type. Var. nitidulum (Wahlb.) L. et J. (Isopterygium nitidulum Lindb.) 
is far rarer, and grows by preference especially in deep shade; but, for 
the rest, this form is not very characteristic and passes without limit 
into the type. 


272. Plagiothecium depressum (Bruch) Dixon. 


S. Iceland: Holt!; Vestmannaey, Heljusdalur!. 

In Heljusdalur it grew in masses on a talus of fallen blocks and 
debris (Urd), everywhere covering the ground between the large blocks 
with its brightly shining mats. Near Holt it was found mixed with 
Brachythecium velutinum at ihe bottom of a tuff cave at an altitude of 
about 300 metres. 


273. Plagiothecium elegans (Hook.) Sull. 
Isopterygium elegans (Hook.) Lindb. 


S. Iceland: Reykirdalur!, at an altitude of 260 metres. 

It was growing here on warm soil in the neighbourhood of a 
sulphur spring, on the ground below a projecting block of rock, in 
company with Diplopyllum albicans and Pellia Neesiana. 


526 A. HESSELBO 


The form found is robust, light green, highly shining and differs 
in habit by the leaves being very decidedly two-rowed, spreading. 
Gemmæ, which were, by the by, quite typical, occurred only extremely 
few in number. 


274. Amblystegium Sprucei (Bruch) Br. eur. 


Commonly distributed over the whole of Iceland. 

Grows in rock-clefts, dark ravines and caves, and among fallen 
blocks of rock, usually scantily woven into the tufts of other mosses 
such as Plagiothecium pulchellum, Amphidium Mougeottii, Lophozia Miilleri, 
L. heterocolpos and Mnium orthorrhynchum, but occasionally also in tiny 
cushions or as delicate, cobweb-like coverings on roofs in caves. In 
Esja it was found up to about 400 metres above sea-level. Only sterile 
specimens have been found. s 


27 


by | 


Amblystegium fluviatile (Sw.) Br. eur. 


SW. Iceland: Tungufoss (F.)!, delermined by C. Jensen. 
Only a few stems were found, intermixed with other mosses, for 
instance Brachythecium rivulare, Hypnum ochraceum, etc. 


276. Amblystegium serpens (L.) Br. eur. 


NETceland:Akureyri se fr (Gel) My vaner (Gr) AW Ce ande 
Reykjavik!. Vestmannaey!, fr. 

In Iceland this species is as variable as it is in other places. The 
specimens from Mvyvatn were very robust, with leaves 1.33 mm. long 
and 0.65 mm. broad, erectly spreading, and distinctly toothed in their 
lower half, with rather long and strong nerve, and cells 0.010—0.012 mm. 
broad and 4—6 times as long. Near Reykjavik it grew in large cushions, 
1—2"cm: deep, on a stone wall near "the sea-side, In the”plantfrom 
the latter locality the leaves were broadly ovate, longly pointed and 
almost entire at the margin, with cells 0.009—0.011 mm. broad and 
2—3 times as long. The form from Vestmannaey is very slender, with 
leaves 0.7 mm. long and 0.3 mm. broad, with very short nerve which 
in the branch leaves was just indicated immediately above the leaf-base. 
On Vestmannaey the fruit was quite unripe on July 6th; in N. Iceland 
it was ripe at the end of July. 


277. £Subsp. Amblystegium Juratzkanum Schimp. 


N. Iceland: Fagriskogur (SE) NE SET eelands FE Drångshlidt tr VEeSE 
mannaey !. 

The plant from Drångshlid was quite typical and bore ripe capsules 
at the end of July. The specimens which were collected on the base 
of a birch-trunk in Fagriskogur agreed fairly well with the form described 
by Limpricht (Kryptogamenflora, vol. III, p. 323) under the name of 
Amblystegium radicale. The leaves taper from an ovate-lanceolate base 


THE BRYOPHYTA OF ICELAND 527 


to a very long and slender point, and are about 1 mm. long and 0.4 mm. 
broad. The branch leaves are lanceolate and toothed along their entire 
margin, with short nerve. The cells at the base of the leaf are quadrate, 
yellow and rather thick-walled, 0.012—0.015 mm. broad; those in the 
upper half of the leaf narrowly prosenchymatous, 0.08 mm. broad, and 
6—8 times as long, and often somewhat wavy. 


278. Subsp. £ Amblystegium littorale (C. Jens.) Hesselbo. 
Syn. Amblystegium serpens var. littoralis C. Jens. in "Færåernes Mosser”. 


N. Iceland: Grimsey (0. D.)!. W. Iceland: Reykjavik!; Brjanslækur 
(HERE 

Both the plants agreed exactly with the specimens collected by 
C.Jensen in the Færåes. 4. littorale, which is a Sub-arectic-Arctic form, 
is probably widely distributed. It has been found not only in the 
Færåes and in Iceland but also on Jan Mayen (Dusén) and in Sweden, 
on Gisld near Troså (Arnell). It has as yet been found sterile only. 


279. Amblystegium compactum (C. M.) Br. eur. 


Vestmannaey (H. J.;!). SW. Iceland: Ondverdarnes, in a well toge- 
ther with Trentepohlia aurea (H. J.)!. 

On Vestmannaey this species occurred rather frequently on the sides 
of clefts and caves in tuff rocks. It grew there in company with 
Trentepohlia, especially in places where water was percolating through, 
and occurred sometimes in compact cushions several cm. deep, some- 
times as a thin covering. The Iceland specimeus agree exactly with 
North American plants of A. compactum; occasionally, however, the 
leaves especially of the forms growing as thin coverings are somewhat 
broader, with shorter cells. 

W. Moånkemeyer and L. Loeske (Revision einiger Amblystegien 
aus dem Herbare Limpricht; Ungarischen bot. Blåtter, 1911, p. 273) refer 
Amblystegium salinum Bryhn to A. compactum as var. salinum (Bryhn 
Mkm. On investigating numerous specimens of Å. salinum I have, how- 
ever, arrived at the conclusion that these two species are in no way 
connected with each other. Amblystegium compactum forms compact, 
dark-green tufts with a somewhat silky lustre, closely interwoven in the 
interior with rusty brown rhizomes which proceed partly from the stem 
and partly, in very great numbers, from the under side of the stem- 
leaves. The stem is prostrate with apex rising upward, and with 
numerous branches. The leaves are very close-set, longly decurrent, 
erectly spreading and secund. The stem leaves taper from a broadly 
ovate base to a long and slender point with strongly curved back apex. 
The branch leaves are narrowly lanceolate, often with apex curved. 
The leaf-margin is sharply toothed along its entire length, and especially 
the lowermost teeth are often turned outwards or towards the base. 
In the branch leaves the nerve reaches to the apex and is of almost 
equal width along its entire length. straight, or sometimes slightly sinuous. 
The leaf-cells are narrower than in any other Amblystegium, 0.005— 


528 ÅA. HESSELBO 


0.006 mm. broad, and 6—10 times as long, thin-walled, pointed and 
wavy. On the whole, both the cell-tissue of the leaf and the form of 
the leaf-teeth call to mind far more an Eurhynchium or a Rhynchostegium 
than an Amblystegium. 

Amblystegium salinum Bryhn grows in loose, yellowish green, hardly 
tomentose tufts.: Rhizoids occur only on the lowermost part of the 
stem and scantily on the. back of the nerve. The stems are erect or 
ascending, slightly and irregularly branched. The leaves are far more 
distant than in A. compactum, are spread out almost horizontally, very 
longly decurrent, ovate-lanceolate, with long, often somewhat obtuse apex; 
all are similar or the branch leaves are somewhat narrower. The leaf- 
margin is finely and obtusely toothed till towards the apex. The leaf- 
cells have the appearance typical of Amblystegium, and in the upper 
half of the leaf are 0.008 mm. broad and 4—6 times as long, with 
sinuose primordial utricle (the latter is usually wanting in A. compactum). 
The nerve is green and sinuose (as in Å. varium) and vanishes just 
below, or in, the leaf-apex itself. A. salinum is undoubtedly a true Eu- 
Amblystegium which is nearest allied to A. serpens, but also approaches 
certain forms of AÅ. varium (Hedw.) as regards the form of the basal 
cells of the leaf and the long sinuous nerve. It would also be remark- 
able if a species like A. compactum, which in North America grows on 
rocks and on damp ground in woods in the interior of the country, 
should in Europe turn into a decidedly salt-soil plant. Å. salinum has 
been collected in many places along the coasts of Sweden and Norway; 
first by Zetlterstedt on Oland and near Bosekop in Finmarken, and 
in the herbarium is named by him Amblystegium serpens var. littorale. 
Hagen has collected the same species in Salten in Nordland (65” 15' 
N. lat.), where it was growing in company with A. littorale (C. J.), and 
in a letter he named it Amblystegium breve n. sp. On the other hand, 
it is not known to me that the real A. compactum has been collected 
in Scandinavia. I have not seen the A. compactum from salty soil in 
North Germany mentioned by Loeske (loc. cit. pp. 272—273), nor have 
I seen Méonkemeyer's Å. compactum from Bornholm. 


280. Amblystegium salinum Bryhn. 


Vestmannaey! on damp sandy soil near the coast, sterile. 


281. Amblystegium trichopodium (Schultz) Br. eur. 


N. Iceland: Grimsey (0. D.)!. 

Judging from the plentiful intermixture of Green Algæ, the plants 
from the above locality must have been growing on wet, probably muddy 
ground. gg and 2 flowers were present in great numbers; fruit, on the 
other hand, had not been developed. The stem leaves were rather close- 
set, horizontally out-spreading, from an ovate base tapering to a long 
and slender point, finely toothed along the entire margin, 1.2—1.5 mm. 
long and 0.50—0.55 mm. broad, with green nerve, 0.04—0.045 mm. wide. 
The branch leaves were lanceolate, 0.95 mm. long and 0.30—0.35 mm. 
broad, with short green nerve. At the base of the leaf the cells were 


THE BRYOPHYTA OF ICELAND 529 


green and rectangular, 0.018 mm. broad and 0.070 mm. long, in its upper half, 
0.010 mm. broad and 4—7 times as long, with sinuous primordial utricle. 





i, st 














Fig. 17. Amblystegium trichopodium (Schultz). Håbit (X 3). 





Fig. 18. Amblystegium trichopodium (Schultz) Br. eur. a, Stem-leaves and b, branch-leaves 
(X 20); c, leaf-cells near margin (X120). 


530 A. HESSELBO 


282. Hypnum Sommerfeltii Myrin. 


N. Iceland: Stora Brekka near Eyjafjordur (O. D.)!. S.Iceland: Fell 
(Harder, 1908)!, a few stems among Hypnum uncinatum, H, stellatum 
and Dilrichum fleæxicaule. 


283. Hypnum chrysophyllum Brid. 
Amblystegium chrysophyllum (Brid.) Lindb. 


E. Iceland: Berufjérdur!. N. Iceland: Near Myvatn (Grl.)!; Hof near 
Eyjafjordur (OD)! —W. Iceland: Esja, several places (Grl::)' "SiTceland: 
Geysir, on warm ground!. Vestmannaey, Heljusdalur'!. 

Var. 8 zenellum Schimp. 

W. Iceland: Esja!. 

On damp gravelly ground and on rocks, rather rare and scanty. 
Near Geysir it grew plentifully on warm damp ground over which the 
tepid water from the basin was flowing. Only sterile specimens have 
been found. 

Hypnum Zemliæ C. Jensen, which was collected by Steenstrup 
(but no habitat is given) is only a form of Hypnum chrysophyllum. 


284. Hypnum protensum Brid. 
Amblystegium protensum Lindb. 


E: Iceland: —Skreiddalur! FSeydisfjordur FN Iceland: Hof (OD! 
frequent in Esja!. 

This species is probably widely distributed, but, on account of its 
great resemblance to the far more commonly occurring H. stellatum, 
cannot always be kept dislinct from the latter species. It grows in 
somewhat damp localities, both on stones and on the ground, but only 
sterile specimens have been found. 


285. Hypnum stellatum Schreb. 
Amblystegium stellaltum (Schreb.) Lindb. 


Very common everywhere on a damp substratum, on rocks, on the 
ground and in bogs, and not rarcly in fruit. It is abundant even at a 
height of 500—600 metres above sea-level. 


286. Hypnum polygamum (Br. eur.) Wils. 
Amblystegium polygamum Br. eur. 


Very common on wet, boggy ground. It occurs especially abundantly 
in the neighbourhood of the coasts, but is also very common in the 
valleys further inland, and hardly ascends higher than about 300 metres. 
Fruit, which is met with very frequently, ripens at the end of June or 
in the beginning of July. 


THE BRYOPHYTA OF ICELAND Sad 


287. Hypnum intermedium Lindb. 
Amblystegium intermedium Lindb. 


This species is no doubt common, at least in N. Iceland, where it 
occurs around Eyjafjårdur up to about 500—600 metres above sea-level. 
Only sterile specimens have been found. 


288. Hypnum revolvens Sw. 
Amblystegium revolvens De Not. 


Very common over the whole of Iceland. 

One of the most frequent and abundantly occurring mosses, which 
often forms the bulk of the vegetation in the bogs, both in the low 
land and in the Alpine region where it is- plentiful even at an altitude 
of about 600 metres, for instance near Eyjafjordur. Fruit, which is 
often present abundantly, ripens in the beginning of July. 

Hypnum revolvens is distinguished from H. intermedium only by the 
fact of its- being monoicous. Therefore, in cases of the frequently 
occurring quite sterile plants, it is really impossible to decide whether 
they belong to the one or to the other species. This is especially true 
of the vigorous, slightly branched aquatic forms, which had previously 
been referred to Hypnum Cossoni Schimp. Both H. revolvens and H. 
intermedium form such Cossoni-forms, but as they are mostly quite sterile 
it is impossible to determine them with any certainty. 


289. Hypnum uncinatum Hedw. 
Amblystegium aduncum (L.) Lindb. 


Very common everywhere upwards to the limit of plant-growth. 

Var. orthothecioides (Lindb.). 

N. Iceland: Grimsey (0. D.)!.. NW. Iceland: Arngardareyri!; Grun- 
navik!. E. Iceland: Kirkjubol (H. J.)!. 

Hypnum uncinatum is one of the most common mosses and occurs 
everywhere both on dry and on damp substrata, on earth, on rocks, 
in bogs, etc.; and often in such quantities that it is the most abundant 
constituent of the moss-carpet. It varics exceedingly in size, habit and 
colour.  Hypnum orthothecioides cannot be maintained as species; in 
addition to the forms found in the above-mentioned habitats, others 
forming all possible transitions to the type have been found in many 
other localities. Fruit, which is rather common, ripens in the first half 
of July. 


290. Hypnum Sendtneri Schimp. 
Amblystegium Sendtneri (Schimp.) De Not. 


N. Iceland: Helgavatnsfloi (Stp.)!. 


Ba A. HESSELBO 


291. Hypnum Kneiffii (Br. eur.) Schimp. 
Amblystegium Kneiffii Br. eur. 


This species. is common, especially in N. and E. Iceland, and is 
somewhat rarer in NW. and S. Iceland. It grows in very wet situations 
in bogs, in pools, along the banks of rivers or covering stones in the 
water. 

Aquatic forms with slightly secund, longly pointed leaves are fre- 
quent; short and broad leaved orthophyllous or drepanophyllous forms 
occur more rarely. The size of the leaf-cells is dependent on the form 
of the leaf; in f. brevifolia they are short in the middle of the leaf, 
about 3—4 times as long as they are broad. In Esja a brevi-orthophyllous 
form was collected in a moss-bog at an altitude of 410 metres. 

It has been collected in fruit only near Husavik in N. Iceland. 


292. Hypnum exannulatum (Gumb.) Br. eur. 


Amblystegium exannulatum De Not. 


Very common. 

Hypnum exannulatum occurs in numerous forms everywhere on a 
damp substratum, often forming the bulk of the vegetation, either alone 
or mixed with other Hypnaceæ (H. stramineum, H. revolvens, etc.);, with 
Cinclidium, Mnium spp., and other bog mosses. It grows not only in 
bogs, but also in pools, moss-bogs, along streams, and occasionally on 
damp rocks. Near Akureyri it was abundant even at an altitude of 
600 metres. 

Var. purpurascens (Schimp.) (Hypnum purpurascens Limpr.) is the 
most frequent form, at any rate in N. and E. Iceland. It is usually 
purplish, more rarely green, erect, as much as about 20 cm. high, and 
often forms large, pure growths on very wet ground, especially in the 
water itself, for instance along the banks of small streams. In this form 
the leaf-base is never decurrent. In the plants which have been in- 
vestigated the cells of the leaf-base are sometimes quite typical, forming 
a single row of large dilated cells across the entire leaf-base, and some- 
times they approach those of the type by forming a more or less large, 
triangular group of empty cells towards the leaf- margin. In typical 
Hypnum exannulatum the leaves are always narrowly decurrent. 

Var. Rotæ (De Not.) is without doubt an extreme aquatic form of 
var. purpurascens, with which it is closely connected by intermediate 
forms. It has the solid, tough stem with long, forwardly directed 
branches and very narrow, slightly secund leaves peculiar to all mosses 
growing in swiftly flowing water. The nerve is very strong, usually 
0.10 mm. wide at the base and, as a rule, vanishing in the apex, more 
rarely excurrent. It grows in running water. often in streams flowing 
through boggy ground. Typical specimens were collected in the following 
localities: E. Iceland: Seydisfjordur!, rather common; Breiddalskard'!; 
Frodarheidi (H. J.)!. NW. Iceland: Kaldalon!; Dynjandi!. OW. Iceland: 
Hvammur (Grl.)!; Esja!, at an altitude of 100 metres. Only sterile 
specimens have been found. 


THE BRYOPHYTA OF ICELAND 533 


Var. serratus (Warnst.). 

Isafjordur!, in pcols. 

In H. exannulatum the length of the leaf-cells varies according to 
the shape of the leaf. In the most commonly occurring forms with 
longly pointed leaves the cells are narrow, 6—10 times as long as broad, 
while short-leaved forms have leaf-cells which are only 4—6 times as 
long as broad (var. brachydictyon (Ren.), such forms are especially met 
with on cold, wet gravelly ground. Fruit is rather frequent, both in 
the type and in var. purpurascens, and ripens at the end of July. 


293. Hypnum fluitans (Dill.) L. 
Amblystegium fluitans (L.) De Not. 


S. Iceland:  Breidabolstadr (F.)!; Breidamerkursandur (Harder)!. 
N. Iceland: Blejkjuholt (St.)!; Myvatn (Grl.)!. 

Monoicous forms, which may be referred to H. fluitans occur only 
very scantily and rarely in Iceland. The numerous older records of 
the occurrence of this species (Vahl, Méårch, Hornemann, Gronlund, 
etc.) are, without exception, due to erroneous determinations, and all 
the older specimens in the collections (except one) belong to H. exannu- 
latum or H. Kneiffii. 

H. fluitans grows everywhere in pools or on inundated ground. 
Near Blejkjuholt Stefånsson has collected a very peculiar form, with 
erect, obtuse leaves - with short nerve. The cell-tissue of the leaves is 
very loose, and the cells of the leaf-base differ only slightly from the 
other cells. 


Var. falcatum Schimp. (Hypnum H. Schultzii Limpr.) 
This very characteristic form was found abundantly, and in fruit, 
on wet boggy ground near Kaldalon in NW. Iceland. 


294. Hypnum filicinum L. 
Amblystegium filicinum (L.) De Not. 


Very common. 

In the greater part of Iceland this species is one of the most fre- 
quent mosses on a damp substratum, especially on wet rocks, but also 
on gravelly ground and in bogs. In most part of N. Iceland it is some- 
what rarer, and usually occurs more scantily. In S. Iceland it often 
covers the faces of wet tuffrocks in enormous mats and not rarely sets 
a great quantity of fruit. In the rest of Iceland fruit is rarer and occurs 
most often in the large, tomentose bog-form. In S. Iceland fruit was 
ripe in the middle of July. 

Hypnum filicinum varies exceedingly. Bog forms and forms from 
wet gravelly ground are usually densely tomentose and regularly pinnately 
branched. Rock forms, when growing on a more dry substratum, are 
slender and slightly branched; on wet rocks, especially on tuff. very 
large and vigorous, often almost like Hypnum commutatum. 


594 Å. HESSELBO 


H. filicinum has its main distribution in the lowlands up to about 
300 metres above sea-level, but is also frequently met with, for instance 
near Berufjordur and in Esja, up to a height af about 500 metres. 
At this altitude it grows on rocks in low, dense cushions, more rarely 
on wet gravelly ground, and these Alpine forms constitute a distinct 
transition to H. curvicaule Jur., which is perhaps only a High Alpine 
form of H. filicinum. 


295. Hypnum curvicaule Jur. 
Amblystegium curvicaule (Jur.) Dicks. & James. 


E. Iceland: Berufjardarskard!, 540 metres above sea-level; Seydis- 
fjordur!, frequent from about 100—500 metres above sea-level. W. Iceland: 
Moådruvellir!, about 400 metres above sea-level. 

Grows on damp gravelly ground and rocks on mountain heights 
up to 500—600 metres. On dry rocks the leaf-cells become longer and 
narrower and the alar cells more thick-walled. , Near Seydisfjordur it 
was collected with quite young fruit (3. 7. 1909). 


296. Hypnum decipiens (De Not.) Limpr. 
Amblystegium glaucum f. decipiens Lindb. 


E. Iceland: Very common near Seydisfjordur!; "Dvergasteinn (H. J.)! 
N. Iceland: Husavik!; Ljosavatn!; Akureyri!; Oxnadalur!. OW. Iceland 
Brattabrekka!. 

Common from Seydisfjordur in E. Iceland throughout the whole of 
N. Iceland. The westernmost habitat of this species is Brattabrekka, 
south of Haukadalur. In thes-rest of Iceland it has not been observed. 

Hypnum decipiens grows on very wet, especially gravelly, ground, 
for instance at the edge of moss-bogs or on inundated gravelly ground, 
and along small streams both on the ground and on stones: Ilt hardly 
ascends higher than about 300 metres, and rarely sets fruit. Near 
Seydisfjårdur, Husavik and Tverå in Oxnadalur it was collected in fruit. 


297. Hypnum commutatum Hedw. 
Amblystegium glaucum (Lam.) Lindb. 


E. Iceland: Seydisfjordur (H. J.;!). N. Iceland: Husavik!; Reykjahlid!. 
W. Iceland: Lundur!; rather frequent in Esja!. S$, Iceland: Rather common !. 

Var. falcatum (Brid.) C. M. Hypnum falcatum Brid. 

Very common. 

H. commutatum is the more rarely occurring form. It is most 
frequent in SW. and S. Iceland, where it is especially abundant on wet 
tuff-rocks. It is rather rare and scanty everywhere else in Iceland. 

Var. falcatum, together with all possible transitions to the type, is 
one of the most common mosses in the greater part of Iceland. Only 
in NW. Iceland is it scanty and met with only here and there. Both 


THE BRYOPHYTA OF ICELAND 535 


forms grow on rocks in and by the water, on wet rock-faces and on 
the ground, both on wet gravelly ground and on the ground in bogs. 
It often forms, especially in N. Iceland, an essential part of the moss- 
carpet in bogs. It hardly ascends higher than about 300 metres above 
sea-level. 2 plants are very common, & plants and fruit are far rarer. 
It has been found in fruit near Stafafell, Seydisfjordur, Hrutafjérdur 
and, rather frequently, in S. Iceland (both forms). 


298. Hypnum molluscum Hedw. 
Ctenidium molluscum (Hedw.) Schimp. 


Iceland (Morch)!; Bardshellir (Grl.)!; Brynjudalur (Grl.)!. "W. Iceland: 
Common near the hot springs in Reykholtdalur!:; Botnsdalur!; common 
about Reykjavik!; common in Esja!; Kolvidarhol!, up to about 350 
metres above sea-level. S. Iceland: Very common!. SE. Iceland: Horna- 
fjordur!. 

This species is very common in S. Iceland, especially on Fljétshlid 
and about Eyjafjall, but is also plentiful near Thingvellir, and frequent 
in SW. Iceland as far as Borgarfjordur, where it is, however, chiefly 
confined to the warm ground. It appears to be quite absent from the 
other parts of Iceland. 

H. molluscum grows on a damp substratum, both on damp rocks, 
especially tuff, and on damp ground. Near Thingvellir it grew abundantly, 
covering the blocks at the bottom of the large lava-clefts. Around the 
hot springs in SW. and W. Iceland it occurs abundantly in several places. 
Thus in Reykholtdalur it is one of the most frequent species of the 
warm clayey flats. 

It occurs almost exclusively in the low land, and only near Kolvi- 
darhol was it found about 350 metres above sea-level. Only sterile 
specimens have been found. 


299. Hypnum imponens Hedw. 
Stereodon imponens (Hedw.) Brid. 


E. Iceland: Hof, on basalt rocks!. W. Iceland: Esja!; Braudarholt!. 
N. Iceland: Vidimyri, near a hot spring (Grl.)!. W. and SW. Iceland: 
Near almost all the hot springs'!. 

In SW. and W. Iceland it grew with a few exceptions on warm 
ground. In Esja it grew in company with Camptothecium lutescens on 
a grass-covered slope, and near Braudarholt intermixed with other 
Hynaceæ on the top of knolls in bogs. 

On a warm substratum Hypnum imponens grows especially on the 
drier clay-flats; there it often forms extensive mats, partly alone, partly 
mixed with Hypnum stramineum, H. Lindbergii, H. molluscum, Hylocomium 
squarrosum and other species. Only sterile specimens have been found. 


536 A. HESSELBO 


300. Hypnum Bambergeri Schimp. 
Stereodon Bambergeri (Schimp.) Lindb. 


W. Iceland: Breidabolstadr in Reykholtdalur'!. 
It was growing there on a damp gravelly flat in company with 
Ditrichum flexicaule, Myurella julacea, Tortella tortuosa and T. fragilis. 


301. Hypnum revolutum (Mitt.) Lindb. 
Stereodon revolutus Mitten. 


Very common over the whole of Iceland. 

It grows both on dry and on more or less damp rocks, occasionally 
also on gravelly soil. and is frequent up to about 600—700 metres above 
sea-level. It varies considerably in size, from quite slender forms (for 
instance on lava blocks) to very robust forms on damp stones in clefts. 
Fruit, which was ripe in the middle of July (but the lids still persisted), 
was found only near Brcidabolstadr in S. Iceland, where Hypnum revo- 
lutum grew in deep cushions, covering stones in a cleft. 


302. Hypnum cupressiforme L. 


E. Iceland: Berufjérdur (Grl.;!); Hof!; Hornafjérdur!.  N. Iceland: 
Vidvik near Skagafjérdur (P. Séféniasson)!; Vidimyri!; common west of 
Bléndudalur!. W. Iceland: Stykkisholmur!; Botnsdalur!; common about 
Esja, Hvalfjordur, Rcykjavik and Hafnarfjordur!. S. Iceland: Common!. 
Vestmannaey !. 

Common from Berufjordur in E. Iceland throughout S. and W. Iceland 
and the western. part of N. Iceland to Bléåndudalur. In E. Iceland it 
has not been found further northwards than Berufjérdur, and in NW. 
Iceland it has not yet been observed. East of Hunafloi it has been 
found only in a few localities near Skagafjordur. 

It usually grows on rather dry rocks or on soil-covered rock-ledges, 
and does not appear to ascend higher than about 300 metres above 
sea-level. Only sterile specimens have been found. 

Var. ericetorum is recorded from Ormarstadir in E. Iceland (leg. H. J., 
det. C. Jensen), but there are no specimens of it in the collections. 


303... Hypnum hamulosum Br. eur. 
Stereodon hamulosus Lindb. 


E. Iceland: Hof!, at an altitude of 100 metres; Hornafjérdur!. 
W. Iceland: Esja, Méådruvellir!, on tuff rocks, 250—400 metres above 
sea-level; Kolvidarhol!; common on tuff, from 250 to 450 metres. 
NW. Iceland: Grunnavik!, at an altitude of about 280 metres. S. Iceland: 
Holt!: Drångshlid!. Vestmannaey!. 

Rather frequent in the South-western and South-eastern parts of 
Iceland. It has been found almost everywhere on tuff-rocks, where it 


THE BRYOPHYTA OF ICELAND 537 


occurred abundantly in several places, for instance in Esja, but only 
sterile specimens were found. 


304... Hypnum callichroum (Brid.) Br. eur. 
Stereodon callichrous Brid. 


E. Iceland: Seydisfjordur!. NW. Iceland: Common everywhere!. 
W. Iceland: Budahraun (H. J.)!; Grundarfjérdur (H. J.)!; common every- 
where in the lava-fields east of Reykjavik and around Hafnarfjordur!; 
Svinahraun!; Esja, in Grimmia-heath at an altitude of about 400 metres, 
and in many places along the banks of streams!. S. Iceland: Skålholt!; 
Thingvellir!; Holt!, common on tuffslopes in clefts. 

This species is common in the south-western part of Iceland, and 
very common in North-west Iceland. In North Iceland it has not been 
found, and in East Iceland only near Seydisfjordur. It occurs in Ice- 
land in two forms. The most frequent form is the typical bog-soil 
form, which is exceedingly common especially in NW. Iceland, and 
sometimes forms the bulk of the vegetation in damp situations, especially 
along streams, on wet slopes and at the edge of bogs, usually in 
association with Harpanthus, Lophozia quinquedentata, Cephalozia bicus- 
pidata, etc. Around Isafjéordur it occurs up to a height of about 
300 metres, and in S. and SW. Iceland up to almost 500 metres above 
sea-level. 

Everywhere in the depressions and at the bottom of the clefts of 
the lava-fields of South-western and West Iceland a mesophilous form 
is found, which is distinguished by its size and by its almost regularly 
two-rowed ramification. It grows in deep, loose and soft cushions, 
either among grass or mixed with Hylocomium spp., Rhacomitrium hyp- 
noides and species of Dicranum and of Lophozia. 

Fruit was collected only in Kaldalon in NW. Iceland: it was quite 
undeveloped even at the end of June. 


305... Hypnum Lindbergii Mitten. 


Stereodon arcuatus Lindb. 


Very common. 

This species grows on wet ground, especially in bogs, associated 
with H. revolvens, H. cuspidatum, Hylocomium squarrosum, Meesea, etc. ; 
but also on damp gravelly ground and on soil-covered rocks. It appears 
lo be of almost equal frequency everywhere. Near Berufjérdur it was 
collected up to a height of about 500 metres above sea-level, and on 
Eyjafjall in S. Iceland it was frequent at the same height. Near Akureyri 
it occurred abundantly on boggy ground even at an altitude of 500- 
600 metres. Only sterile specimens have been found. 

Hypnum pratense is recorded by Gronlund from several localities, 
but the record is due to a confusion with H. Lindbergili. 

Note... Hypnum incurvalum is recorded from Iceland by Moårch, 
but the specimens in his collection belong to Plagiothecium pulchellum. 


The Botany of Iceland. Vol. 1, part. II. 35 


538 A. HESSELBO 


306... Hypnum palustre Huds. 


Amblystegium palustre (Huds.) Lindb. 


E. Iceland: Seydisfjordur!. UN. Iceland: Akureyri (Grl.)!; Vidimyri 
(Grl.)!; Grimstunga (Grl.)!; Ljosavatn!, at an altitude of about 300 metres; 
Melar (Grl.)!.. W. Iceland: Gilsbakki!; Reykholtdalur (Grl.;!); Lundur- 
reykjadalur!; Reykjavik!; common in Esja!; Alafoss!. S. Iceland: Common!. 

This species is common in S. Iceland (Eyjafjall, Fljétshlid, etc.), 
SW. and W. Iceland, is found here and there in N. Iceland and is rare 
in E. Iceland. It has not yet been found in NW. Iceland. It grows 
everywhere on rocks and stones in and by rivers, and almost always 
sets fruit, which ripens during July. It does not appear to ascend higher 
than about 300 metres above sea-level. 

Of the varieties only var. d subsphæricarpon (Schleich.) Br. eur. is 
of any interest. It occasionally occurs in company with the type on 
stones in swiftly flowing rivers. 


307... Hypnum arcticum Sommerf. 
Amblystegium Smithii (Sw.) Lindb. 


W. Iceland: Hvammur (Grl.)!.. NW. Iceland: Bæir! fr. It is, more- 
over, recorded from Iceland by Méårch and from Akureyri (Grl.), but 
both the plants are wrongly determined. 

Near Bæir it grew on wet rocks by a small waterfall. 


308... Hypnum alpestre Sw. 
Amblystegium rivulare (Sw.) Lindb. 


E. Iceland: Berufjérdur!, common; Seydisfjordur!, common up. to 
about 200—300 metres above sea-level; Breiddalskard!, at an altitude of 
about 400 metres; Vallanes (H. J.;!), common on rocks along the sea; 
Skriddalur!. N. Iceland: Oxnadalur!, very common in the river. NW. Ice- 
land: Isafjéordur!, frequent up to about 200 metres; Sugandafjordur!: 
Arngerdareyri!. W. Iceland: Lundur!. SW. Iceland: The Laxå (F.)!; the 
Langå (H. J.)!; the Oxararå !. 

Common in E., N. and NW. Iceland, rarer in W. and SW. Iceland. 
This species has its main distribution in E. Iceland, where it occurs 
everywhere in the rivers and often covers the stones along the banks 
for long distances. Fruit, which occurs rather commonly, ripens in 
July (Oxnadalur, July 24; Vallanes, hardly ripe on June 28th). 


309... Hypnum alpinum Schimp. 
Amblystegium molle f alpinum Lindb. 


E. Iceland: Berufjérdur!" N. Iceland: Akureyri (Grl.;!); Hof near 
Eyjafjordur (0. D.)!: Holar (Grl.)! fr; "Lækjarmot! fr; common in Oxna- 
dalur! fr., NW. Iceland: Isafjéårdur! fr.; Dyrafjérdur! fr.; Arngardareyri ! 
fr.; Dynjandi! fr. 

Figured in Flora Dan. (tab. 2621, fig. 2) as Hypnum molle, leg Morch. 


THE BRYOPHYTA OF ICELAND 539 


This specimen is not to be found in the collections; on the other hand, 
a specimen from Iceland, determined by Mårch as Hypnum arcticum, 
belongs to H. alpinum. 

Rather common in N. and NW. Iceland, where it grows on stones 
in rivers, like the other species of Hygrohypnum. It does not ascend 
higher than about 200 metres above sea-level (Isafjérdur), and often 
sets fruit which ripens in N. Iceland in the latter half of July. 

Hypnum alpinum varies considerably in leaf-form, leaf-tissue and 
habit. 

The leaves, which are most often spreading, are occasionally fal- 
cato-secund along the entire length of the stem, in which feature it 
approaches H. dilafatum in habit. The Iceland specimens of H. alpinum 
approach, on the whole, H. dilatatum, both in habit, leaf form and struc- 
ture of cell-tissue. The cells of the base of the leaves are often thickened 
and slightly porous, and the alar cells are sometimes colourless and 
thin-walled, sometimes yellowish with slightly thickened walls. The 
leaf-form varies from almost orbicular to elliptic. The leaf-margin is 
almost always finely toothed all round. 

I have, however, thought it more correct to refer all the Iceland 
plants to H. alpinum, as at any rate the capsule and the perichætium 
corresponded exactly to those in this species. Thus, fully developed 
cilia have never been found in the peristome. The leaf-cells are also 
shorter than in H. dilatatum and in size agree most nearly with those 
in H. alpinum. 


310. Hypnum ochraceum Turn. 
Amblystegium ochraceum Lindb. 


Very common on stones in rivers up to 300—400 metres above 
sea-level, but only sterile specimens have been found. 

Varies considerably in size, habit, etc. 

Var. uncinatum Milde often occurs in company with the type. 

Var. Zfiliforme Limpr. is rather common in swiftly flowing rivers, 
where it often covers the bottom for long distances. 

Note. Amblystegium polare Lindb. is recorded by Gronlund (Tillæg 
til Islands Kryptogamflora, Bot. Tids., 20, p. 113) to have been collected 
near Vallanes by H.Jønsson. The specimen in question is in the 
Botanical Museum in Copenhagen and is labelled "Dichodontium pellu- 
cidum, Polytrichum alpinum, Amblystegium polare, det. C. Jensen”, but in 
spite of a careful search it was not possible to find Å. polare among 
the specimens, so for the present the occurrence of this species in Ice- 
land must be regarded as uncertain. 


311. Hypnum cordifolium Hedw. 
Amblystegium cordifolium De Not. 


N. Iceland: Myvatn. (Grl.)!.. NW. Iceland: Grunnavik!, fr. 
This species is also recorded from the Olfuså, Nesland, Litlu Bor- 
garkatlar and Thorodstadaengjar, but the plants from all these localities 


355 


540 A. HESSELBO 


are only forms of H. giganteum, which often occurs in slender, slightly 
branched forms, which are very difficult to distinguish from H. cordi- 
folium. As a rule, however, the old typical shoots from the previous 
year may be found at the bottom of the tuft, from whence the cordi- 
folinm-like shoots proceed. Near Grunnavik H. cordifolium grew abun- 
dantly on boggy, partly inundated ground, and there it was richly 
fruiting. On the damp ground it was flaccid and prostrate, in water 
erect in large, deep cushions. 


312... Hypnum Richardsonii (Mitt.) Lesq. et James. 
Amblystegium Richardsonii Lindb. 


E. Iceland: Skorastadr (H. J.)! fr.; Seydisfjordur!, abundantly at an 
altitude of 350 metres. N. Iceland: Thorodstadaengjar (St.)!; Ljosavatn! 
fr. at an altitude of about 250 metres; Husavik!. NW. Iceland: År- 
muli!. W. Iceland: Méødruvellir!, at an altitude of about 200 metres; 
Ijallahåls (H.J.)!. S. Iceland: Barkastadr!, at an altitude of 530 metres. 

This species is probably more frequent than may be believed, 
judging from the few habitats, but it is often overlooked on account of 
its resemblance to the exceedingly common H. cuspidatum and H. gigan- 
teum. It grows in bogs, often abundantly, associated with H. strami- 
neum, H. sarmentosum, H. revolvens, H. exannulatum, etc., and often sets 
fruit which ripens in the middle of July. 


313... Hypnum giganteum Schimp. 
Amblystegium giganteum De Not. 


Very common. 

Grows in bogs, where it is often the most abundant constituent of the 
vegetation, especially in the wettest situations, in channels and pools. 
Near Akureyri it was common in bogs even at an elevation of 600— 
700 metres. Fruit, which occurs rather frequently, ripens in the first 
halffor July: 


314... Hypnum stramineum Dicks. 
Amblystegium stramineum De Not. 


Very common over the whole of Iceland. 

It grows on damp ground, usually intermixed in tufts of other 
Hypnum spp., Sphagna, Cinclidium, Mnium, etc. On very wet ground 
it often forms pure extensive growths. Thus, near Ljosavatn, it grew 
in the wettest parts of a bog at an altitude of 400 metres, in extensive, 
richly fruiting mats, 10—20 cm. deep. In depressions in the rocky 
flat it forms in association with H. sarmentosum, H. revolvens and 
H. exannulatum low, dense almost black growths. 

Fruit was ripe near Ljosavatn on July 18th. 


THE BRYOPHYTA OF ICELAND 541 


315... Hypnum sarmentosum Wahlb. 
Amblystegium sarmentosum De Not. 


Commonly distributed on boggy ground, but as a rule not plenti- 
fully, mixed with other Hypnaceæ or in unmixed tufts. 

It varies considerably in size and colour. The slender High Alpine 
forms, which occasionally form the bulk of the vegetation at an allitude 
of 500—700 metres, sometimes approach so close to H. stramineum 
that it may be difficult to keep these two species distinct from each 
other. 

In S. Iceland it has been found in many places from about 300 me- 
tres upwards. but it has not been found in the low land. In the other 
paris of Iceland it is distributed from the low land up to about 
700 metres above sea-level, but appears to be most frequent above a 
height of about 200—300 metres. It has been collected in fruit near 
Hof in E. Iceland, Ljosavatn in N. Iceland where it occurred abundantly 
in bogs at an altitude of about 250—300 metres, and near Grunnavik 
in NW. Iceland where it was the most abundant constituent of the 
vegetation in a bog. The fruit, in the specimens from all these localities, 
was green in the middle of July. 


316... Hypnum trifarium W. et M. 
Amblystegium trifarium De Not. 


E. Iceland: Seydisfjordur!; N. Iceland: Vatnsskard !. 

In both the above localities it was growing rather scantily on very 
wet boggy ground, in company with Hypnum scorpioides, H. stramineum 
and H. revolvens. 


317... Hypnum turgescens Th. Jensen. 
Amblystegium turgescens Lindb. 


E. Iceland: Djupivogur!. N. Iceland: Svinadalur!. S. Iceland: Bar- 
karstadr!, at an altitude of 530 metres. 

It grew everywhere abundantly on boggy ground, in company with 
Hypnum revolvens, H. scorpioides and H. polygamum. 


318. Acrocladium cuspidatum (L.) Lindb. 


Hypnum cuspidatum L. 

Very common. 

This species grows on more or less damp ground, especially in 
bogs and damp meadows, but also in the wøter itself in moss bogs 
and pools, or on damp rocks. Ås a rule it does not ascend higher 
than 250—300 metres above sea-level. In S. Iceland it has been found. 
however, in several places up to a height of about 500 metres. 

Fruit, which occurs rather frequently, was ripe about July ist. 


549 A. HESSELBO 


319. Scorpidium scorpioides (L.) Limpr. 
Amblystegium scorpioides Lindb. 


Very common. 

It usually grows in the wettest parts of the bogs, often in the 
water itself, in association with Hypnum giganteum and H. exannulatum. 
In the greater part of the country it is met with everywhere in such 
localities; only in SW. Iceland (Fljotshlid and Eyjafjall) does it appear 
to be less frequent. It appears only by exception to ascend higher than 
about 300 metres, near Seydisfjordur it was collecied at an altitude of 
400 metres. Only sterile specimens have been found. 


320... Hylocomium proliferum (L.) Lindb. 
Hylocomium splendens (Hedw.) Br. eur. 


Very common. 

This species is found especially on a dry substratum, for instance 
on slopes, amongst grass, in heath-soil or as the most abundant consti- 
tuent of the moss-covering on stones and in coppices. It occurs how- 
ever also on damper ground, intermixed in tufts of Sphagnum, Sphæ- 
rocephalus palustris, Rhacomitrium hypnoides, etc., in bogs. Hylocomium 
proliferum is of almost equal frequency at all altitudes. Near Akureyri 
it was collected at an altitude of 990 metres. Only sterile specimens 
have been found. 


321... Hylocomium pyrenaicum (Spruce) Lindb. 
Hylocomium Oakesii (Sull.) Schimp. 


S. Iceland:' Krisuvik (Stp.)! 


322... Hylocomium parietinum (L.) Lindb. 
Hylocomium Schreberi (Willd.) De Not. 


E. Iceland: Berufjérdur!; Hornafjérdur!. S. Iceland: Common every- 
where!. Vestmannaey!. W. Iceland: Esja (Grl.;!); Stykkisholmur!; Bor- 
garfjordur!. NW. Iceland: Common !. 

Widely distributed in SE., S. and W. Iceland and on the peninsu- 
las of NW. Iceland. In E. Iceland it has not been found north of 
Berufjérdur, and it appears to be absent from the whole of N. Iceland. 
It usually grows on rather dry ground, for instance in birch coppices, 
where it almost everywhere forms an essential part of the vegetation, 
and on moss-grown slopes and in the Rhacomitrium-heath, but it occurs 
also on damper ground, especially on knolls in bogs. It rarely ascends 
higher than the birch. In NW. Iceland it was collected up to about 
300 metres above sea-level, and in Esja it grew among Rhacomitrium 


THE BRYOPHYTA OF ICELAND. b43 


hypnoides at an altitude of about 450 metres. Only sterile specimens 
have been found. 


323... Hylocomium loreum (L.) Br. eur. 


W. Iceland: Esja (Grl.;'), common up to about 400 metres above 
sea-level; commion in the coppices of the district of Borgarfjordur!: 
Berserkjarhraun; Arnarbotn; Budahraun (H. J.)!; Stykkisholmur!; Bryn- 
judalur (Grl.)!; Hafnarfjéårdur (Grl.;!); Svinahraun!; Kolvidarhol!, up to 
about 350 metres. NW. Iceland: Sugandafjérdur!, up to about 250 me- 
tres; Dyrafjérdur!, up to about 200 metres. S. Iceland: Widely distri- 
buted!; Thingvellir!; Laugardalur!. 

Widely distributed in the whole of the southern and south-western 
part of Iceland as far as Borgarfjordur and Snæfellsnes. It also occurs 
rather plentifully in some of the fjords of the north-west. It grows 
sometimes in clefts among fallen blocks of rock, sometimes on slopes 
together with other Hylocomium spp., Dicranum scoparium, Rhacomi- 
trium spp., etc., or in the birch coppices. It was collected in fruit 
near Holt in S. Iceland, where a few old, empty capsules were found. 


324... Hylocomium triquetrum (L.) Br. eur. 


Iceland (Koenig). E. Iceland: Frequent from Berufjérdur south- 
wards!:… N. Iceland:  Asbyrgi!; near Eyjafjordur (O. D.;!); Ljosavatn 
Grl.;!); common west of Bléndudalur!. NW. Iceland: Rather common 
in all the fjords!. W. Iceland: Rather common everywhere (Grl.;!). Vest- 
mannaecy!. S. Iceland: Common!. 

Commonly distributed from Berufjérdur in E. Iceland throughout the 
whole of S., SW. and W. Iceland, and in the majority of the fjords of 
the: north-west.… It is rather rare in North Iceland proper, cast of 
Blåndudalur, and occurs, as a rule, only scantily in the coppices or in 
the Betula nana heath. 

Its growth is similar to the other species of Hylocomium, with 
which it is almost always found associated, especially in coppices and 
on moss-grown slopes. It has the same altitudinal distribution as H. 
loreum and H. parietinum, as it hardly ascends higher than to the upper 
limit of the birch. In Esja it is common up to 350—400 metres, near 
Kolvidarhol it was found in company with H. loreum at 350 metres, 
and in NW. Iceland it everywhere accompanies the birch to its upper 
limit at about 250 metres. Only sterile specimens have bcen found. 


325... Hylocomium squarrosum (L.) Br. eur. 


Very common over the whole of Iceland. 

It grows both on a damp substratum, on stony flats as on boggy 
ground, and on a more dry substratum, associated with other Hyloco- 
mium spp., Hypnum uncinatum, etc. It is most frequent in the low 
land, but occurs, however, also at rather considerable altitudes. In 
Berufjardarskard it was found even at 500—600 metres, near Dyra- 
fjordur at 330 metres and in S. Iceland in many places up to about 


544 A. HESSELBO 


500—550 metres above sea-level. Only sterile specimens have been 
found. 


326... Hylocomium rugosum (Ehrh.) De Not. 


Widely distributed on dry grass or moss-grown slopes or flats, 
usually mixed with other Bryophyta such as Hylocomium parietinum, H. 
proliferum, H, squarrosum, Rhacomitrium hypnoides, Dicranum scoparium, 
Ptilidium ciliare, etc., rarely unmixed in large mats. It has the same 
altitudinal distribution as H. triquetrum, and hardly ascends higher 
than about 300—400 metres above sea-level. Only sterile specimens 
have been found. 


KEE FHESBRYOPHVSTESCOMMUNIRIES: 


sin the other Arctic and Sub-arctic countries, the Bryophyta play a 
Å very important part in the plant-covering of Iceland. They occur 
either as an essential component of, so to speak, all plant associations, 
and often in far greater numbers as regards species and individuals, 
than do the higher plants, or as distinct Bryophyte associations from 
which other plants are entirely absent, or in which they occur only as 
a subordinate component. 

As a basis of classification it comes natural to distinguish 
between (1) Lowland Formations and (2) Highland Formations. 
Within these two main groups the formations are again arranged 
according to their water-requirements, commencing with those which 
are the most water-loving. Naturally, no sharp limit can be drawn 
between these formations; on the contrary, all possible transitions 
between them are constantly met with, which makes a comprehen- 
sive survey extremely difficult. In addition to this, the investiga- 
tions are in many respects very incomplete, especially with regard 
to the history of development of the Bryophyte associations. Also, 
several of the formations occur both in the Highlands and in the 
Lowlands. This is especially the case with decidedly xerophilous 
formations like the Rhacomitrium heath and, partly, also other 
moss-heaths. 

Mountain plateaus and slopes situated above the growth-limit of 
the birch and the heather moor are in this paper considered as be- 
longing to the Highlands (the Alpine Region). The limit in question 
may lie at various levels and can, of course, never be drawn 
sharply, but is in many cases dependent on the structure of the 
mountains. The flat land along the coasts and at the bottom of 
the valleys rises, as a rule, in terraces inwards towards the high 


546 A. HESSELBO 


land. Ås far upwards as about 200—400 metres the slopes and 
flats are covered with heaths and birch-coppices or bogs respec- 
tively. . Then usually follow steep, bare slopes of gravel or abrupt 
cliffs, and the plateaus which follow these bear the scanty plant- 
covering of the rocky flat. 

The conditions, however, differ somewhat in different parts of 
the country. In NW. Iceland there is almost no low land with the 
exception of the narrow stretch of coast along the narrow, deep 
fjords. The steep, often vertical cliff-faces rise to a height of 500— 
600 metres and have a talus of débris (Urd) at their foot, and cliffs 
rising in terraces occur almost exclusively at the head of some of 
the fjords. The entire surface at the top of the mountain belongs 
lo the rocky flat, and the associations characteristic of the latter 
often occur on the slopes and flats right down to the level of the 
sea, especially on slopes facing north. 

In S. Iceland, on the other hand, in sheltered valleys, localities 
are met with as far upwards as 500—600 metres above sea-level, 
which, according to the composition of the plant-covering, must be 
referred to the Lowland formations. 


I. THE LOWLAND FORMATIONS. 
The Littoral Bryophyte Vegetation. 


There are only very few species which are restricted to the 
immediate vicinity of the sea, or which grow by preference in 
the neighbourhood of the coast. Schistidium maritimum and Ulota 
phyllantha doubtless occur wherever there are rocky coasts, and 
are rarely found many hundred metres from the coast. They 
chiefly occur along the sea-coast itself, and at the entrance to 
the larger fjords, and decrease in frequency inwards towards the 
head of the fjord. Near Berufjérdur, for instance, both species are 
very common around Djupivogur, but are entirely absent from the 
inner part of the fjord, and this is also the case in Seydisfjordur, 
Åround the broad Isafjérdur they are common as far inwards as Arn- 
gerdareyri, but are, as a rule, absent from the narrow branch-fjords. 
Both species grow on the coast rocks, but also frequently descend 
to the ground. On the island of Vestmannaey Ulota was very 
common on knolls in fields, where the grass-covering had been 
torn up by the wind, and Schistidium maritimum has been found 
in several places on damp sandy soil. 


THE BRYOPHYTA OF ICELAND 547 


Ørthotrichum Blyttii occurs here and there on coast rocks, or at 
any rate in the immediate neighbourhood of the coast. Pottia Heimii 
is commonly distributed along all flat coasts and Amblystegium 
littorale occurs here and there on damp sandy soil. 

Besides these littoral species proper, there åre several others 
which occur especially abundantly on the coast, but are also distri- 
buted in many other localities. Bryum argenteum and B. capillare 
grow especially below cliffs inhabited by sea-fowl, but their pre- 
sence there is, no doubt, as much due to the abundant supply of 
organic manure at io the proximity of the sea. Desmatodon lati- 
folius, Leptobryum pyriforme and Tortula ruralis are also typical of 
damp sandy soil and Hypnum polygamum of saline, boggy soil. 
On damp sandy soil on the beach low, scattered moss-cushions 
consisting of stunted individuals of many different and more casual 
species are often met with, for instance, Distichium montanum, Di- 
dymodon rubellus, Ceratodon purpureus, Encalypta rhabdocarpa, 
Myurella julacea, Bryum spp., etc. 


Hydrophilous Bryophyte Formations. 


The water originating from rain and mist or from melting 
snow will, according to the local conditions and the inclination, 
eilher spread out over the ground and — if it consists of humus 
or sand — sink into it, or seek outlets in brooks and rivers. Where 
the water remains standing without being able to find a sufficient 
outlet, a bog vegetation is developed, or, in deeper hollows, a lake 
is formed. Part of the water which sinks down to the solid rocky 
substratum continues its way downwards along the latter until it 
sooner or later issues as a spring. Stagnant water contains humus 
substances, is not well aerated, and is warmed by the sun, while 
running water, both in streams and springs, is pure and clear and 
contains oxygen and carbonic acid. Running water has a very low 
temperature during the summer, at the most 42—6?. Therefore 
the composition of the vegetation on boggy ground differs entirely 
from that on ground along streams and near springs, inundated or 
saturated by the pure water. 

The hydrophilous formations may be classified as follows: — 


548 A. HESSELBO 


AA The Bryophytenvesetation of Pure" Water: 
a. The Bryophyte Vegetation of Running Water and of 
Lakes. 
b. The Bryophyte Vegetation of Inundated Gravelly Soil 
(slightly inclined ground and banks of streams). 
c. The Bryophyte Vegetation on Muddy Soil near Springs (Dy). 
BEM ERB myop Inkyate VIE se ta brom kob oss yes om 
CÆTherBryoplyternvesetatronfofd am pis andjÆæson 
DEM blerB rype se Fat one arket penmes 


The Bryophyte Vegetation of Running Water and of Lakes. 


The Bryophyte Vegetation of Lakes”is”extremely ”scanty, 
and often entirely absent. Along the shores of shallow lakes the 
surrounding swamp-vegetation may extend right into the water, 
especially species such as Hypnum giganteum, H. exannulatum and 
H. scorpioides, while rocky shores are usually quite bare of sub- 
merged Bryophyte vegetation or bear, here and there, one or other 
of the species growing in streams, such as Fontinalis antipyretica or 
a Hypnum-species.…. How far Bryophyta (for instance, Fontinalis) 
may occur in deeper water has not been investigated. 

Glacier-rivers, on account of their torrential current and 
changeable course, as well as of the low temperature of their clay- 
containing water, are entirely devoid both of Bryophyte vegetation 
and of higher plants. 

Brooks and Rivers. Where the bottom consists of loose 
gravel and smaller stones which are kept by the water in constant 
motion, Bryophyte vegetation is entirely absent. But if the bottom 
is of firm rock or larger stones it is, as a rule, covered with ex- 
tensive moss-carpets which usually consist of Hypnum ochraceum 
or Fontinalis antipyretica, which may, either separately or together, 
cover large areas of the bottom. In very strong currents H. ochra- 
ceum forms the highly elongated and often thread-like var. filiforme 
which, together with Rhynchostegium rusciforme var. atlanticum, is 
a characteristic species of torrential rivers on much inclined ground. 
H. ochraceum var. filiforme is found especially in larger streams, 
where it is then the only species occurring, while Rhynchostegium 

7 In this paper no distinction has been made between swampy soil, marshy 


soil and boggy soil, the term "boggy” includes all three kinds of soil. The Danish 
term in all three cases is "sumpet”. 


THE BRYOPHYTA OF ICELAND 549 


appears to prefer smaller rivers with large loose blocks of rock at 
the bottom. 

Several other Bryophyta occur either mixed with the above or 
occasionally as dominants in the Bryophyte vegetation. Schistidium 
rivulare is very common and often occurs abundantly. Hypnum 
dilatatum, H. alpinum and H. alpestre are common in the northern 
and eastern parts of the country, but are absent from or are rare in 
other parts. In the river Ellidarå near Reykjavik the bottom was 
in several places covered with Fontinalis androgyna. In shallower 
water, and especially on inundated rock-surfaces in waterfalls, 
several other species also occur. Almost everywhere in such localities 
are found: Haplozia cordifolia, Scapania undulata, Brachythecium 
rivulare, Rhynchostegium rusciforme, Hypnum palustre, H. falcatum, 
Philonotis fontana, Mniobryum albicans var. glacialis, Bryum Duvalii 
and Bryum ventricosum together with some of the, species which 
thrive best in places where they are outside the water during a part 
of the year. 

Stones and rocks which protrude above the surface of the 
water and are inundated only during specially high water-levels 
are usually covered with a dense moss-carpet consisting of nume- 
rous species. In addition to the majority of the species of river- 
bottoms there occurs also an abundance of those belonging to damp 
rocks. Among the species which are rarely absent from such lo- 
calities may be meutioned: Hypnum falcatum, H. filicinum , Schisti- 
dium rivulare, S. gracile, Rhacomitrium aciculare, R. sudeticum, Blindia 
acuta, Dichodontium pellucidum, Amphidium Mougeottii, Haplozia 
atrovirens and Mnium punctatum, and in North and East Iceland, in 
addition to the above-mentioned, Philonotis seriata and Hypnum 
decipiens are to be found. ÅA somewhat different vegetation occurred 
in the river Rekyirdalså in SW. Iceland, which is so well supplied 
from hot springs that in the beginning of June the water had a 
temperature of about 12%, while that of the streams flowing into 
it on both sides was only 42—5?. Hypnum ochraceum, which is else- 
where so common, was entirely absent, and Schistidium rivulare 
occurred very scantily. On rock-surfaces and larger stones Hypnum 
palustre, Brachythecium rivulare and Rhynchostegium rusciforme grew 
abundantly, the last had also set fruit. Fontinalis longifolia covered 
the bottom in many places, and on stones protruding above the 
surface of the water Hypnum filicinum and Schistidium gracile were 
growing. In the cold tributary streams the usual vegetation occur- 


550 A. HESSELBO 


red, consisting of Hypnum ochraceum, H. falcatum, Schistidium ri- 
vulare, Rhynchostegium rusciforme var. atlanticum and Scapania 
undulata. 


The Bryophyte Vegetation of Inundated Gravelly Soil (slightly 
inclined ground and banks of streams). 


Where the water flows down the mountain sides and spreads 
over slightly inclined gravelly ground, and along the flat banks of 
streams which are periodically inundated, a vegetation composed 
mainly of Bryophyta is developed. The most conspicuous spe- 
cies growing on this gravelly ground, and one which is hardly 
ever absent, and is often the most abundant constituent of the 
moss carpet, is Philonotis fontana, and after it this community 
may be designated the Philonotis association (Ostf., 1907). 
Very frequently many other species also occur, either as an ad- 
mixture in the Philonotis carpet, or replacing it over smaller or 
larger areas. In NW. Iceland P. fontana is often replaced by P. 
seriata.. Mniobryum albicans var. glacialis is almost as common 
as Philonotis; and Bryum ventricosum, B. Duvalii, Brachythecium 
rivulare and Acrocladium cuspidatum are also among those species 
which are rarely absent. 

Ås a few examples will best show the composition of this ve- 
getation, the following are given: — 

1. The Ellidarå near Reykjavik: The mossy fringe along 
the banks consisted of Philonotis fontana, Dichodontium pellucidum 
and Acrocladium cuspidatum. 

2. Husavik, along aa" stream: A- broad belt "of Cinclidium 
stygium together with some Paludella squarrosa. 

3. Husavik, along the banks of smaller streams on a slope: 

Mniobryum albicans var. glacialis, Mnium punctatum, Philonotis 
fontana, Marchantia polymorpha together with some Bryum Duvalii 
and Thuidium lanatum. 

4. Berufjårdur, along a stream: Philonotis fontana, Mnium 
Seligeri, Hypnum giganteum and Mniobryum albicans var. glacialis. 

5. Svinaskard, along a small stream: Philonotis seriata, 
Scapania undutata, Chiloscyphus polyanthus and Pellia Neesiana. 

6. Kaldalon (NW. Iceland), im a small stream and on 
gravelly ground over which water was flowing: Philonotis seri- 
ata, P. fontana, Dicranella squarrosa, Mniobryum albicans var. 


THE BRYOPHYTA OF ICELAND SET 


glacialis, Bryum Duvalii, Chiloscyphus polyanthus var. fragilis, Harpan- 
thus Flotowianus and Pellia Neesiana. 

7... Kaldalon, along the banks of a stream: Hypnum calli- 
chroum, Sphagnum teres, Harpanthus Flotowianus and Cephalozia 
bicuspidata var. Lammersiana. 

8. Reykjavik, on ground over which water was flowing: 
slightly inclined surface with streamlets. The banks of the streamlets 
were fringed with Philonotis fontana, Mnibryum albicans var. gla- 
cialis, Bryum ventricosum, B. Duvalii and Acrocladium cuspidatum. 

9. Gnupsdalur (NW. Iceland), gravelly ground, over which 
water was flowing, streamlets flowing down the sides of the valley: 
Philonotis seriata, P. fontana, Dicranella squarrosa, Haplozia cordi- 
folia, Chiloscyphus polyanthus var. fragilis, Scapania undulala and 
S. paludosa. Haplozia formed large black or greenish-black, and 
Scapania paludosa very large reddish-brown patches in the water, 
which was a few centimetres deep. 

In North Iceland several species of Hypnum occurred abundantly 
in gravelly soil over which water was flowing, and often formed pure 
Hypneta. Especially do Hypnum exannulatum var. purpurascens, 
H. falcatum, H. decipiens and sometimes H. stramineum and H. sar- 
mentosum occur in such localities. 


The Bryophyte Vegetation on Muddy Soil near Springs (Dv). 


The bright-green moss-carpet around and below the point of 
issue of the springs described by Helgi Jønsson (1900, p. 24; 
1895p:73) "as! moss bogs” (Icelandic Dy) are one of the” most 
frequently occurring and characteristic plant communities of the 
Arctic and Sub-arctic regions. They were first mentioned by Gr&n- 
lund (1877, p. 330) as mats of sapgreen mosses along small streams 
on mountain sides, consisting of Bartramia fontana, Webera albi- 
cans, Brachythecium rutabulum var. rivulare and Hypnum uncina- 
tum, a description which includes both the moss bogs and the 
moss vegetation along smaller streams, which have also much in 
common. 

Moss bogs develop in muddy soil, in all places where the 
ground water emerges as springs. They occur both on mountain 
slopes and at the bottom of valleys, occasionally also at higher 
levels, on rocky flats, but they are especially numerous on the 
lowest mountain slopes and in the flat boggy land below these 


DØ A. HESSELBO. 


slopes. There they lie irregularly scattered in bright-green patches, 
differing greatly in size, and conspicuous even at a distance 
by their colour which is distinct from that of their surround- 
ings. Higher up on the slopes at the foot of the bluffs, they. fre- 
quently form continuous belts in connection with the mossy fringe 
along the downward -flowing streams (Helgi Jonsson, 1900, 
p. 25). 

The vegetation consists of nearly the same species as those 
found on inundated gravelly soil, and the external conditions for 
the plants are also essentially the same in both places, since it is 
the cold, well-aerated water which determines the composition of 
the vegetation. During summer the temperature of the water is 
usually only 49—5%, and this low temperature acts as a great 
check to the growth of higher plants, and is therefore indirectly 
favourable to the moss vegetation. The difference between the two 
vegetations is especially due to the substratum, that of the vege- 
tation of inundated ground and stream-banks consisting of gravel, 
while the moss bog developes on mud. For this reason also, some 
of the species which occur in the gravelly soil are absent from the 
moss-bogs proper, for instance Hypnum. falcatum, H. decipiens, Di- 
cranella squarrosa, Philonotis seriata, Haplozia cordifolia, Scapania 
paludosa and Chiloscyphus polyanthus var. fragilis. 

On flat ground moss bogs are, as a rule, almost circular in 
shape, and the species are arranged concentrically around the point 
of issue of the spring, whence the water gradually oozes through 
the moss carpet, and spreads out over the surrounding boggy 
ground. On sloping ground the moss bog is usually oblong in 
shape, and the spring emerges at its upper end (Fig. 19). 

In the majority of cases the Bryophyte vegetation near springs 
(the Dy) is a Philonotis-Mniobryum association, in which other species 
of Bryophyta and a few higher plants occur scattered (Ostf., 1907, 
p. 69). On the soft mud around the point of issue of the spring 
Mniobryum albicans var. glacialis 





— and often covering the latter 
usually grows, and outside the latter there is a broad belt of Phi- 
lonotis fontana, which may be intermixed with or replaced by other 
mosses; and scattered in the moss carpet are a few flowering plants 
and vascular cryptogams, especially Montia rivularis, Sazifraga 
stellaris, Epilobium spp. and Equisetum palustre. 


THE BRYOPHYTA OF ICELAND BAG 





Fig. 19. Kaldalon (NW. Iceland).  Slightly inclined ground below the stony slope. 


The light-coloured stripes are moss bogs, the darker, boggy ground or, on somewhat 


higher levels. patches covered with Vaccinium-Empetrum heath, 


Besides the above-mentioned mosses the following plants are 
characteristic of the moss bog: — 


Brachythecium rivulare M. Seligeri 

Acrocladium cuspidatum M. cinclidioides 
Hypnum Kneiffii M. affine var. elatum 
Bryum ventricosum Marchantia polymorpha 
B. Duvalii Pellia Neesiana 


Mnium punctatum 


But also other Bryophyta from bogs may occur intermixed 
with these, especially towards the edge, where the moss bog gradu- 
ally passes into the common bog-vegetation. But of course all pos- 
sible transitions also occur, between moss bog and bog vegetation 
as well as between moss bog and the vegetation of the banks of 
streams, all according to the nature of the underlying substratum. 

Ås some examples will best show the composition of the moss 
bogs, the following are given: — 

1. Seydisfjårdur, numerous moss bogs on an almost hori- 

36 


5H4 A. HESSELBO 


zontal, boggy surface, below the slope of the mountain (Fig. 20). 
The vegetation was almost identical in all of them: In the middle, 
Mniobryum albicans var. glacialis was found intermixed with Montia 
rivularis, then came a belt of Philonotis fontana, often greatly inter- 
mixed with Marchantia polymorpha and with scattered plants of 
Sawifraga stellaris, Equisetum palustre and Epilobium spp. In some 
moss bogs a pure Hypnetum composed of H. exannulatum var. 
purpurascens or H. Kneiffii had been developed. 





Fig. 20. Moss bog near Scydisfjårdur. The light-coloured patch in the middle is 


Mniobryum albicans var. glacialis. In the foreground and on the sides Philonotis fontana. 


2. Husavik, moss bog on slightly inclined ground: In the 
middle Mniobryum albicans var. glacialis and Marchantia polymorpha 
were found, and around this a belt of Bryum ventricosum f. major, 
Mnium affine var. elatum and Philonotis fontana, and outside it all 
an almost pure belt of Philonotis. 

3. Breidabolstadr, slightly inclined grassland with scattered 
moss bogs: In the middle Mniobryum albicans var. glacialis grew inter- 
mixed with Bryum ventricosum f. major; around this was a belt of 
Philonotis fontana, and outside it all Mnium affine var. elatum and 
Acrocladium cuspidatum. Some smaller moss bogs below the slope 


THE BRYOPHYTA OF ICELAND 555 


were composed of Brachythecium rivulare, Mniobryum albicans var. 
glacialis, Philonotis fontana, Bryum ventricosum, Mnium Seligeri and 
Mnium affine var. elatum, which grew in patches intermixed. 

4. Holt (S. Iceland), moss bog below a slope: Acrocladium 
cuspidatum, Mniobryum albicans var. glacialis, Bryum ventricosum f. 
major, Mnium punctatum, M. Seligeri and Pellia Neesiana grew inter- 
mixed, sometimes as single plants and sometimes in patches. In 
the middle Montia rivularis grew among the mosses. 

The majority of the mosses growing near springs form special 
cold-water forms which are particularly characterized by their 
highly elongated, slightly branched stems of almost equal height, 
by which feature they all assume a homogeneous habit. Brachythe- 
cium rivulare forms a form with long, erects shoots of equal height 
and with appressed leaves. Philonotis fontana forms, when growing 
in very cold water, a highly elongated, hardly tomentose form with 
appressed, shortly pointed leaves with short leaf-celis and very broad 
nerve (forma adpressa). The cold-water form of Bryum ventricosum 
(f. major) is also higher in growth and more vigorous than is the 
bog-form, has stems bare of tomentum with large and broad, 
spreading leaves. Mniobryum albicans var. glacialis is also much more 
vigorous than the type, with larger and broader leaves. Mnium 
spp. develop elatum-forms with erect, sterile shoots of equal height. 


The Bryophyte Vegetation of Boggy Soil. 


The bog vegetation is extensively distributed about Iceland, 
especially in the low land, and in many places it occupies almost 
the whole tract of land between the mountain slopes and the sea. 
The lower mountain plateaus and slightly inclined slopes up to a 
height of about 300—400 metres are also to a great extent covered 
with bogs (Sedge-bogs). The mode of formation of these bogs has 
been described by Helgi Jonsson (1895, p.45). They originate 
partly from the further development of the vegetation of the springs 
and the banks of streams, and partly from shallow lakes first 
forming in depressions and afterwards becoming gradually filled up 
with plants in connection with sand transported by the wind. In 
the former case a drier bog (myri, pl. myrar) is formed, the sedges 
superseding the mosses; in the latter case a wet bog (fl6i, pl. flåar) 
is first formed, which, by gradually becoming overgrown with plants, 
develops into a ''fmyri”?, 


1 For a description of "myri” and "fléi” see also Thoroddsen's Physical 
Geography of Iceland, ante p. 323. 36” 


590 A. HESSELBO 


In "Die Moose des Sarekgebietes” (1910, p. 248) C. Jensen 
classifies the bog vegetation in four formations: The Willow bog, 
the Sedge bog, the Moss bog and the Peat bog. This classification 
cannot be entirely adopted as regards Iceland. Willow bogs proper 
(willow coppices in boggy soil) do not appear to be developed there. 
True, scattered individuals or small groups of Salix (S. lanata, S. 
phylicifolia and S$. glauca) are often found in the sedge bogs, espe- 
cially in NW. Iceland, but the moss vegetation there does not 
differ in any respect from that of the surrounding sedge bog. 

The vegetation of boggy soil is composed mainly of mosses 
and sedges, greatly varying in abundance. Sometimes the sedges 
are the dominant plants, and then the moss carpet is less conspicuous 
at the bottom, among the Carex and Eriophorum spp.; sometimes 
there is a continuous moss-carpet in which the higher plants grow 
scattered, and then the one formation may be termed a sedge bog, 
and the other a moss bog, which, however, does not correspond 
with Helgi Jånsson's definition of a moss bog, but appears to 
agree with Warming's description (1887, p. 132). They are the 
same mosses which occur in the same manner in both formations, 
and all possible transitions are found between these. 

Helgi Jénsson (1900, p. 20; 1905, p.9) classifies the bog 
vegetation into Star-grass bogs and Moss bogs. The latter, which 
include the vegetation along streams, on inundated ground and near 
springs (Dy), and also some of the moss bogs in Warmings defi- 
nition, have been treated in the preceding section. The star-grass 
bogs are again divided into "”Myrar” and "”Fléar”. In the Myri the 
ground water stands on a level with the surface, the soil contains 
acid humus and is so closely interwoven with rhizomes and roots 
that it bears one like a carpet. In the Fléi the ground water 
stands above the surface, the soil is muddy and produces a more 
scattered vegetation of Carex and Eriophorum tufts. As it is chiefly 
the height of the ground water which determines whether a Myri 
or a Fléi is developed, and the Fl6i is. formed by the filling up of 
a shallow lake, there occur all transitions between lake and Fl6i, 
and through further development, between Fl6i and Myri. 

The Elo Fis spoork inkmosses Rhh er soft km uddyæsolkisænot 
favourable to their growth, and it is only as an exception that 
some of the species are found which grow along the banks of 
lakes, but often the moss-covering is entirely absent. Hypnum 
scorpioides and H. giganteum are the most frequent species which 


THE BRYOPHYTA OF ICELAND DD 


occur everywhere in pools and channels. Hypnum exannulatum, H. 
revolvens var. Cossoni and Acrocladium cuspidatum are also frequent. 
In many places in East Iceland Cinclidium styginm was found 
abundantly in Fléar, while Hypnum cordifolium and H. fluitans were 
found in a few places only. 

The moss-covering of the Myri is characterized by the fact of 
its being rich in species, which grow sometimes greatly intermixed, 
sometimes separately in large and small patches. The ground is 
usually knolly, and there is a great difference in the vegetation 
between and upon the knolls, the most hygrophilous species growing 
on the wet ground between the knolls, while on the top of the 
knolls mesophilous and xerophilous forms are met with, and be- 
tween these all transitions occur, according to the height of the 
knoll and the degree of moisture. They are nearly the same 
species which are everywhere the most abundant constituents of 
the Myri vegetation. The following species occur on the ground 
between the knolls: — 


Musci veri 


"Hypnum revolvens Meesea triquetra 
Fy TES intermedium £Paludella squarrosa 
= stramineum "Catoscopium nigritum 
= stellatum =Cinclidium stygium 
É = polygamum =Mnium cinclidioides 
Foo sarmentosum —— punciatum 
= Lindbergii oz affine 
 r= exannulatum Fo Seligeri 
Fo == giganteum Sy subglobosum 
= Richardsonii =Bryum ventricosum 
= falcatum "Splachnum vasculosum 
= trifarium g == sphæricum 
— turgescens "NE.) Dissodon splachnoides 
€Camptothecium nitens DSEGj TØ AES Wahlenbersgii 
=£Brachythecium rivulare = virens 
ENE.) Thuidium lanatum "| Dicranum angustum 
"Acrocladium cuspidatum "Fissidens osmundoides 
=Philonotis fontana "Dichodontium pellucidum 
=Meesea trichoides Campylopus Schimperi 
Sphagna 
FBb-snun teres SPhagnum papillosum 
= rubellum inundatum 
E == Warnstorffii squarrosum 
— fimbriatum — acutifolium 
z = medium riparium 
ig = Girgensohnii — Lindbergii 


— subnitens 


558 A. HESSELBO 


Hepaticæ 


=Pellia Neesiana =Lophozia quinquedentata 
"Aneura pinguis = quadriloba 

£ —. multifida = Schultzii 
=Cephalozia bicuspidata — Hornschuchiana 

ig = pleniceps ”Scapania irrigua 
=Cephaloziella Hampeana = uliginosa 

ag — rubella = undulata 
Odontoschisma elongatus =Blepharostoma trichophyllum 
=Alicularia scalaris =Anthelia julacea 


=Lophozia Kunzeana 


In the above list an asterisk is prefixed to those species which 
are common over the whole of Iceland, and which are met with 
in almost every considerable boggy tract. Some of the species are 
common in some districts and absent from others, for instance 
Thuidium lanatum and Dissodon splachnoides, which are very com- 
mon in North and East Iceland, but are rare or entirely absent 
from other parts of the country, and Campylopus Schimperi, which 
is found only in the southern part. 

The mosses often grow so much intermixed that a small collec- 
tion may contain 15—20 or even more species, but often smaller 
and larger patches are found dominated by a single species. Thus 
Catoscopium, Philonotis, Cinclidium or Mnium bogs occur, in each 
of which one of these species is the dominant member. On very 
damp ground the mass of the vegetation is formed by, for instance, 
Mnium cinclidioides or Cinclidium stygium, and in South Iceland by 
Mnium Seligeri. In North and East Iceland it is especially Thu- 
idium lanatum which is often the most conspicuous species. 

Sphagnum spp. grow almost always in scattered tufts, and 
Sphagnum bogs proper are rarely met with. Helgi Jonsson 
(1900, p. 25) records such bogs from Snæfellsnes, where they oc- 
curred on sloping ground and were composed of Sphagnum teres, 
S. Warnstorffii and S. Girgensohnii intermixed with Paludella squar- 
rosa, Hypnum stramineum, Hylocomium squarrosum and Polylrichum 
commune. I have only seen a similar vegetation near Barkarstadr 
in South Iceland, where, on a wet boggy slope stretching down 
towards a river, there was an almost continuous growth of Sphag- 
num rubellum and SS. Warnstorffii intermixed with a few other 
mosses, especially Thuidium lanatum, Hylocomium squarrosum and 
Hypnum stramineum. 

Sphagnum spp. often play an important part in the knoll- 
formation of the myri, the large Sphagnum-cushions being inter- 


THE BRYOPHYTA OF ICELAND ' 559 


woven with mosses and hepatics, and different plants such as 
Vaccinium spp., Betula nana, Comarum palustre and several Carices 
then establishing themselves upon these cushions. 

The numerous hepatics which occur are everywhere found 
interspersed in the moss carpet, especially in that upon the knolls, 
and, as regards all the species mentioned below, doubtless also in 
the ground vegetation between the knolls. 

The Bryophyte Vegetation of Knolls! is dependent upon 
the varying degree of moisture, which is greatest on the ground 
between the knolls and least in their uppermost part. Some of the 
Bryophyta of the ground do not ascend higher than to the base 
of the knolls, while others, especially many of the pleurocarpous 
mosses, occur with equal frequency upon and between the knolls. 
In addition to the latter mosses the following species are found upon 
the knolls: — 


=Hypnum uncinatum =Mnium hornum 

== imponens "Rhacomitrium canescens 
=Climacium dendroides £ — hypnoides 
=Hylocomium squarrosum =Ditrichum flexicaule 
ig — parietinum £Dicranum scoparium 
x — proliferum 5 — congestum 
Orthothecium chryseum (N. Icel.) É — elongatum 
=Polytrichum alpinum £Lophozia lycopodioides 
x >= strictum £Plagiochila asplenioides 
s = juniperinum =Ptilidium ciliare, 


=Aulacomnium palustre 
and besides these, numerous other species which belong to other 
formations and must rather be regarded as casual visitors. 

The following examples will show the composition of the bog 
vegetation in the different parts of Iceland: — 

1... Skålbolt im SW. Iceland. The country here over a wide 
expanse is covered with vast boggy areas in which small, protrud- 


7 According to my observations these knolls are formed either by soil accu- 
mulating in tufts of Sphagnum, etc., in which other plants take root, among 
others sedges, when soil again accumulates, or by soil accumulating directly 
around roots of sedge-tussocks. This accumulation of soil and plants continues 
till big knolls are formed, some 60—70 cm. in height. These knolls occur in all 
stages of development, from tufts or tussocks with a slight accumulation of soil 
around their roots to completely rounded knolls in which the tussocks have en- 
tirely disappeared. In Danish "Tue” stands both for "knoll” and for "tuft” and 
tussock”, which is very convenient when referring to one of the various stages 
of these "knolls”. But the rendering into English has been difficult, though in each 
case the term most appropriate to the stage of development of the "knoll” has 
been carefully chosen. 


560 A. HESSELBO 


ing masses of rock (holt) occur scattered. In the wettest parts of 
the myri grew Hypnum scorpioides, H. giganteum and some H. re- 
volvens v. Cossoni. The moss-covering of the ground between the 
knolls consisted mainly of Hypnum revolvens,, H. stellatum, H. Lind- 
bergii, H. stramineum, Acrocladium cuspidatum, Hypnum polygamum, 
Hylocomium squarrosum, Camptothecium nitens, Mnium Seligeri, M. cin- 
clidioides, Philonotis fontana and Bryum ventricosum, all of which grew 
intermixed. Of more scanty occurrence were Hypnum sarmentosum, 
Paludella squarrosa, Meesea trichoides, M. triquetra, Cinclidium stygium, 
Mnium punctatum, Splachnum vasculosum, S. sphæricum, Fissidens 
osmundoides, Oncophorus Wahlenbergii, Alicularia scalaris and Aneura 
pinguis. The following hepatics grew both between and upon the 
knolls: — Lophozia Kunzeana, L. quinquedentata, Scapania irrigua, 
Cephalozia pleniceps, C. bicuspidata and Cephaloziella Hampeana. 
The knolls were usually formed by Sphagna, especially by S. teres, 
S. Warnstorffii and S. rubellum, and in very wet parts of the myri 
also by $. medium. Woven into and above the Sphagnum-cushions 
grew Hypnum uncinatum, Hylocomium squarrosum, H. proliferum, 
H. parietinum, Climacium dendroides, Aulacomnium palustre, Dicra- 
num scoparium var. turfosum, Ptilidium ciliare, Plagiochila asplenioides, 
and, uppermost, Rhacomitrium hypnoides and R. canescens. 

2. Håskuldstadr in E. Iceland. Knolly sedge-bogs with a 
rich vegetation of mosses between the knolls, along the river at the 
bottom of the valley. The moss carpet of the ground between the 
knolls consisted of Hypnum revolvens, H. giganteum, H. stellatum, Åcro- 
cladium cuspidatum, Mnium cinclidioides, Cinclidium stygium, Mnium 
affine var. elatum, Paludella squarrosa, Philonotis fontana, Meesea tricho- 
ides, M. triquetra, Bryum ventricosum, Aulacomnium palustre, Catoscopium 
nigritum, Oncophorus Wahlenbergii, Dichodontium pellucidum and 
Aneura pinguis. All the Bryophyta, as a rule, grew intermixed, 
but some of them frequently formed large, pure growths, especially 
Catoscopium nigritum and in the wettest parts Mnium cinclidioides 
and Cinclidium stygium. Upon the knolls, which were sometimes 
formed by Carex, sometimes by Sphagnum, grew Camptothecium ni- 
tens, Thuidium lanatum, Hypnum stramineum, H. uncinatum, Hylo- 
comium squarrosum, Sphagnum teres, S. Warnstorffii, and, upper- 
most, Hylocomium parietinum, H. proliferum, Dicranum scoparium, 
Rhacomitrium hypnoides and Scapania curta. Woven into the moss- 
covering, especially into that upon the knolls, were found Lophozia 
Kunzeana, L. quinquedentata, Scapania irrigua, Blepharostoma tricho- 


THE BRYOPHYTA OF ICELAND 561 


phyllum, Plagiochila asplenioides, Cephalozia bicuspidata and C. 
Hampeana. The two sedge bogs described above may be regarded 
as types, and by far the greater majority of the sedge bogs have 
very nearly the same vegetation. 

SØE Breidabolstadronrn tt Eljotshlide (SS Iceland) PA flat, 
boggy tract of meadow-land beneath the low mountain-slope, inter- 
sected by ditches and, lowest of all, by a low dike in order to keep 
the water-level at a suitable height. The ground was even, without 
knolls, and covered by a vigorous vegetation of Carices, with a moss 
carpet at the bottom. Occasionally there were also patches where 
the moss-covering was dominant, and the species of Carex occurred 
in this as scattered individuals. The species were comparatively 
few in number and were usually dominant in spots. The most 
frequent species were Philonotis fontana, Mnium Seligeri, M. cincli- 
dioides, Cinclidium stygium, Bryum ventricosum and ÅAcrocladium 
cuspidatum. Here and there patches occurred with a more mixed 
vegetation, which, in addition to the plants mentioned above, con- 
sisted of Hypnum revolvens, Meesea trichoides, Splachnum vasculosum, 
Lophozia quinquedentata, and perhaps of several other species. Here 
and there were tufts formed by Sphagnum rubellum and $. teres. 

A vegetation of about the same composition as that described 
above is very commonly met with in South Iceland, where tracts 
of meadow-land, by means of draining, irrigation or damming up, 
are kept at about the same ground-water level during the whole 
summer. The knolls disappear — or are not formed — and the 
moss-covering becomes poorer in species. 

4 Ljøsavatn in North Iceland... A rather .large-'sedge 
bog on very wet ground, on the slightly inclined slope of the moun- 
tain. The moss-carpet was composed of Cinclidium stygium, Mnium 
cinclidioides, M. Seligeri, Paludella squarrosa, Camptothecium nitens, 
Hypnum spp., Philonotis fontana, Bryum ventricosum, Thuidium 
lanatum, Lophozia Kunzeana and L. quinquedentata.  Sphagnum 
rubellum, S. teres, S. Warnstorffii and S. Girgensohnii occurred abun- 
dantly in large tufts. In the wettest parts, where the water reached 
above the surface, grew thick, extensive carpets of Hypnum exan- 
nulatum, H. stramineum, H. sarmentosum, H.giganteum and H. Ri- 
chardsonii, all in very vigorous development, and in fruit. The 
Hypnaceæ sometimes grew intermixed, but usually they occurred 
separately, and were only slightly mixed with other Bryophyta and 
Cyperaceæ. This formation corresponds undoubtedly with C. Jen- 


562 A. HESSELBO 


sen”s Amblystegium-bog (1910, p. 253) and occurs rather commonly 
in North Iceland, but more rarely in the other parts of the country, 
probably because in North Iceland the climate is more continental 
than in the rest of Iceland. These moss bogs are most frequently 
composed of Hypnaceæ, but often Paludella squarrosa, Cinclidium 
stygium or Mnium cinclidioides forms an essential part of them. 

5. Isafjårdur. The narrow strip of land, below the steep moun- 
tain slope (almost 600 metres in height) along the western side of 
the fjord, is covered with sedge bogs right to the head of the fjord. 
The most abundant constituents of the moss carpet between the 
knolls were Hypnum revolvens, H. giganteum, H. exannulatum, Palu- 
della squarrosa, Philonotis fontana, Mnium cinclidioides, M. punctatum, 
Cinclidium stygium and Bryum ventricosum. Here and there in the 
moss carpet grew Thuidium lanatum, Catoscopium nigritum, Meesea 
trichoides, M. triquetra (scantily) and Oncophorus Wahlenbergii. Splach- 
num sphæricum and S$. vasculosum grew on cow-dung. The knolls 
were formed by Sphagnum teres, S. Warnstorffi, Polytrichum strictum, 
P. alpinum, Mnium hornum and Dicranum scoparium. The following 
hepatics occurred woven into the moss-cushions: Lophozia quin- 
quedentata, L. Kunzeana, L. Hornschuchiana, L. quadriloba, Scapania 
irrigua, Blepharostoma trichophyllam, Harpanthus Flotowianus, Ce- 
phalozia pleniceps, Cephaloziella Hampeana and C. rubella. 

6. Seydisfjårdur. A boggy flat at an altitude of about 
350 metres. The ground was covered with a blackish-brown moss 
carpet composed of Hypnum giganteum, H. stramineum, H. sarmen- 
tosum, H. revolvens, H. Richardsonii, H. exannulatum and H. uncina- 
tum, occasionally alternating with patches of Mnium cinclidioides. 
Tufts of Sphagnum teres and S. Girgensohnii occurred frequently, 
and Splachnum vasculosum was common on dung. Carex spp. and 
Eriophorum were abundant without, however, occurring so densely 
as to cover the ground. This type of Hypnum bog, from which 
other plants are most frequently entirely absent, is common in 
North and East Iceland at elevations of about 200—500 metres, on 
flats where the snow lies long. The moss carpet is, as a rule, thin 
and bears distinct traces of having been long subject to the pres- 
sure of the snow-covering, the old stems being pressed closely 
together, and only the annual shoots growing up into the air. 


Peat Bogs. In the course of time sedge bogs usually develop into 
peat bogs. Gradually, as the peat layer becomes thicker, the dampness 


THE BRYOPHYTA OF ICELAND 563 


of the surface decreases, the most hygrophilous species disappear 
and some other species, chiefly mesophilous, take their place. Some 
of the latter are especially connected with this soil. The most 
marked species of peaty soil are Psilopilum lævigatum, Polytrichum 
gracile, P. juniperinum, Pogonatum polytrichoides, Mnium hornum, 
Catharinea undulata (also in warm soil) and Dicranella cerviculata. 
Some species which occur on other soils, but most abundantly on 
peat, are Philonotis tomentella, Pogonatum urnigerum, Funaria hy- 
grometrica, many Bryum spp. (especially B. inclinatum, B. affine, 
B. arcticum, B. lacustre and B. pallens), Pohlia nutans, Leptobryum 
pyriforme, Distichium inclinatum, Ceratodon purpureus and Dicranella 
crispa. The above-mentioned species are all mesophilous, a few 
are xerophilous also (for instance P.juniperinum and Ceratodon), 
and the majority of them occur also in sandy soil, but in Iceland 
the peat is often abundantly mixed with fine sand, transported by 
the wind. The two rare species, Trematodon ambiguus and Pogo- 
natum capillare var. dentatum, have also been found on peat. 

The most decided character-plant of peaty soil is Psilopilum 
lævigatum, which occurs everywhere on peat-flats which have been 
laid bare, and may, for instance, entirely cover the piled-up heaps 
of cut peat with its numerous capsules, often in association with 
Funaria hygrometrica. Another characteristic vegetation of localities 
like these consists of extensive bluish-green mats of low-growing, 
sterile Pogonatum urnigerum, interwoven with Alicularia scalaris. 

The following examples will show the composition of the peat 
bogs in the different parts of Iceland: — 

1. Seydisfjårdur. In a peat bog intersected by ditches: 
Psilopilum lævigatum, Leptobryum pyriforme, Pogonatum urnigerum, 
Polytrichum juniperinum and P. alpinum grew abundantly. Inter- 
mixed with the above occurred Bryum inclinatum, B. arcticum, B. 
purpurascens, Distichium montanum, Ceratodon purpureus, Scapania 
curta, Alicularia scalaris and Lophozia Wenzelii. 

2. Akureyri. Behind the town, at the foot of the mountain, 
were large expanses of bogs from which great quantities of peat 
had been removed. The bared flats and the piled-up heaps of cut 
peat were often entirely covered with Psilopilum and Funaria. 
Commonly occurring species were Polytrichum gracile, P. alpinum, 
P. juniperinum, Leptobryum pyriforme, Bryum inclinatum, B. affine 
B. lacustre, B. arcticum, Philonotis tomentella, Ceratodom purpureus. 
Dicranella crispa, Didymodon rubellus and Scapania irrigua. 


564 A. HESSELBO 


30 Grunnavikknearnkokulskjordursedee pil åayertor 
peat, here and there intersected with ditches. The following mosses 
were observed: Polytrichum alpinum, P. Swartzii, Mnium hornum, 
Pohlia nutans, P. cucullata (both common), Bryum purpurascens, 
Ceratodon purpureus, Dicranella cerviculata, D. crispa, Lophozia quin- 
quedentata and a few tufts of Tetraplodon bryoides. Psilopilum lævi- 
gatum was found only scantily and after a long search. 

4. Lundur in W. Iceland. On a damp peatflat, the upper- 
most layer of which was pared off, grew Dicranella Schreberi, 
Aongstræmia longipes, Bryum lacustre, B. inclinatum, B. æneum, Di- 
chodontium pellucidum, Philonotis fontana, Meesea trichoides and 
AÅneura pinguis. 


The Bryophyte Vegetation of Damp Sandy Soil. 


The damp sandy or gravelly ground along streams, inundated 
only in the case of especially high water-levels, is often covered 
with a moss carpet of a very mixed composition. Here liverworts 
play an especially important part, and very frequently pure liver- 
wort associations are met with, or other associations in which the 
liverworts are almost dominant. They are almost everywhere the 
same species which are the most abundant constituents of the moss 
carpet. The following Hepaticæ are the most frequent: Scapania 
subalpina, Alicularia scalaris, A. geoscypha, Lophozia quinquedentata 
Cephalozia bicuspidata var. Lammersiana, Anthelia julacea, A. Ju- 
ratzkana, and, especially in NW. Iceland, Harpanthus Flotowianus. 
Intermixed with the above occur a very great number of species 
of which the most frequent are Pellia Neesiana, Aneura pinguis, 
Haplozia atrovirens, Cephalozia pleniceps, Dicranella crispa, Dichodon- 
tium pellucidum, Aongstræmia longipes, Didymodon rubellus, Disti- 
chium montanum, Pohlia commutata, P. gracilis, Bryum pallens, B. 
inclinatum, B. affine, Philonotis tomentella, P. fontana, Pogonatum 
urnigerum and Timmia austriaca. A few less common species, 
such as Blasia pusilla, Eucalyx subellipticus and Distichium inclina- 
tum, also belong to this community. Sometimes a moss-covering 
of another composition is also met with, Rhacomitrium canescens 
often forms extensive mats on wet sandy ground. In North Ice- 
land Timmia austriaca and Philonotis tomentella are found plenti- 
fully in many places on the damp ground above the. Philonotis- 
belt, as a rule abundantly mixed with other species. Dicranella 
crispa is also very often found in great quantities on damp sandy 


THE BRYOPHYTA OF ICELAND 565 


ground. In North-west Iceland the development of the liverwort- 
mats was especially typical in many places (for instance in Gnups- 
dalur and near Isafjérdur); they consisted chiefly of Harpanthus 
Flotowianus, Alicularia geoscypha, A. scalaris, Cephalozia bicuspidata 
var. Lammersiana and Lophozia quinquedentata. Here Oligotrichum 
hercynicum also occurred abundantly in damp sandy localities. 

On a somewhat damp, cold flat at the bottom of the fjord 
Kaldalon, below the Jokull, grew Pohlia gracilis, Hypnum exannu- 
latum var. brachydictyon, Philonotis tomentella and Eucalyx subel- 
lipticus. 


The Bryophyte Vegetation near Hot Springs. 


Iceland abounds in hot springs: they are not, however, equally 
distributed over the whole country, but are most numerous in dis- 
tricts in which the most volcanic activity occurs. The majority 
of these hot springs are found in the south-eastern part of the 
country, on Reykjanes, around Reykjavik, in the south-western low 
land, and in several of the valleys of West Iceland, especially in 
Reykholtdalur. In North Iceland, the district around Myvatn is 
rich in hot springs, but they are also found scattered singly over 
the whole of this part of the country. They are absent from East 
Iceland, but a few occur in North-west Iceland, especially around 
Isafjardardjup. Groups of hot springs are found in several spots 
in the inner high land, but the Bryophyte vegetation around these 
is quite unknown. 

The majority of the hot springs of Iceland have a rather high 
temperature, the water being most frequently at about boiling point. 
Springs, the temperature of which is not much higher than that of 
their surroundings, are found for instance around Isafjardardjup 
near Laugarbol (507—55?) and Nauteyri (2537—30?), yet no charac- 
teristic vegetation had developed there. According to Thoroddsen 
such springs also occur north of Vatnajåkull, and have a tempera- 
ture which is not many degrees higher than their surroundings. 

The hot springs in Iceland may be classified as belonging to 
two different groups, viz. the Alkaline Springs and, the Sulphur 
Springs. 

The Alkaline Springs (thermæ) usually occur in districts where 
there has been no recent volcanic activity. They contain pure, 
clear water which often deposits siliceous sinter, and are always 
surrounded by a luxuriant vegetation. When the temperature reaches 





566 A. HESSELBO 


boiling point, they are called in Icelandic ”hver” (plur. ”"hverar”), 
if the temperature is lower, they are called aug” (plur. "augar”). 

The Sulphur Springs (Icelandic ””brennistenshverar”) are most 
numerous in districts where the effect of the volcanic activity is 
still felt, for instance near Myvatn and in many parts of South 
Iceland. The water contains sulphuretted hydrogen. These springs 
deposit sulphur, and the soil around them is greatly decomposed 
by the acid vapours which arise, and -is transformed to red or 
yellowish clay-masses. Sometimes, as for instance in Reykirdalur, 
they also deposit siliceous sinter. As a rule, a group of springs 
consists of either the one or the other kind of spring, occasionally, 
however, a few springs containing sulphuretted hydrogen may be 
found together with the alkaline springs. The vegetation around 
sulphur springs is always very scanty, since only a few species of 
plants can thrive were they are exposed to hot vapours containing 
sulphuretted hydrogen. 

The ""Laugar”” near Reykjavik. The vegetation has been described 
by-Oostenfeldi(borslidsrvols22 pp 235 ket sen) NEwhokhasknor 
however, mentioned the moss vegetation. When I visited the 
springs in 1909 and 1912 I found no characteristic moss vege- 
tation. Since the warm water had been used for washing purposes 
the ground surrounding the spring had been much trampled upon, 
and further downwards towards the warm outlet only the usual 
bog species were found. Here Philonotis fontana alone formed the 
elongated slender form known also from other hot springs. 

The Springs on Biskupstungur. On the low boggy tract of 
land between the rivers Bruarå and Tiångufljéåt in South Iceland 
(Biskupstungur, so called because the old episcopal residence — 
Skålholt — was situated there), there are, on both sides, along the 
rivers several hot springs of considerable size, or groups of hot 
springs of which Thorlåkshver and Laugaråshver in the southern- 
most part, near Skålholt, and Sydri Reykjahver, further northwards 
near the Bruarå, have been investigated. They all contain pure 
water free from sulphuretted hydrogen, with the exception of a few 
small holes near Thorlåkshver. 

Thorlåkshver (Fig. 21) is situated a few kilometres west 
of Skålholt, close to the east bank of the Bruarå. It consists of 
one larger and several smaller holes, from whence the boiling 
water flows out and gathers together into a rather large brook, 
which flows into the Bruarå. The vegetation along the sloping 


THE BRYOPHYTA OF ICELAND j 567 


banks of this brook and of the flat boggy surroundings (which are 
partly influenced by the heat of the soil, partly inundated by the 
hot water) is very luxuriant. Both around the basins of the springs 
and along the brook formed by the outlets of the springs, grew a 





Fig. 21. Thorlåkshver near Skålholt. In the background the river Bruarå; behind 


the fence to the left is a potato field, where the temperature of the ground was about 
252. The chief outlet of the spring is just behind the small elevation to the left, and 
the outlet-channel is from the right straight across the foreground. The” "peninsula”, 
in the middle of the figure, surrounded by the hot water, is partly covered with 
Sphagnum (the light patch) and partly with Hypnum-mats (the dark patch). 


broad belt of enormous yellowish brown and pale-green Søhagnum- 
cushions, 23 cm. deep, chiefly formed by $. imbricatum (yellowish 
brown), $. feres and S$. cymbifolium (whitish green) interwoven with 
Hypnum stramineum, Hylocomium squarrosum and here and there with 
Acrocladium cuspidatum, Hypnum imponens and H. Lindbergii. The 


568 A. HESSELBO 


surface of the Sphagnum cushions was usually covered with the 
orbicular, closely appressed leaves of Hydrocotyle vulgaris. The 
temperature in the Sphagnum cushions was rather high, on the 
surface of the ground as much as 50”, chiefly on account of their 
feeble heat-conducting power, and the tufts were quite saturated with 
the moisture from the warm vapours. The Sphagnum-belt did not 
extend so far as to the hot water, but everywhere there was a strip 
(10—20 cm. broad), nearest the water, covered with low dark-green 
mats of Fossombronia Dumortieri, occasionally intermixed with 
Haplozia crenulata or Alicularia scalaris. The same vegetation of 
Hepaticæ was in fact found in all places where the banks were too 
steep to allow Sphagna to gain foothold. The temperature under 
the Fossombronia-covering was, as a rule, about 40” (347—437). 
Where the bank was flatter and on the warm, dry, clayey flats in 
the neighbourhood of the springs Archidium phascoides grew in 
great masses, and often formed, without intermixture of other 
species, extensive, dark-green mats, 1—2 cm. deep. It occurred also 
on the siliceous sinter deposited by the water, and here and there 
among the Hepaticeæ. The temperature of the soil under the 
AÅrchidium-mat was almost everywhere about 40”. In the brook 
formed by the outlet of the spring there was a small island which 
was partially inundated by the hot water, and here grew a great 
many large cushions of a peculiar form, a Barbula fallax (var. læ- 
vifolia n. var.), and in one spot, on the slope, in the neighbourhood 
of the basin, grew Anthoceros punctatus together with Fossombronia. 

Outside the Sphagnum-belt the ground was perceptibly warm 
even for a tolerable distance, and the vegetation there was likewise 
greatly exposed to the warm aqueous vapours. There Hypnaceæ, in 
particular, grew very luxuriantly on the rather dry substratum. 
Hypnum imponens and H. Lindbergii were often almost the only 
species to be found there, the latter growing in a peculiar, erect 
form, almost branchless. The temperature of the ground under the 
Hypnum-covering was, as a rule, 257—30”. 

Å little below the spring the brook flowed along a steep bank 
about a metre high. Here the Sphagnum-belt disappeared and was 
replaced by a more mixed vegetation. Lowermost, at a height of 
about 10 cm. above the water, grew Fossombronia, Haplozia crenu- 
lata, Alicularia scalaris, and Anthoceros, then followed thick cushions 
of Hylocomium squarrosum, Hypnum stramineum, Thuidium delica- 
tulum, Sphagnum teres, S$. imbricatum and Catharinea undulata, 


THE BRYOPHYTA OF ICELAND 569 


moreover, Haplozia sphærocarpa woven into the Catharinea-cushions. 
All the mosses had developed to an unusual height and vigour, 
especially Catharinea which occurred in a peculiar form (f. thermo- 
phila) and formed large cushions more than 10 cm. deep. The same 
vigorous moss vegetation, associated with grasses and sedges, was 
also found on the flat above the steep bank, as far as the influence 
of the warm vapours was felt. 

Where the banks were low, so that the hot water partially 
flowed over and mixed with the cold water of the boggy ground, 
the usual bog or stream-bank vegetation was developed, which 
nearest to the outlet, in hot water of a temperature of about 25”— 
30”, consisted of a very slender form of Philonotis fontana, associa- 
ted with some Hypnum stramineum and Catoscopium nigritum. 

The Hypnum-belt mentioned above gradually merged into the 
usual grassland végetation; most of the species of which (for in- 
stance Rhacomitrium canescens, R. hypnoides, Philonotis, Mniobryum 
albicans and Hypnum uncinatum) were also found intermixed with 
the thermophilous vegetation proper. 

ÅA short distance from the main group of the hot springs, on a 
small, flat piece of ground near the river, there are some small 
holes, some of which emit boiling water, and some vapours im- 
pregnated with sulphuretted hydrogen. The ground around these 
holes consists of a rather dry, warm (30?7—407), clayey soil, with a 
very sparse vegetation. Archidium is common everywhere; Fossom- 
bronia often grows around the holes, and here and there on the 
clayey flats Riccia crystallina and Anthoceros punctatus. 

Laugaråshverir (Fig. 22) are situated in the southernmost part 
of Biskupstångur, near the farm Laugarås. At the foot of the slope 
on which the farm is situated, and in the boggy ground below, 
there issue several boiling hot springs which, through several 
channels, unite and form a large brook which flows into the Tun- 
gufljåt. The vegetation is very much like that in Thorlåkshver. 
Next to the water there is a belt of Fossombronia, Haplozia crenu- 
lata, ÅAlicularia and Archidium with a ground temperature of as much 
as 43”; then comes the Sphagnum-belt which is composed of $. sub- 
nitens var. coerulescens, S.teres, S. angustifolium, S$. cymbifolium, S. 
imbricatum and $. inundatum in association with Hypnum strami- 
neum. The warm ground between the channels was covered with 
extensive mats of Hypnum imponens, H. Lindbergii, H. stramineum, 
Hylocomium squarrosum, Climacium dendroides, Thuidium delicatu- 

The Botany of Iceland. Vol. I, part II. 37 


570 A. HESSELBO 


lum, Calypogeia Trichomanis and Scapania irrigua. Aulacomnium 
palustre was occasionally met with on wetter ground, and Archi- 
dium phascoides, Ditrichum homomallum and, on a single spot, 
Campylopus fragilis on rather dry, clayey soil. Under the Hyp- 








Fig. 22. Laugaråshverir. The springs are situated at the foot of a 


hill: along the outlet-channel and next to the water there is a narrow, 
dark belt of Hepatice and Archidium, then comes a broad belt of 
Sphagnum-cushions, and outside that (the dark patch in the fore- 


ground) are Hypnum-carpets. 


num-mats there was an almost constant temperature of 32", and 
under Archidium as much as 46”. On the slope above one of the 
largest springs Riccia sorocarpa grew in abundance, and where 
steep banks were formed along the outlets grew Hypnum, Thui- 


dium and Catharinea undulata f. thermophila; as was the case near 
Thorlåkshver. 


THE BRYOPHYTA OF ICELAND GVa | 


Further northwards near the Bruarå is Sydri Reykjahver. 
This consists of a large round basin from the centre of which the 
spring rises, bubbling up at short intervals and ejecting a consider- 
able amount of boiling hot water which flows over the edge of 
the basin and gathers together into a brook which flows into 
the river. 

On the somewhat higher and eastern side of the basin there is 
luxuriant grassland, while the lower western side bears a vigorous 
Bryophyte vegetation. Here, next to the water, grew extensive 
dark-green and reddish-brown mats of Alicularia scalaris, Haplozia 
crenulata and some Catoscopium nigritum in which bluish-green 
patches of Fossombronia occurred interspersed, and occasionally ro- 
settes of Anthoceros punctatus. Here the temperature of the ground 
was 257—27" just below the surface. Then followed a belt, about 
2 metres broad, of Sphagnum papillosum, S. cymbifolium and fruit- 
ing specimens of Polytrichum commune, with a ground temperature 
of about 20”. 

Lowermost on the steep bank (50—70 metres high) along the 
outlet grew the usual belt of Hepaticæ consisting of Haplozia, Ali- 
cularia and Anthoceros, then an abundance of Oligotrichum hercy- 
nicum interwoven with Haplozia sphærocarpa, and uppermost along 
the edge large cushions of Hypnum stramineum, Hylocomium squar- 
rosum and some Sphagnum cymbifolium, and here and there Caly- 
pogeia Trichomanis. 

In a small bog by the side of the basin, into which the hot 
water no doubt frequently flowed, grew, in addition to the usual 
bog vegetation, numerous large cushions of Sphagnum teres and 
S. inundatum. 

The Hot Springs in Reykholtdalur. Throughout the whole of 
Reykholtdalur, on both sides of the river Reykjadalså, there 
occur very many hot. springs, some of which discharge a great 
amount of water. Near the vicarage of Reykholt rise Dynkur, 
Skribla, Snorralaug and other springs, which in part contain 
sulphuretted hydrogen. Some hundred metres more to the south, 
on the other side of the river, is Hagindishver, and farther 
down the valley are other hot springs, of which the largest are 
Deildatunguhver, Snældubeinstadahver, Stårlurreykjahver, Klepp- 
jarnsreykir (probably Gråniund's Kleppholtsreykir) and Vellindis- 
hver; the last-mentioned is a fountain (geysir), which is situated 
on a small rock in the river, and is quite bare of vegetation. The 


54 le 


72 A. HESSELBO 


water in all these latter springs is pure and free from sulphuret- 
ted hydrogen. 

Near the springs around Reykholt the Bryophyte vegetation. is 
rather scanty, partly on account of the sulphuretted hydrogen, and 
partly on account of the constant disturbances caused by the 
traffic and the use of the hot water. Sphagnum is practically ab- 
seni. Along the outlets of the springs liverworts grew almost ex- 
clusively. Fossombronia and Blasia pusilla were very common, the 
latter grew in the form of flattened rosettes. Haplozia crenulata 
and Alicularia scalaris were also frequent. 

Åround all the other springs in Reykholtdalur there was a 
luxuriant vegetation of Sphagna. S. papillosum was especially do- 
minant and occurred in very large cushions, often 20—25 cm. deep, 
but besides this many other species were found either intermixed 
or separately in yellow, greenish, red or brownish cushions. The 
following species have been found on warm ground in Reykholt- 
dalur: Sphagnum papillosum, S. cymbifolium, S$. angustifolium, S. ru- 
bellum, S. Warnstorffii and S. teres were all common, S. subnitens 
(fr.), $. acutifolium, and S$. Dusenii var. falcatum occurred only now 
and then. 

The Sphagnum spp. grew partly as a belt along the outlets of 
the springs, partly on the boggy ground over which the hot water 
was flowing.  Intermixed with these grew everywhere Hypnum 
stramineum, and in many places Polytrichum commune with ripe 
capsules. Near Snældubeinstadahver Odontoschisma Sphagni 
was scantily woven into tufts of S$.rubellum. 

The warm clayey flats next to the water, and especially the 
steep banks stretching down towards the outlets, were covered with 
a dense carpet of mosses and liverworts, in which the latter were 
usually in the majority. The bluegreen rosettes of Anthoceros were 
found almost everywhere, usually together with Fossombronia, Blasia, 
Alicularia scalaris and Haplozia crenulata. In such places Archidium 
also often grew in great abundance. 

Gronlund records Catoscopium nigritum as occurring abundantly 
around hot springs in Reykholtdalur, but he has confused it with 
Archidium phascoides.  Catoscopium is found only exceptionally, 
and, as it were, accidentally on warm ground, sometimes in a 
somewhat divergent, slightly tomentose form (var. Grånlundii C. J.). 

On warm boggy ground the Sphagnum carpet was often greatly 
intermixed with, or partly replaced by, a Hypnum vegetation to- 


THE BRYOPHYTA OF ICELAND BLE. 


gether with some other Bryophyta. Hypnum revolvens, H. ex- 
annulatum, H. stramineum, H, stellatum, Acrocladium cuspidatum, 
Scapania irrigua, Pellia Neesiana and Alicularia scalaris were very 
common, Åneura multifida occurred here and there, especially in 
the Sphagnum rubellum cushions. Where the ground was less 
damp, especially on slightly inclined slopes stretching down towards 
the hot water, it was covered with a thick carpet of Hypnum im- 
ponens, H. molluscum, H. stramineum, H. revolvens, Hylocomium 
squarrosum, H. parietinum, H, proliferum and Camptothecium. lutes- 
cens, in which were scattered cushions of Sphagnum, especially S$. 
rubellum and S.teres. Sometimes Preissia commutata also could 
form bluish-green patches, about a metre in breadth, usually inter- 
mixed with some Hypnum molluscum. 

Sturlurreykjahver is situated on a slightly inclined slope 
stretching down towards the river Reykjadalså. At the top, near 
the spring, there is the usual Sphagnum vegetation which, farther 
downwards where the ground on both sides of the outlet of the 
spring is rather boggy, is replaced by a greatly mixed vegetation, 
composed principally of Hypnum imponens, Hylocomium squarrosum, 
Polytrichum gracile, Dicranum scoparium var. orthophyllum, Caly- 
pogeia Trichomanis and Alicularia scalaris. Alternating with the 
Hypnum carpet, extensive, dark-green mats of Archidium are found. 
The surface temperature of the ground under the Sphagnum carpet 
was, as a rule, 40”, whilst under the Hypnum-Archidium carpet 
there was an almost constant temperature of 37”. On stones in 
the tepid water, the temperature of which was at this spot 347, 
grew large cushions of Dicranella squarrosa. 

Kleppjarnsreykir is undoubtedly the same spring that 
Gronlund (1877, p. 350) calls Kleppholtsreykir. He states that at 
this spring the same two species of Søohagnum occur that are found 
near Tunguhver, and that near the spring and its outlet, among 
others, ""Distichium capillaceum, Mnium serratum, Hypnum ochraceum 
and Catoscopium nigritum” are found. The two last-mentioned have 
been wrongly determined, and the two first do not at any rate be- 
long to the species characteristic of the warm ground. 

The outlet has for a tolerably long distance steep banks, about 
half a metre high, which are, at the foot, covered with the usual 
liverwort-vegetation: Haplozia crenulata, Fossombronia, Anthoceros, 
Ålicularia and Archidium. The uppermost part is covered with a 
dense moss-carpet consisting of Hylocomium squarrosum, H. proli- 


574 A. HESSELBO 


ferum, Hypnum imponens, Mnium hornum, M. undulatum, M. serra- 
tum, Fissidens osmundoides and Enthostodon ericetorum. On a stone 
partly inundated by the hot water, which had a temperature of 
abon 0se Fissidens osmundoides, Blindia acuta, Dicranella squar- 
rosa, Enthostodon ericetorum, Anthoceros punctatus, Aneura multifida 
and Scapania irrigua grew intermixed. 





Fig. 23. Deildatunguhver. To the left is the slope with the hot springs. The part 
in the middle is saturated with the hot water and covered with Sphagnum papillo- 
sum, Polytrichum commune and Hydrocotyle. 

Deildatiunguhver (Gronlund's Tiånguhver) (Fig. 23) is 
situated north of the river and opposite to Kleppjarnsreykir. It is 
mentioned by Groånlund (1877, p. 349), who states that the surface 
temperature of the ground at the top of the hill was 23” R, and 
that there Polytrichum commune, Sphagnum cymbifolium, S$. cuspida- 
tum and a Campylopus (C. flexuosus) -— not determined with cer- 
tainty — grew in the warm ground. Deildattnguhver is one of 
the hot springs of Iceland which discharge the greatest amount 
of water, and consists of a row of holes at the foot of a hill 
6—8 metres high, whence an enormous quantity of water and 
steam is ejected with a great noise. The masses of steam, 
which can be seen for miles, enshroud the surroundings of the 
spring with so dense a cloud that it is hardly possible to ap- 
proach without getting soaked through. Below the hill the water 


THE BRYOPHYTA OF ICELAND Bo 


gathers into a broad brook which flows into the river. The whole 
hill, which consists of reddish clay, is warm throughout, and at 
the top the temperature was 367—40” just below the surface of the 
ground. 

On the slope stretching down towards the springs Blasia pu- 
silla grew abundantly, occasionally in company with Fossombronia 
Dumortieri. The dry ground on the top of the hill was covered 
with large patches of Campylopus flexuosus and Hypnum imponens 
intermixed with scattered tufts of Catharinea undulata and Oligo- 
trichum hercynicam and entirely interwoven with Gymnocolea in- 
flata. At the foot of the hill, but on the opposite side to that on 
which the springs lie and where the ground was damper, grew, in 
addition to the above-mentioned species, Sphagnum cymbifolium 
and Polytrichum gracile. Here some 5—6 small plants of the fern 
Blechnum spicant were also found. 

The warm brook below the hill divides into several branches, 
which flow round some large islands saturating them with the 
warm water. Here Sphagnum papillosum, S$. angustifolium and Po- 
lytrichum commune grew in enormous cushions, which were every- 
where covered by the orbicular leaves of Hydrocotyle vulgaris and 
entirely enshrouded in hot vapours. 

Geysir. On a low mound about one kilometre long and 
about ”/> a kilometre broad, situated about 100 metres above sea- 
level, there are a great number of hot springs (about 40) besides 
the well-known Great Geysir. The majority of these springs con- 
sist of a larger or smaller hole, but sometimes of a basin, several 
metres in diameter, in which the boiling water usually reaches to 
the upper edge and flows over the somewhat raised rim, or is 
ejected at shorter or longer intervals. The soil consists of reddish 
clay, or nearest to the springs — especially to Geysir — of sili- 
ceous sinter. The water in the majority of the springs smells 
slightly of sulphuretted hydrogen. The whole district around the 
springs is warm, ånd in many places the temperature is so high 
that the tenants of the neighbouring farm (Laug) bake their bread 
by putting the dough into a pot and burying it about 30—40 cm. 
below the surface of the ground. 

The siliceous sinters are quite bare of vegetation. The warm 
clay-flats situated on a higher level are, as a rule, dry, and either 
bear a greatly scattered Bryophyte vegetation or are quite bare. Un- 
der the scattered, low cushions of Ditrichum homomallum, Archi- 


576 A. HESSELBO 


dium and Fossombronia which were growing here, the thermometer 
showed as high a temperature as 34” on the surface of the ground. 
On a slightly damp flat which was covered with a low-growing, 
dense form of Climacium dendroides, the surface temperature of the 
ground was 43”. 

In the more low-lying tracts, where the ground was rather 
damp, or partially inundated with hot water, there had, in several 
places, developed a luxuriant vegetation of phanerogams and moss- 
es, which was, however, often much trodden down. Here, in the 
boggy parts, grew a low vegetation of Carex-Viola palustris, greatly 
mixed with Bryophyta, of which the following species, which -grew 
intermixed, were observed: Hypnum stramineum, H. sarmentosum, 
H. imponens, Hylocomium squarrosum, Sphagnum cymbifolium, Ca- 
tharinea undulata, Calypogeia Trichomanis, ÅAlicularia scalaris, Ha- 
plozia crenulata, Scapania irrigua, Åneura pinguis and an undeter- 
minable Pohlia. On damp clayey flats Archidium, Fossombronia 
Dumortieri, Haplozia crenulata and Ålicularia scalaris formed low, 
dense mats in which Anthoceros punctatus occasionally occurred in 
small rosettes. 

Below Great Geysir, Hypnum chrysophyllum grew on a clayey 
flat inundated with tepid water. 

Laugarvatnshverir. By the shore of lake Laugarvatn, below 
the farm of the same name and along the water's edge, there is 
a row of springs containing boiling water saturated with sulphur- 
etted hydrogen. The surrounding ground which consists of black 
basalt gravel is, owing to its ex- 
posed situation, quite bare of Bryo- 
phyte vegetation; and of higher 
plants only a few specimens of 
Polygonum Persicaria were found. 
By the northernmost of the springs 
only, which is situated at some dis- 
tance from the lake shore, a warm, 
damp clayey flat was covered with 
a carpet of Sphagnum cymbifolium, 
S. Girgensohnii, Polytrichum- com- a a 
mune, Hylocomium squarrosum, Hyp- 
num stramineum, Archidium phas- 








Fig. 24. Bryum sp. a, Older leaves deve- 
loped before the plant was inundated with 
coides, Pohlia grandiflora % Åong- not water; b, leaves from shoot developed 


sltræmia longipes, Gymnocolea inflata in hot water (Geysir). 


-= 


THE BRYOPHYTA OF ICELAND 571 


c. coles., Haplozia sphærocarpa, H. crenulata, Alicularia scalaris and 
Scapania irrigua. 

At Englandshverir, some small hot springs which rise in 
Lundurreykjadalur, there was only opportunity for quite a short 
stay on the journey past. Here, on the warm ground along the 
outlet of the springs the following species were observed: Sphag- 
num teres, Hypnum imponens, Catharinea undulata, Fissidens os- 
mundoides, Enthostodon ericetorum, Catoscopium nigritum, Årchidium 
phascoides, AÅneura pinguis, Pellia Neesiana and Fossombronia Du- 
mortieri. 

The Hot Springs near Reykir. In the south-western part of 
Iceland, around the farms Reykir and Reykirfoss, about 45 kilo- 
metres south-east of Reykjavik, there is a very large group of hot 
springs which stretches towards the north and north-west, through 
the valleys, right up upon Hengill to a height of 400—500 metres 
above sea-level. To this group must also be reckoned the hot springs 
near Kolvidarhol and on Hellisheidi. The majority of the springs 
are situated around the farm Reykirfoss, and north of the latter, 
on both sides of the river from a height of about 50 to 90 metres 
above sea-level; most of them are in the neighbourhood of the river, 
and are in part situated directly upon the banks of the latter, several, 
however, lie upon the slope of Reykjafjall, up to about 100 metres 
above sea-level. Where the naked hasalt does not protrude, the 
ground around the springs consists of greyish-white siliceous sinter 
almost bare of vegetation, and large tracts of it are warm and, so 
to speak, perforated like a sieve with numerous hot springs of all 
possible sizes. These appear in the form of (1) steaming holes 
(fumaroles) which range in size from quite fine pores to rather large 
funnels, (2) fountains (geysirs) which at short intervals eject boiling 
water ranging from quite a slender jet one foot in height to mighty 
columns of roaring water from 10 to 15 feet in height (Little Geysir), 
(3) mud-pools with bubbling, bluish-grey mud which is often ejected 
to a distance far beyond the surroundings of the pool, and (4) 
boiling basins or cauldrons which may measure 6—8 metres in 
diameter, and either have a quiet surface or boil up at short inter- 
vals, so that the water partially flows over the edge of the basin. 
The steam is everywhere impregnated with sulphureited hydrogen, 
with the result that the vegetation is very poor and homogeneous. 
The luxuriant Sphagnum-vegetation known from the alkaline springs 
does not occur near these springs. Around these steaming holes, 


578 A. HESSELBO 


geysirs and mud pools the ground is usually quite bare as far as 
the boiling water and the mud are able to spurt. 

Near the boiling basins which occur in great number and gene- 
rally have a diameter of 2—4 metres, the ground forms, as a rule, 
a slope on the one side with an angle of 257—40f, whilst on the 
opposite side the water flows down a gentle slope which stretches 
as far as to the river. The ground for the first 10—20 cm. of this 
slope, which corresponds approximately to the different water-levels, 
is of a greyish-white colour and quite bare of vegetation. Then 
comes a belt of varying breadth formed of liverworts which, next 
to the water, are black or of a brownish black, above this of a 
reddish-brown or brownish-green, and at the top, green. This liver- 
wort-carpet was everywhere composed of Gymnocolea inflata, Haplo- 
zia crenulata, occasionally also Alicularia scalaris f. rufescens and at 
the top Cephalozia bicuspidata. The temperature under the moss- 
covering was on an average 207—235”. 

Polytrichum commune grew scattered in the liverwort carpet, 
from which its bluish-green tops protruded several centimetres, but frequ- 
entlyitalso formed large growths abovethe liverworts, partly interwoven 
with the latter. Only very few phanerogams were able to grow 
there, usually only flowerless specimens of Viola palustris, the short- 
stemmed leaves of which rested on the liverwort carpet. Besides 
the above-mentioned Bryophyta several others were found on the 
warm clayey flats, although far more scantily. Anthoceros puncta- 
tus, Fossombronia Dumortieri, Blasia pusilla and Archidium phascoides 
grew here and there. In several places Oligotrichum hercynicum 
occurred in tufts a few mm. high, and in one single locality Alicu- 
laria geoscypha and Aongstræmia longipes were found in company 
with it. On warm ground between loose blocks of rock, at an alti- 
tude of about 260 metres, the following were found among others: 
Polytrichum commune, Hylocomium loreum, Plagiothecium elegans 
and Diplophyllum albicans. 

In the neighbourhood of the farm Reykirfoss, on the western 
side of the river, and at the edge of the lava field, there is a rather 
large spring with pure water of a temperature of about 37”. Here 
the blocks of stone in the water were quite covered with Pellia 
Neesiana, in the tufts of which Veronica Anagallis had taken root. 

Where the boiling water flowed over the slightly inclined slopes, 
it deposited greyish-white siliceous sinter, upon which grew blue- 
green algæ as soon as the water had cooled down to 607—70". The 


THE BRYOPHYTA OF ICELAND 579 


Bryophyta appeared first at the edge of the outlet of the spring where 
the temperature was 25?—40"”. The vegetation here resembled that 
of wet, cold ground, and was usually composed of Philonotis fon- 
tana, Mnium Seligeri, Hypnum stramineum, Catoscopium nigritum, 
and in a single spot of some Sphagnum angustifolium. On drier, 
warmer ground Hypnum imponens was found here and there in 
company with Hypnum stramineum, Hylocomium squarrosum, Årchi- 
dium phascoides, etc. 

Of the hot springs situated at higher levels only a few have 
been investigated. In the neighbourhood of Kolvidarhol, at an 
altitude of about 350 metres, on the warm ground around some 
holes whence issued vapours impregnated with sulphuretted hy- 
drogen and where sulphur had been deposited, the following species 
were growing: Gymnocolea inflata, Haplozia crenulata, Fossombronia 
Dumortieri, Åneura pinguis, Archidium phascoides and Polytrichum 
commune. Here Gymnocolea inflata was not strictly confined to the 
warm ground, but appeared rather to prefer clay, as it also grew 
on clayey flats which had probably been warm previously, but were 
now quite cold. 

The same species together with Blasia pusilla grew also in Rey- 
kjadalur (at an altitude of about 260 metres) on the damp hase of 
a warm slope from which vapours, containing sulphuretted hydrogen, 
and water issued sparsely from numerous small holes. Near some 
mud-pools on Hellisheidi only Gymnocolea inflata and Haplozia 
crenulata occurred. 

The Sulphur Springs on Reykjanes — which I myself had no 
opportunity of visiting — have been described by Ostenfeld (Bot. 
Tids., vol: 22, p. 239). They occur in two groups, near Reykjanes 
lighthouse and also near Krisuvik. 

The Solfataras near Reykjanes lighthouse form a large 
group with numerous orifices of discharge, partly mud-pools, partly 
steaming holes, everywhere around which the ground is warm 
(207—309). Near "the outlet of the largest mud-pool, Gunna, the 
damp ground, which had a temperature of about 30”, was covered 
with Haplozia crenulata to which, along the margin, came several 
other species, viz. Fossombronia Dumortieri, Archidium phascoides 
(recorded by Ostenfeld as Pohlia nutans v. filicaulis), Trichostomum 
littorale and Bryum ventricosum, all of which, however, only occur- 
red sparingly. Around the steaming holes grew Riccia bifurca, 
Preissia commutata, Fossombronia Dumortieri, Archidium phascoides, 


580 A. HESSELBO 


Bryum ventricosum, Fissidens osmundoides and Philonotis fontana. 

The Sulphur Springs near Krisuvik are, according to 
Ostenfeld, rather bare of vegetation, and he mentions only Haplozia 
crenulata as growing near a few of these. The collections contain 
also Gymnocolea inflata and Polytrichum commune which were 
gathered here by Steenstrup, so probably the same species occur 
on this spot as are, for instance, to be found near Kolvidarhol. 

In the remaining part of Iceland the vegetation of the warm 
ground has hardly been investigated at all. During my stay at 
Myvatn in 1909 continual rain and fog prevented me from making 
a closer investigation of the numerous sulphur springs which occur 
there. The warm clayey flats were mostly quite bare, only around 
a single steaming hole, where the temperature of the surface was 
about 40”, grew Riccia bifurca, Haplozia crenulata and Fossombronia 
Dumortieri. 

Å moss-cushion from one of the Solfataras near Kverkfjall, 
(Eyafjallajokul) collected by Andr. Lundager, consisted of Philo- 
notis fontana and Riccia bifurca. 

By comparing what is known regarding the moss vegetation of 
warm ground, it is seen that the composition of the vegetation is 
dependent on the chemical nature of both the water and the es- 
caping vapours, and also on conditions pertaining to soil. 

Where the water is free from sulphuretted hydrogen, or where it, 
at most, contains a trace of it, the Sphagnum vegetation is very 
luxuriant, especially when the surroundings of the spring are boggy, 
as is the case in Biskupstångur and near most of the springs in 
Reykholtdalur. The condition for the formation of a continuous 
Sphagnetum, in addition to a warm, damp substratum, is an abun- 
dant development of vapour, which envelops the moss-cushions in 
a damp, warm atmosphere. The most frequent species is S$. papil- 
losum, which often constitutes the bulk of the Sphagnum carpet, in 
association with S$. cymbifolium, S. angustifolium and S. teres, fre- 
quently replaced by, or intermixed in patches with, some other 
species. Scattered in the Sphagnum carpet grew, as a rule, a very 
vigorous, fruiting form of Polytrichum commune, and usually also 
Hypnum stramineum, Hylocomium squarrosum or Acrocladium cuspi- 
datum. 

Outside the Sphagnum-belt there occurs, on warm, boggy ground, 
a moss carpet interspersed with Careæ spp. and other flowering plants, 
and chiefly composed of Hypnaceæ, occasionally also of Philonotis 


THE BRYOPHYTA OF ICELAND 581 


fontana, Aulacomnium palustre or of an abundance of species inter- 
mixed. The commonest of these species are: Hypnum imponens 
and H. Lindbergii, which are often the most abundant constituents 
of the carpet, H. stramineum, Hylocomium squarrosum, Acrocladium 
cuspidatum, Thuidium delicatulum, and numerous other bog-mosses 
such as Hypnum revolvens, H. molluscum, Catoscopium nigritum, 
Fissidens osmundoides, Scapania irrigua, Pellia Neesiana, Aneura 
pinguis, etc. Especially the first six species, however, occur every- 
where in abundance, and often attain an unusual size and development. 

Ås an intermixture in this moss carpet some species, for in- 
stance Mnium hornum, Catharinea undulata and Polytrichum gracile, 
are occasionally met with which partly occur also in other localities 
and are partly characteristic of warm ground, and in Iceland only 
occur in such. To the latter group belong — in addition to Hyp- 
num imponens, which is rare elsewhere — Enthostodon ericetorum, 
Calypogeia Trichomanis, Aneura multifida (found scantily in one spot 
on cold boggy ground) and Odontoschisma Sphagni. Some species 
develop special warm-soil forms: Catharinea undulata f. thermophila 
is exceedingly vigorous, as much as 10 cm. high, with long, ex- 
tremely wavy, secund leaves right up from the base. Philonotis fon- 
tana becomes thread-shaped in warm ground, with long, branchless, 
barely tomentose shoots (Fig. 8), Hypnum Lindbergii develops a similar 
form with almost branchless, erect stems with scattered leaves, and 
Hypnum imponens is also, as a rule, much elongated and slightly 
branched. Comp. further Bryum sp. (Fig. 24). On drier, clayey 
ground the Hypnum carpet is replaced by other species. Archidium 
phascoides is extremely common, and often forms extensive growths 
both near alkaline springs and near springs containing sulphuretted 
hydrogen. Here Campylopus flexuosus, C. fragilis, Oligotrichum hercy- 
nicum, Catharinea undulata, Gymnocolea inflata and several other 
species are also to be found. 

In damp clayey soil, especially along the ouilets of the hot 
springs, both where the water is pure and where it contains sul- 
phuretted hydrogen in small quantities, without however depositing 
sulphur, a low Bryophyte carpet occurs composed of specie—smostly 
Hepaticæ — the majority of which are only met with in warm soil. 
Haplozia crenulata, Fossombronia Dumortieri and Oligotrichum hercy- 
nicum are also common. Preissia commutata is common in Reyk- 
holtdalur, Alicularia geoscypha and Aongstræmia longipes and a few 
other species occur here and there. 


582 A. HESSELBO 


Where the surroundings of the spring consist of siliceous sinter, 
as is the case with the majority of the springs of the Geysir-com- 
plex, and in Reykirdalur, all species of Sphagnum are absent, or 
are to be found only as a subordinate component, and the moss 
vegetation is, on the whole, scanty, and consists of the same species 
as are to be found on clayey soil, together with some Hypnaceæ, 
Philonotis, etc. on damper soil. 

Near the Sulphur Springs proper the Bryophyte vegetation 
is extremely scanty, and the ground is often quite bare of vegeta- 
tion as far as the hot vapours extend. Sphagnum and Hypnaceæ 
are entirely absent, whereas the majority of the Hepaticæ are to 
be found, especially Haplozia crenulata, Fossombronia Dumortieri, 
Gymnocolea inflata, Alicularia scalaris, Preissia commutata together 
with Archidium phascoides, Polytrichum commune and a few other 
mosses. Riccia spp. (bifurca, sorocarpa and crystallina) belong spe- 
cially to the dry, warm ground in the neighbourhood of sulphur 
springs. 


The following species have been found on warm ground: 

1. Species which occur only on warm ground. 

Riccia crystallina L. Thorlåkshver. 

Riccia sorocarpa Bisch. Laugaråshverir. 

—  bifurca (Hoffm.) Lindenb. Myvatn; Reykjanes: Kverkfjåll. 

Anthoceros punctatus L. Common. 

Fossombronia Dumortieri (Huben.) Lindb. Everywhere. 

Calypogeia Trichomanis (L). S. F. Gray. Myvatn; Reykholtdalur; 
Englandshverir; Sydri Reykjahver; Geysir. 

Odontoschisma Sphagni (Dicks.) Dum. Reykholtdalur. 

Sphagnum imbricatum Hornsch. Thorlåkshver; Laugaråhverir. 

Sphagnum cymbifolium (Ehrh.) Hedw. Common. 

Sphagnum Dusenii C. Jens. v. falcatum Reykholtdalur. 

Archidium phascoides Brid. Very common everywhere. 

Entosthodon ericetorum (Bals. et de Not.) C. M. Reykholtdalur ; 
Englandshverir. 

Campylopus flexuosus (L.) Brid. Reykholtdalur. 

Campylopus fragilis (Dicks.) Br. eur. Laugaråshverir. 

Barbula fallax Hedw. var lævifolia n. var. Thorlåkshver. 

Catharinea undulata (L.) Ehrh. f. thermophila Thorlåkshver ; 
Laugaråshverir. 

2. Species which are mainly distributed over warm 


THE BRYOPHYTA OF ICELAND 583 


ground and only occur asan exceptionin other localities. 

AÅneura multifida (L.) Dum. Reykholtdalur. 

Haplozia crenulata (Sm.) Dum. Near all hot springs. 

Gymnocolea inflata (Huds.) Dum. Reykholtdalur; Reykirdalur; 
Krisuvik. 

Sphagnum papillosum Lindb. Very common. 

S. angustifolium C. Jens. Common near alkaline springs. 

S. inundatum Russ. Laugaråshverir; Sydri Reykjahver. 

Hypnum imponens Hedw. Common near most hot springs. 

Haplozia sphærocarpa (Hook.) Dum. and Oligotrichum hercynicum. 
These two species have a most peculiar distribution. 

Haplozia has been found near several hot springs in SW. Ice- 
land and also on mountain heights near Isafjérdur. 

Oligotrichum hercynicum is common near most alkaline springs, 
and also near a few containing sulphuretted hydrogen. It is, 
moreover, common on rocky flats at an altitude of 400—500 metres, 
and also in NW. Iceland; but is otherwise very rare. 

sæ Speceres whrchareralso common inother lolealitres: 

Åneura pinguis Dum. Common. 

Pellia Neesiana (Gottsche) Limpr. Common. 

Blasia pusilla L. Near many springs. 

Preissia commutata (L.) N. v. Es. Reykholtdalur; Reykjanes. 

Alicularia scalaris (Schrad.) Corda. Very common everywhere. 

— geoscypha de Not. Reykirdalur. 

Cephalozin bicuspidata (L.) Dum. Reykirdalur. 

Scapania irrigua (Nees) Dum. Frequent. 

Sphagnum subnitens Russ. et Warnst. Laugaråshverir; Reyk- 
holtdalur. i 

S. teres (Schimp.) Ångstr. Very common. 

S. rubellum Wils. Reykholtdalur. 

S. Warnstorffii Russ. Reykholtdalur. 

S. Girgensohnii Russ. Laugarvatnshverir. 

Dicranella squarrosa (Schrad.) Sch. Reykholtdalur. 

Fissidens osmundoides (Sw.) Hedw. Frequent near alkaline springs. 

Blindia acuta (Huds.) Br. eur. Reykholtdalur. 

Ditrichum homomallum (Hedw.) Hampe. Geysir; Reykirdalur. 

Trichostomum littorale Mitten. Reykjanes. 

Catoscopium nigritum (Hedw.) Brid. Frequent. 

Mnium undulatum (L.) Weiss. Reykholtdalur. 

Mnium hornum L. Common near alkaline springs. 


584 A. HESSELBO 


Mnium serratum Schrad. Reykholtdalur. 
Aulacomnium palustre (L.) Schwågr. Common, several places in 
abundance. 
Philonotis fontana (L.) Brid. Very common, often in masses. 
Catharinea undulata (L.) W.et M. Common. 
Polytrichum commune L. Common. 
— gracile Dicks. Reykholtdalur. 
Thuidium delicatulum (L.) Mitten. Frequent. 
Hypnum molluscum Hedw. Abundant near all the hot springs 
in Reykholtdalur. 
Hypnum Lindbergii (Lindb.) Mitten. Common everywhere. 
Hypnum revolvens Sw. Common near many springs, especially 
in Reykholtdalur. 
Hypnum chrysophyllum Brid. Geysir. 
— … sarmentosum Whilb. Geysir. 
—  stramineum Dicks. Very common everywhere. 
—  exannulatum Gumb. Reykholtdalur. 
Acrocladium cuspidatum (L.) Lindb. Common. 
Climacium dendroides (L.) W. & M. In many places, plentiful near 
Geysir. 
Hylocomium squarrosum (L.) Br. eur. Common everywhere. 
— proliferum (L.) Lindb. Common in Reykholtdalur. 
— parietinum (L.) Lindb. In several places plentiful. 


Besides the species mentioned in the above list, several other 
species are, however, also found as more casual intermixtures in 
the moss carpet, especially species from boggy soil and from grass- 
land, for instance Rhacomitrium canescens, R. hypnoides and Hypnum 
uncinatum. 


Mesophilous Bryophyte Formations. 


The Bryophyta, as a rule, play a less prominent part in the 
plant-covering of soil of a middle degree of dampness, than in that 
of wet soil, because in the former the higher plants find more 
favourable conditions of life, and the Bryophyta are consequently 
driven more into the background and occur only as ground vege- 
tation between the higher plants. Several other conditions are, how- 
ever, of essential importance in this connection, for instance con- 
ditions pertaining to the light, the direction of the slope and the 
snow-covering. Bryophyta bear shade better than do the higher 


THE BRYOPHYTA OF ICELAND 585 


plants and consequently develop better than the latter in loca- 
lities which are less favourable as regards the light, for instance, 
on the northern sides of clefts and in caves. In valleys southern 
slopes are covered with grass or other herbaceous plants, while 
northern slopes are moss-grown. In the narrow tuff-clefis of South 
Iceland the sides are often covered with a luxuriant moss carpet 
consisting of Hylocomium spp., Thuidium, Polytrichum, etc. - Here 
the snow-covering no doubt also plays an important part, since by 
remaining until far into the spring, it checks the growth of the 
higher plants far more than it does that of the mosses. 

Grassland is mostly knolly, and the mosses grow abun- 
dantly on the ground between the grasses. Here it is especially 
species such as Hypnum uncinatum, Hylocomium squarrosum, Cli- 
macium dendroides, Polytrichum alpinum and Rhacomitrium canes- 
cens which are the most abundant constituenis of the vegeta- 
tion, but in addition to ihese many other species occur, for 
instance Catharinea undulata, Timmia austriaca, Tortella tortuosa, 
Dicranum congestum, Ditrichum flexicaule, Distichium montanum, 
Bartramia ityphylla, Rhacomitrium hypnoides, Hylocomium spp., 
Hypnum Lindbergii and Lophozia quinquedentata. The composi- 
tion of the vegetation varies according to the degree of dampness, 
therefore, we sometimes find species from boggy soil and sometimes 
xerophilous species of heathy soil intermixed in greater and smaller 
quantities. 

Grass-slopenandkHerb-slope Ek Op the herb=slopesthe 
Bryophyta are very scantily represented. On the grass-slope the 
bottom is covered with a more or less dense carpet of mosses and 
liverwort& in which the chief species are Hylocomium proliferum, 
H. parietinum, H. squarrosum, H. rugosum, H. loreum, H. triquetrum, 
Hypnum uncinatum, Climacium dendroides, Camptlothecium lutescens, 
Thuidium delicatulum (S. Icel.), Mnium affine, Timmia austriaca, Po- 
lytrichum alpinum, Pogonatum urnigerum, Rhacomitrium canescens, 
R. hypnoides, Ditrichum flexicaule, Lophozia lycopodioides and Pla- 
giochila asplenioides, but besides these, many other species occur as 
a more casual admixture. 

On a stony substratum with a thin layer of soil mosses are 
dominant, and phanerogams grow scattered in the moss carpet. 
A "moss-slope” is then developed which, as a rule, is composed of 
the same species as those found on the grass-slope, most frequently 


1 See footnote in Thoroddsen”s Physical Geography of Iceland, ante p. 330. 
The Botany of Iceland. Vol, I, part II. 38 


586 A. HESSELBO 


Hypnum uncinatum, Hylocomium spp., Rhacomitrium spp. and Thu- 
idium delicatulum (S. Icel.). These are often associated with several 
of the xerophilous species, such as Dicranum scoparium, Frullania 
Tamarisci, Ptilidium ciliare and Antitrichia curtipendula, through which 
are formed transitional stages to the moss-heaths mentioned below. 
In South Iceland several of the southern species have their greatest 
distribution on well-sheltered, grass-covered slopes with a southern 
exposure, for instance Scleropodium purum, Eurhynchium piliferum, 
E. Stockesii, Thuidium tamariscinum, T. Philiberti, T. delicatulum and 
Mnium undulatum. 

Birchcoppicesterontontafrathetdsyæsubstratummeke 
wood-floor is either occupied by grassland or by heather moor, 
and the Bryophytle vegetation does not differ in any respect from 
that which occurs on a corresponding substratum outside the 
coppices. Most frequently it consists of Hylocomium spp. in asso- 
ciation with Hypnum uncinatum and Rhacomitrium spp. (R. canes- 
cens and R. hypnoides). 

Bryophyla grow only rarely and scantily on tree trunks. Ra- 
dula complanata and Frullania dilatata have been found in one 
single spot on a tree trunk, but, as a rule, it is some ground species 
or other which grows a little way up on the slanting trunks. In 
some few places Rhacomitrium fasciculare has been found on tree 
trunks to which it had spread from neighbouring stones. 

Laugardalsskågur. This coppice, which is open and of 
about a man's height, has its floor mostly covered with heather 
moor, with a thick moss carpet composed of Hylocomium parieti- 
num, H. proliferum, H. triquetrum, H. squarrosum, Hypnum uncina- 
tum and Rhacomitrium hypnoides, occasionally Dicranum scoparium, 
Ptilidium ciliare, Lophozia lycopodioides and L. quinquedentata also 
occurred. Of quite a similar composition was the moss carpet in 
almost all the coppices of South and West Iceland. A few other 
species also belong to this community, but are more rarely met 
with, for instance Bryum cæspiticium, Pohlia nutans, P. acuminata 
and Lophozia barbata. 

Hallormstadaskågur (East Iceland). The floor here rather 
varies in nalure, and is sometimes covered with heather moor and 
sometimes with grassland or peat. In heathy soil Hylocomium spp. 
were dominant in connection with Rhacomitrium hypnoides, while 
especially Hypnum uncinatum grew abundantly on grassy ground. 
Where it was very shady Timmia austriaca grew abundantly on knolls. 


THE BRYOPHYTA OF ICELAND 387 


Of other species the following were observed: Pohlia nutans, P. 
cruda, Bryum cæspiticium, B. affine, Bartramia ilyphylla, Polytrichum 
alpinum, P. juniperinum, Distichium montanum, Ditrichum flexicaule, 
Sælania cæsia, Tortula subulata, T. ruralis, Dicranum scoparium, 
Ptilidium ciliare, Lophozia lycopodioides, L. quinquedentata, Frullania 
Tamarisci and Scapania curta. 

Birch coppice in Kaldalon (NW. Iceland). The south-west- 
ern slope was covered with a low-growing and very dense birch 
coppice with a ground vegetation of numerous flowering plants 
(Geranium silvaticum, Bartsia alpina, Taraxacum, etc.) and a Bryo- 
phyte carpet of Hylocomium proliferum, H. parietinum, H. squarrosum 
and Hypnum uncinatum intermixed with Dicranum scoparium, D. 
fuscescens, Pogonatum urnigerum, Polytrichum alpinum, Lophozia ly- 
copodioides, L. quinquedentata, L. Flærckei, L. ventricosa, Cephalozia 
bicuspidata and C. pleniceps. In open spots Pohlia nutans was very 
common, and Polytrichum gracile occurred frequently on knolls on 
the floor of the coppice. 

Of a similar composition is the moss carpet of all the coppices 
of NW. Iceland. In several spots Dicranum majus and Hylocomium 
triquetrum also occurred. 

BryophytanofsSoilfrecently laid. bare. On slopes, on 
the sides of clefts, on soil-covered rock-ledges and, on the whole, 
in all places where by weathering, by land-slips or by the action 
of water new soil is produced, which has not yet become inhabited 
by higher plants, a luxuriant Bryophyte vegetation occurs, which 
C. Jensen (1910, p. 258) includes in a group within the mesophi- 
lous associations, under the name "”Moosvereine des frischen Erd- 
bodens”. A number of species are here met with, especially of the 
commonly occurring mesophilous forms, but also, several species 
belonging to damp sandy soil and to rocks. It is especially liver- 
worts and the acrocarpous mosses which are represented here, 
while the pleurocarpous mosses occur but scantily. It is a con- 
dition for the growth of mosses that there is an abundant snow- 
covering during winter, which affords them protection from de- 
siccation by wind. 

The composition of the moss-covering varies greatly according 
to the prevailing degrce of humidity, and the local conditions. 

In such localities as those mentioned above the following 
Hepaticæ occur: — ÅAneura pinguis, Marchantia polymorpha, Gym- 
nomitrium concinnatum, Alicularia scalaris, A.geoscypha, Lophozia 

387 


588 A. HESSELBO 


quinquedentala, L. ventricosa, L. Miilleri, L. alpestris, Cephalozia bicus- 
pidata, Cephaloziella Hampeana, Blepharostoma trichophyllum, An- 
thelia julacea, Å.Juratzkana, Scapania subalpina, S$. curta, more 
rarely Preissia commutala, Blasia pusilla, Eucalyx subellipticus, Lo- 
phozia excisa, L. heterocolpos and Lophocolea minor. The following 
are (he commonest mosses: — Dichodontium pellucidum, Dicranella 
crispa, Distichium montanum, Didymodon rubellus, Tortula subulata, 
Desmatodon latifolius, Tortella tortuosa, T. fragilis, Schistidium apo- 
carpum, S. gracile, Encalypta ciliata, E.rhabdocarpa, the majority of 
the species of Bryum, Leptobryum pyriforme, Pohlia commutata, P. cruda, 
Mniobryum albicans, Mnium punctatum, Bartramia ityphylla, Philo- 
notis tomentella, Timmia austriaca, Pogonatum urnigerum, Myurella 
julacea, M. tenerrima, Brachythecium reflexum, Plagiothecium Roese- 
anum, Hypnum hamulosum, H. callichroum, H. uncinatum, Hyloco- 
mium loreum. Some rare species, for instance Eurhynchium stri- 
gosum var. præcox, Heterocladium squarrosulum, Distichium inclina- 
tum and Weisia viridula are found especially in such localities as 
those mentioned above. 


Xerophilous Bryophyte Associations. 
Heaths. 

The heath formation is extensively distributed about Iceland. 
It occurs especially on dry slopes, and hardly ascends higher than 
about 300—-400 metres above sea-level, where it is succeeded by the 
rocky-flat association. 

Heather-moor (H. Jønsson, 1900, p. 69). The chief heath- 
forming plants are Empetrum nigrum, Vaccinium uliginosum, V. 
Myrtillus, Arctostaphylos uva ursi, Betula nana and, especially in 
E. Iceland, Calluna vulgaris. Mosses and liverworts often grow 
abundantly on the ground. The Bryophyte carpet is chiefly com- 
posed of Hylocomium spp. (especially H. proliferum and H. parieti- 
num), Rhacomitrium hypnoides, R. canescens and Hypnum uncinatum, 
but Dicranum spp. (D. scoparium, D. fuscescens and others), Lopho- 
zia spp. Ptilidium ciliare, Frullania Tamarisci and several other 
species often occur in abundance. As examples will best show 
the composition of this vegetation, the following are given: — 

In North-west Iceland heath-vegetation is extensively distributed, 
especially around Dfrafjérdur and Isafjérdur. Here the heath-for- 
ming plants are mostly Vaccinium Myrtillus, V. uliginosum, Empe- 
trum nigrum and occasionally Betula nana. 


THE BRYOPHYTA OF ICELAND 589 


Onkbeatbyksroyndken GrupdalurmearDyrafjordurta 
rather scattered Bryophyte vegetation was found, composed of Hylo- 
comium proliferum, H. parietinum, Hypnum uncinalum, Dicranum 
scoparium, D. molle, D. fuscescens, D. Starckei, Conostomum boreale, 
Lophozia Flærckei, L. lycopodioides, L. Kunzeana, L. ventricosa and 


- 





Fig. 25. Grunnavik near Jåkulsfiårdur in NW. Iceland. Stony slope. The low-lying parts 
are boggy, the somewhat higher ridges are covered wilh heath vegetation (Empetrum, 
Vaccinium, Dicranum spp., etc.), whilst the stony ridges situated highest (in the back-- 


ground) are bare of vegetation. 


L. alpestris.. The Dicranum spp. formed thick, extensive cushions, 
and the Lophozia spp. grew woven into these cushions. 

Grunnavik near Jåkulsfjårdur (Fig. 25). Up to a height of 
100—200 metres ihe mountain slopes were partly covered with 
heather-moor and partly wilh bog-vegetation or moss-bogs. The 
stony ridges, situated higher up, which had been formed hy stones 
sliding down from the mountain above, were covered with heath, 
while the intervening depressions were boggy, and there along the 
channels, where the water formed streamlets or issued as springs, 
the usual "Dy” vegetation consisting of Mniobryum albicans, Philo- 
notis, etc., had developed. 


590 A. HESSELBO 


The Bryophyte carpet of the heath was composed of Hyloco- 
mium parietinum, H. proliferum, Hypnum uncinatum, Rhacomitrium 
hypnoides, Dicranum scoparium, D. fuscescens, D. molle, Plilidium 
ciliare, Lophozia Flærckei, L. lycopodioides, L. ventricosa, Cephalozia 
bicuspidata, C. pleniceps and scattered cushions of Sphagnum Gir- 








Fig. 26... Knolly moor of Betula nana intermixed with Salix lanata and Salix phylicifolia. 
Ground vegetation: Empetrum, Arctostaphylos uva ursi, Anthoxanthum odoratum and 


Polygonum viviparum. 


gensohnii. The Dicranum spp. especially D. scoparium often formed 
continuous carpets, which appears to correspond with the "”Dicra- 
numfheathssædeseribedshyd Jensens op 262: 

Knolly Betula nana Heath near Ljosavatn (Fig. 26). In 
addition to Betula nana, which formed the bulk of the vegetation, 
there occurred scattered shrubs of Salix phylicifolia, S. glauca and 
numerous grasses and flowering plants. Mosses and liverworts 
grew abundantly on the knolls and on the ground between them. 
The following species were observed: Hypnum uncinatum, Rhaco- 
mitrium canescens, R. hypnoides (all in abundance), Hylocomium pro- 
liferum, H. rugosum, H.squarrosum, Camptothecium nitens, Heterocla- 
dium squarrosulum, Brachythecium erythrorrhizon, Polytrichum. juni- 
perinum, Timmia austriaca, Conostomum boreale, Pohlia cruda, Bar- 
lramia ityphylla, Tortella tortuosa, Dicranum congestum, Ditrichum 


THE BRYOPHYTA OF ICELAND 591 


flexicaule, Lophozia lycopodioides, L. Kunzeana and Plagiochila as- 
plenioides. 

The Moss Heath. On dry stony tracts there often occurs a 
continuous growth of mosses, in which higher plants grow scat- 
tered without being able to give character to the vegetation. In 
heather-moors mosses often form a continuous carpet, and then all 
transitions between typical heather-moor and moss-heaths are found. 
By the decay of the mosses a humus-layer is gradually formed, 
and the moss-heath is thus the pioneer of other associations such 
as heather-moors, coppices and grassland (H. Jonsson, 1905, p. 41). 

The moss heath occurs most frequently as a Rhacomitrium- 
heath (Grimmia-heath, H. J., 1895, p. 70; 1900, pp. 68 and 85; 1905, 
p. 40). It is found in the low land — where it is typically developed, 
especially in the lava fields, which are often entirely covered 
by it — as well as on mountain slopes, and in the Alpine region 
as far upwards as 600—700 metres above sea-level (Fig. 27). It also 
occurs in patches in other formations, for instance in grassland, 
where it is usually developed on the top of the knolls.  Rhacomi- 
trium hypnoides is the dominant specics, and is often the only one 
found over considerable tracts; it forms a uniformly grey carpet, 
as much as a foot deep. There is only a very small intermixture of 
other Bryophyta. The most frequent åre Rhacomitrium canescens, 
Hylocomium proliferum, H. parietinum, Dicranum scoparium, Pfili- 
dium ciliare, Frullania Tamarisci, rarer are Lophozia lycopodioides, 
L. barbata, and other species. The Rhacomitrium heath is extensively 
distributed in many places on the taluses of fallen blocks ånd dé- 
bris (Urd), and there — in case conditions are favourable — by 
the mosses decaying and. forming humus, it quickly passes into other 
associations, especially into grass-slopes and herb-slopes. 

The Rhacomitrium canescens heath is developed on a somewhat 
damper substratum than is the R. hypnoides heath, and is met with, 
for instance, in patches in depressions in the latter, where it is 
immediately distinguished by its light greyish-green colour, which 
differs distinectly from the whitish-grey colour of its surroundings. 
Here it must frequently share the locality with Hypnum uncinatum 
and is often greally intermixed with other Bryophyta, for instance 
Polytrichum alpinum, Hylocomium spp., Lophozia lycopodioides and 
L. quinquedentata, and several other species. Ås an example of 
such a Rhacomitrium canescens heath may be mentioned a slightly 
inclined gravelly flat near Holt in South Iceland, about 400 me- 


Å. HESSELBO 


592 





THE BRYOPHYTA OF ICELAND 593 


tres above sea-level, which was covered with a continuous carpet 
of R. canescens, in which Hylocomium parietinum, Dicranum Starckei, 
Conostomum boreale, Lophozia alpestris and L. lycopodioides occurred 
scantily interspersed (Fig. 28). 





Fig. 28... Rhacomitrium canescens heath near Holt in South Iceland. In the background 


is seen one of the deep clefts with vertical sides, so common in this district. 


Near Hof in SE. Iceland there was a horizontal, somewhat 
damp gravelly flat covered wilh a continuous moss-carpet, of which 
the chief species were Rhacomitrium canescens and Hylocomium pro- 
liferum, abundantly mixed with Hylocomium parietinum, H. rugosum, 
H. squarrosum and Hypnum uncinatum. Spread about in the moss 
carpet there grew Dicranum scoparium, D. congestum, Ditrichum 
flexicaule, Tortella tortuosa, Polytrichum juniperinum, P. alpinum 
and Aulacomnium turgidum. Conostomum boreale was common and 
occurred in small, compact tufts, usually interwoven with Lophozia 
ventricosa, Gymnomitrium concinnatum, Anthelia and species of 
Cephalozia, and on prominent knolls of earth Diphyscium. sessile 
in association with Gymnomitfrium concinnalum formed compact 
blackish-brown coverings... Aulacomnium turgidum occurred abund- 
antly in several spots, more or less intermixed with Hylocomium 
squarrosum, Hypnum uncinatam and Rhacomitrium canescens, and 
formed carpets. Spread about in the moss carpet grew numerous 


294 A. HESSELBO 


Juncus and Carex tufts, and this formation must most nearly be 
termed a transitional form between moss-heath and meadow-land. 
On the whole, Rhacomitrium canescens thrives best in Iceland on 
somewhat damp ground, under quite similar conditions as Aula- 
comnium turgidum. 

The Mixed Moss-heath (Hylocomium-Rhacomitrium heath) 
occurs especially on slightly inclined gravelly flats, under similar 
conditions as the heather moor, and is, as a rule, a pioneer for 
the latter. It may, however, also develop into a grass or herb 
slope, where the ground is wetter. The chief species of this moss 
carpet are Rhacomitrium canescens, R. hypnoides and species of 
Hylocomium, but as a rule many other species are found intermixed 
with these. 

In a moss-heath near Moådruvellir (Esja), on a slightly inclined 
gravelly flat, at an altitude of about 100 metres, the chief species 
of the Bryophyte carpet were Hylocomium proliferum, Rhacomitrium 
hypnoides, R. canescens and Hypnum uncinatum; more scantily oc- 
curred: Hylocomium loreum, H. squarrosum, Climacium dendroides, 
Antitrichia curtipendula, Brachythecium albicans, Polytrichum alpi- 
num, P, urnigerum, Timmia austriaca, Pohlia cruda, Ditrichum flexi- 
caule, Bartramia ilyphylla, Distichium montanum, Ptilidium ciliare, 
Frullania Tamarisci, Lophozia lycopodioides, L. quinquedentata and 
L. quadriloba. Scattered in the Bryophyte carpet grew a few flower- 
ing plants such as Anthoxanthum and Galium verum, and this 
moss-heath will undoubtedly develop into grassland. 

On a flat interspersed with stones at an altitude of about 
400 metres the Bryophyte carpet was composed of Rhacomitrium 
hypnoides abundantly mixed with R. canescens, Hylocomium parie- 
tinum, Hypnum uncinatum and Dicranum Starckei. As an inter- 
mixture in the Bryophyte carpet occurred: Timmia austriaca, Pohlia 
commutlata, Distichium montanum, Lophozia quinquedentata, L. qua- 
driloba, L. alpestris, Plagiochila asplenioides, Blepharostoma tricho- 
phyllum, Anthelia Juratzkana, Alicularia scalaris, Cephalozia pleni- 
ceps and C. bicuspidata. Flowering plants were almost entirely 
absent. 

On prominent knolls of earth occurring on heathland and 
grassland, on dry ledges and slopes, a very peculiar Bryphyte 
covering is often found: low, very compact cushions of Diphyscium 
sessile in connection with several liverworts, especially Gymnomi- 
triam concinnatum, are found covering the soil with a very dense 


THE BRYOPHYTA OF ICELAND 595 


brownish-black carpet. Conostomum boreale also — usually inter- 
woven with liverworts such as Gymnomitrium concinnatum, Lopho- 
zia ventricosa, Pleuroclada albescens and species of Cephalozia — 
forms exceedingly compact cushions on dry, stony ground. Both 
these species grow in particularly exposed localities, where the 
snow-covering is absent during winter, but their compact tufts, 
densely covered with tomentum to an unusual degree, protect them 
from being destroyed by strong winds. Diphyscium forms quite 
low (only one cm. high), but rather wide tufts, while Conostomum 
forms tufts a few cm. in diameter, but often more than 10 cm. in 
height. In the latter case the basal porlion of the tuft is securely 
wedged into the gravelly substratum and attached to the stones 
with numerous rhizoids. Diphyscium is found at all levels, while 
Conostomum is most widely distributed on the rocky flats, but in 
the low land occurs only somewhat scantily in the heath formation. 


The Bryophyte Vegetation of the Rocks. 


The rock vegetation includes formations of very various com- 
position, with extremely different demands as regards light, mois- 
ture, etc. The substratum may consist of solid rock, for in- 
stance rock-faces, flats or large blocks, or it may consist of de- 
tached stones of various sizes, as on gravelly and stony slopes and on 
taluses. The distribution of the water is very unequal, since a 
quite dry part with decidedly xerophilous species may be met with 
next to a part where the water flows over the rock, and provides 
conditions for the growth of hygrophilous species. 

The chemical and physical nature of the substratum exercises 
great influence upon ihe composition of the Bryophyte covering. 
On tuff and breccia the vegetation is far more luxuriant than on 
hasalt, and the reason for this is partly because the uneven sur- 
face of the tuff affords better habitats for the plants, partly because 
the water penetrates into the tuff comparatively casily, and after- 
wards reappears through the fissures and porose parts of the rock- 
surface so that these are, as a rule, rather damp. The greater or 
smaller amount of limc present is also of importance for the fre- 
quency of many species. The direction of the exposure and the 
conditions pertaining to shelter play a great part with regard to 
the Bryophyle vegetalion. Species growing on exposed rock-faces 
and flats must be able to endure complete desiccation by the sun 
during the summer, and by strong winds during the winter, while 


596 AÅ. HESSELBO 


the Bryophytes of the clefts and Urds live in a comparatively hu- 
mid atmosphere and are protected by the snow-covering during the 
winter. The conditions pertaining to light are also of importance 
with regard to the composition of the Bryophyte vegetation. On 
the ground between the stones of the talus, grow other species than 
those to be found upon the stones. Caves and dark clefts contain 
species which belong specially to these localities, and the vegeta- 
tion on vertical surfaces or slopes with a southern exposure is 
different to that found on the same kind of ground with a northern 
exposure. 

Owing to the extremely different conditions to which plants 
are exposed when growing on rocks, all kinds of associations are 
met with there, from decidedly xerophilous to hygrophilous, light- 
forms, and species which thrive best in dark caves. The rocky 
substratum may be compared to a mosaic pavement, in which 
each parlicular piece of homogeneous conditions is exceedingly 
small. 

The mosses grow partly on the rock itself (lithophytes) and 
parlly in soil which has accumulated in fissures and depressions, 
or on ledges (chomophytes). The boundary line between these two 
groups has, however, been effaced to a great extent as many of 
the lithophytes proper also occasionally grow on gravel or even 
on soil, and it is not always possible to decide whether a moss- 
cushion has originally developed on humus-covered rock, or has 
itself formed humus by a process of decay. Especially on mountain 
heights the two groups merge one into the other, as species such 
as Andreæa petrophila, Dicranoweisia crispula, Rhacomitrium. fas- 
ciculare and R. sudeticum, which in low-lying regions are decidedly 
lithophytes, descend to the ground on mountain heights and often 
form extensive growlhs on rocky flats. The following species may 
be referred to the lithophytes proper (those marked ” are common): 


Species belonging to Coastal Rocks. 


”Ulota phyllantha ”Schistidium maritimum 


Water Lithophytes. 


”Schistidium rivulare =Fontinalis longifolia 
£Fontinalis antipyretica =Rhynchostegium rusciforme 
= androgyna ”"Hypnum ochraceum 
thulensis = alpinum 


islandica = alpestre 


THE BRYOPHYTA OF ICELAND 597 


Theremalnins Lithophytes: 


=Metzgeria furcata ӯRhacomitrium fasciculare 
£Radula complanata É = aciculare 
Gymnomitrium coralloides Hedwigia albicans 
=Madotheca Cordæana fOrthotrichum rupestre 
Frullania fragilifolia anomalum 
=Andreæa petrophila = saxatile 
=Anæctangium compactum == cupulatum 
Weisia crispata = Sturmii 
=Dicranoweisia crispula — Killiasii 
=Blindia acuta = Blyttii 
Bryoxiphium norvegicum = lævigatum 
Tortula obtusifolia Leucodon sciuroides 

— muralis Neckera complanata 
Schistidium confertum Leskea nervosa 
Grimmia commutata =Pterigynandrum filiforme 

= alpestris Lescuræa patens 

= Doniana Isothecium myurum 

== ovata = tenuinerve 

= incurva =Homalothecium sericeum 

patens Thamnium alopecurum 

— funalis Hypnum Bambersgeri 

— torquata Er evolntum 
”Rhacomitrium sudeticum 5 = cupressiforme 
v — heterostichum 


In addition to the species mentioned above there are a great 
number of other species which are of equal frequency on bare 
rocks and on humus-covered rocks. To this group belong for in- 
stance Haplozia atrovirens, Diplophyllum albicans, Amphidium Mou- 
geotti, Hymenostylium curvirostre, Schistidium spp. and Hypnum 
hamulosum. 

To the chomophytes may be reckoned a numerous group 
of species which partly contain the chomophytes proper, i. e. species 
which properly speaking belong to humus-covered rock-ledges or to 
clefts, and partly a great number of species which are as frequently 
met with in other formations. The following species may be re- 
garded as belonging to the chomophytes proper: — 


Reboulia hemisphærica £Haplozia pumila 
Sauteria alpina Lophozia heterocolpos 
Fimbriaria pilosa Tortula mucronifolia 
Fegatella conica Z£Encalypta rhabdocarpa 
Preissia commutata g = ciliata 
Haplozia riparia £Amphidium lapponicum 


= atrovirens Hymenostylium curvirostre 


598 A. HESSELBO 


Sælania cæsia "Orthothecium chryseum 
Anomobryum filiforme = intricatum 
=Plagiobryum Zierii Eurhynchium strigosum 
Bryum æneum == Swartzii 

— pallescens =Plagiothecium pulchellum 

capillare = depressum 

=Mnium orthorrhynchum = Ræseanum 

== serratum =Amblystegium Sprucei 

= stellare Hypnum hamulosum 


Among the species met. with here, which however cannot be 
reckoned to the chomophytes proper, the following are the most 
frequent: Bartramia ityphylla, Pohlia cruda, Bryum inclinatum, B. 
ventricosum, ØOncophorus virens, O. Wahlenbergii, Didymodon ru- 
bellus, Distichium montanum, Schistidium apocarpum, S$. gracile, Di- 
trichum flexicaule, Philonotis tomentella, Marchantia polymorpha, 
Lophozia spp., Scapania curta, S$. subalpina and Blepharostoma tricho- 
phyllum. 

Stony and Gravelly Slopes. The Bryophyte vegetation 
is, as a rule, rather poor in species, and not very characteristic. 
Rhacomitrium heaths are common where the slope is not too steep, 
but stony and gravelly slopes are often almost bare of vegetation, 
only here and there occur some cushions of Dicranoweisia crispula 
and species of Rhacomitrium. 

A Stony Slope near Grunnavik in NW. Icéland consisted 
of large blocks at the foot, and in the upper and steeper part of 
smaller stones. Vegetation was absent at the top, but increased in 
abundance lower down as the ground gradually become more level, 
and the conditions thereby became more stable. Scaltered among 
the stones grew plants of the rocky flat, for instance Salix herba- 
cea, Papaver radicatum., Oxyria digyna, Sibbaldia procumbens and in 
the shelter afforded by larger stones Athyrium alpestre, Alchemilla 
alpina, etc. Upon the stones grew Rhacomitrium hypnoides, R. fas- 
ciculare and Dicranoweisia crispula and on the ground between the 
stones Brachythecium reflexum, Hypnum uncinatum, Lescuræa pa- 
tens, L. filamentosa, Hylocomium squarrosum, Polytrichum alpinum, 
Dicranum fuscescens, Lophozia ventricosa, L. lycopodioides and Anthelia 
Julacea. 

Stony Slope near Berufjoårdur. On the steep somewhat 
damp slope there grew Rhacomitrium fasciculare, Schistidium gracile, 
Dicranoweisia crispula, Lescuræa decipiens, Heterocladium squarrosu- 
lum and Timmia austriaca. 


THE BRYOPHYTA OF ICELAND 599 


HaulhrsForsraltentblockstandtdebrrse(U nd) ERE con 
ditions for the development of planis is far more favourable here 
than on the stony slope, which is constantly exposed to disturbances 
through the down-sliding of stones and earth. The most xerophi- 
lous species, viz. species of Rhacomitrium and Grimmia, Dicrano- 
weisia crispula, Orthotrichum rupestre and Hypnum revolutum grow 
on the stones, while on the ground between the stones there are 
favourable conditions for a number of species which have greater 
requirements with regard to shelter, moisture and shade. The Urd 
is therefore very rich .in species, since lithophilous species are 
found there on the blocks, and species, which grow on the ground, 
occur between the stones, and Bryophytes belonging to heathland 
and grassland, are found on the soil-covered stones and ledges. 

In order to show how heterogeneous the composition of the 
Bryophyte vegetation may be, the Urd in Heljusdalur on Vest- 
mannaey may serve as an example. The valley is almost semicir- 
cular in shape, and open towards the south, and in consequence of 
this favourable situation combined with the comparatively mild, 
damp climate of the island, a considerable number of southern 
species are found there, while species such as Ulota maritima and 
Schistidium maritimum betoken the proximity of the sea. The rocks 
around the valley are inhabited by numerous sea-fowls, and they 
often rest on the blocks of the Urd, and on these spots, manured 
by the birds, it is species of Bryum (B. argenteum and B. capillare) 
which particularly make their appearance. 

The fallem blocks are everywhere covered with cushions of 
mosses and liverworts. The commonest species are Schistidium 
apoøcarpum, Ulola maritima, Hypnum cupressiforme, H. uncinatum, 
Orthotrichum rupestre and Ceratodon purpureus.. The following 
species were also observed on the blocks: Radula complanata, Ma- 
dotheca Cordæana, Plagiochila asplenioides, Cephaloziella Hampeana, 
Didymodon rubellus, ”Barbula cylindrica, Tortella tortuosa, T. fragilis, 
Tortula subulata, ET. mucronifolia, Distichium montanum, FOrthotri- 
chum anomalum, Encalypla. ciliata, E. rhabdocarpa, Pohlia cruda, 
Bryum capillare, B. elegans, B. inclinatum, B. æneum, Mnium serra- 
tum, M. stellare, M. orthorrhynchum, Myurella julacea, Orthothecium 
intricatum, ”Isothecuum myurum, Homalothecium sericeum, Amblyste- 
gium Sprucei, ZA. serpens, Hypnum hamulosum and H. revolutum. In 
the caves formed among the stones, into which the light penetrated 
but scantily, the walls were covered with light-green shining mats 


600 A. HESSELBO 


of =Plagiothecium depressum, the ground between the blocks was 
covered with a thick, dark-green carpet of Hypnum uncinatum, 
Hylocomium squarrosum, Brachythecium salebrosum, ”Eurhynchium 
Stockesii, Plagiothecium Roeseanum, ”Thuidium tamariscinum, T. 
abietinum, Mnium undulatum, Ditrichum flexicaule, Dichodontium 
pellucidum, =Lophocolea cuspidata, "L. minor and Lophozia quinque- 
dentata. On the soil-covered grassy ledges and slopes, between the 
stones, grew Brachythecium albicans, B.salebrosum (in masses), Cli- 
macium dendroides (in abundance), Camptothecium lutescens, Hyloco- 
mium proliferum, H. parietinum (sparingly) and Mnium affine (in 
abundance). 

All the species marked ” are southern forms the majority of 
which have only been found in South Iceland. On the whole, the 
Urd affords favourable habitats for many species which do not 
thrive in more exposed localities. 

On a dry Urd consisting of large blocks of basalt, near'Holt, 
in South Iceland, the following species were growing: — Rhacomi- 
trium fasciculare, R. heterostichum, R. hypnoides, Grimmia Doniana, 
Schistidium apocarpum, Andreæa petrophila, Antitrichia curtipendula, 
Hypnum uncinatum, Hylocomium loreum, H. proliferum, H. parieti- 
num and Frullania Tamarisci. 

Almost the same species are found on larger detached blocks. 
On blocks of lava round about Reykjavik there grew particularly 
Rhacomitrium heterostichum, R. fasciculare, Andreæa petrophila, Schi- 
stidium apocarpum, Grimmia Doniana and Dicranoweisia crispula. 

On larger stones in the home-field (Tunet) near Vallanes 
(East Iceland) there grew Pterigynandrum filiforme, Hypnum revolu- 
tum, Tortula ruralis and Encalypta rhabdocarpa. Some large blåcks 
near Hafursholt in South Iceland were quite overgrown with 
Grimmia alpestris. On some large blocks of basalt near Seljaland 
there grew Grimmia Doniana, Rhacomitrium fasciculare, R. hypnoides, 
R. sudeticum, Dicranoweisia crispula and Gymnomitrium coralloides. 

The Vertical Rock-belts. Owing to the division into layers 
peculiar to the basalt, the mountain sides consist of numerous 
vertical scarps of varying height, alternating with slopes formed by 
fallen blocks and debris. Ås a rule, the first 100—200 metres of the 
mountain form a slope which is only slightly inclined at the bottom 
and becomes steeper higher up, and where only here and there 
small portions of the primitive rock protrude. This slope is limited 
by a precipitous rock-face of very varying height, sometimes ris- 


THE BRYOPHYTA OF ICELAND 601 


ing to a height of several hundred metres, right up to the plateau 
on the mountain top, but usually divided into numerous steps, 
consisting of shelves or ledges of varying breadth. The rock-face 
is here and there intersected by deep fissures and clefts, excavated 
by the downward-flowing water, which partly drains away in 





Fig. 29. Face of basalt rock (E. Iceland). Everywhere in the crevices small, black, moss- 


cushions (Andreæa, Dicranoweisia crispula and Grimmia spp.). 


these channels, partly spreads over the ledges and from thence 
flows down the rock-faces or penetrates into the crevices of the 
rock, and then reappears once more further down. Consequently, 
the conditions pertaining to moisture vary greatly, and therefore 
the moss-covering also. As some examples will best show the com- 
position of the latter, the following are given: — 


1. Esja, low damp rock-face just above the slope: 
The Botany of Iceland. Vol, I, part. II. 39 


602 A. HESSELBO 


Philonotis tomentella, Mnium punctaltum, M. orthorrhynchum, Amphi- 
dium Mougeottii, A. lapponicum, AÅnæctangium compactum, Didymodon 
rubellus and Schistidium gracile grew abundantly in extensive cushions. 
Lescuræa filamentosa was found abundantly at the basal part of 
the rock-face. Pohlia cruda, Bartramia ityphylla, Plagiothecium pul- 
chellum, Myurella julacea, M. tenerrima, Radula complanata, Haplozia 
atrovirens and Marchantia polymorpha grew partly in small tufts, 
partly intermixed with other specics in the tufts formed by them. 

27 BotnsdalurtHvalfjordurtdampirockskacerlle base 
of the rock-face, up to a height of about a few metres, was covered 
with extensive cushions of Hypnum cupressiforme, H. molluscum, 
Brachythecium glareosum, Orthothecium chryseum, Philonotis tomen- 
tella and Distichium montanum. Where the rock-face was wet with 
the downward-oozing water, large patches of it were covered with 
Anomobryum concinnum. Partly in the crevices of the rock-face, 
partly scattered in the tufts there occurred Amblystegium Sprucei, 
Bryum pallescens, B. inclinatum, Lophozia quinquedentata, L. Miilleri, 
Blepharostoma trichophyllam and Aneura pinguis. 

3. Berufjårdur, rock-face, where the water was oozing 
down from the ledge above: Hypnum alpestre, Orthothecium 
chryseum, Blindia acuta, Onchophorus Wahlenbergii, Plagiopus C-deri, 
Philonotis fontana, Distichium montanum, Myurella julacea, M. tener- 
rima, Scapania subalpina, Lophozia quinquedentata, Haplozia atrovi- 
rens and H. cordifolia. 

AE Semdrskord ure dryrole kt hacerwehaknorbrernkere 
posure. On the dry rock face itself there grew: Prerigynandrum 
filiforme, Homalothecium sericeum, Grimmia funalis, G. torquata, 
Hypnum revolutum and Ortholtrichum rupestre. The basal part of 
the rock-face was much weathered, presenting numerous crevices 
and small ledges. There grew: Hypnum revolutum, H. filicinum, 
Orthothecium chryseum, Plagiothecium pulchellum, P. Roeseanum, 
Myurella julacea, M. tenerrima, Amphidium Mougeottii, Mnium orthor- 
rhynchum, Bartramia ityphylla, Pohlia cruda, Plagiobryum Zierii, 
Meesea trichoides, Orthotrichum Blyttii, Distichium montanum, Di- 
trichum  flexicaule, Fissidens osmundoides, Schistidium  confertum, 
Didymodon rubellus, Metzgeria furcata, Blepharostoma trichophyllum 
and  Odontoschisma elongatum. 

SEA kureyriedrytr ok facerness metres ablosveris es 
level. The rock-face was covered with extensive mats of Homalo- 
thecium sericeum and Plerigynandrum filiforme mixed with cus- 


THE BRYOPHYTA OF ICELAND 603 


hions of Grimmia funalis, Hypnum revolutum, Rhacomitrium. hete- 
rostichum and Orthotrichum rupestre. On the basal and somewhat 
damper portion of the rock-face there grew Gymnostomum rupestre, 
Philonotis tomentella, Orthothecium chryseum, Hypnum uncinatum, 
Distichium montanum, Ditrichum flexicaule, Amphidium lapponicum, 
Pohlia cruda, Bartramia ityphylla, Ceratodon purpureus, Tortella tor- 





Fig. 30.  Grimmia torquata on one of the sides of Flokastadagil. 


tuosa, Lophozia quinquedentata, L. quadriloba and everywhere on 
the ledges large cushions of fruiting Tortula ruralis. 

The two last quoted examples are typical of the vegetation of 
dry rock-faces. Homalothecium sericeum occurs there almost always, 
and is often dominant on exposed rock-faces as far upwards as 
400—500 metres above sea-level, Pterigynandrum is also extremely 
common, . especially in the low land. Grimmia funalis is rarely ab-… 
sent and often forms very large and deep, fruiting tufts, and espe- 
cially in South Iceland G. torquata is extremely common and often 
covers large portions of the rock-faces with its irregularly-rounded 
cushions (Fig. 30). 

6.- Low, much weathered and fissured basalt rocks 
near Hof in SE Tceland. -On the: top of the rocks.it was"espe- 


397 


604 A. HESSELBO 


cially Rhacomitrium hypnoides which grew abundantly on débris pro- 
duced by disintegration. On the rocks there grew Hypnum cupressi- 
forme, H. revolutum, H. imponens, Pterigynandrum filiforme, Leskea ner- 
vosa, Rhacomitrium fasciculare, R. heterostichum, Grimmia funalis, G. Do- 
niana, Schistidium apocarpum, S$. confertaum, Dicranoweisia crispula, 
Andreæa petrophila, Orthotrichum rupestre, O. Sturmii, Gymnomitrium 
coralloides, Frullania Tamarisci, Radula complanata, Metzgeria furcata 
and at the base of the rock, Madotheca Cordæana. Å southern rock- 
face was covered with Leucodon sciuroides var. morensis. In clefts 
and crevices and on soil-covered ledges there grew Mnium orthor- 
rhynchum, M. stellare, Pohlia cruda, Encalypta ciliata, E. rhabdo- 
carpa, Bartramia ityphylla, Dicranum Andersonii, Distichinm monta- 
num, Ditrichum flexicaule, Didymodon rubellus, Amphidium lapponi- 
cum, Plagiochila asplenioides, Marsupella Funckii and Lophozia al- 
pestris. 

The locality just described comes nearest to that known in 
Iceland as "Holt”, i. e. low, stony ridges or protruding masses of 
rock in the low land. The vegetation upon these ""Holts” is gene- 
rally xerophilous in character and is, as a rule, not so rich in 
species as is the vegetation mentioned above. The ridges are usu- 
ally more or less covered with soil which bears heath-vegetation, or 
are occupied by gravelly flats, therefore the Bryophyte vegetation 
consists chiefly of species belonging to heathland or grassland (Hy- 
locomium spp., Polytrichum alpinum, P.juniperinum, P. pilosum, Hyp- 
num uncinalum, Frullania Tamarisci, Rhacomitrium spp., etc.), while 
directly upon the rocks there grow species such as Hypnum cu- 
pressiforme, H. hamulosum, H. revolutum, Orthotrichum and Grimmia 
spp., and in the crevices Bartramia, Pohlia cruda, Amphidium lap- 
ponicum, Mnium orthorrhynchum, Myurella, Distichium montanum 
and other of the species found in rock-clefts in greater or smaller 
quantities. 


The Bryophyte Vegetation of the Tuff Rocks. 


The reason why the Bryophyte vegetation of the tuff rocks is 
here treated in a separate section is that the life-conditions of the 
plants are essentially different on a basalt and on a tuff substratum. 
Tuff consists of consolidated fine-grained material (volcanic ash and 
dust) through which are scattered larger and smaller blocks of 
rock. While the basalt has a smooth surface, with fissures and de- 
pressions only here and there, the surface of the tuff is uneven and 


THE BRYOPHYTA OF ICELAND 605 


rough, and therefore offers more favourable conditions to the mosses 
for attaching themselves to it. When a basalt and a tuff surface 
occur "side by side, as is often. the -case, the difference is very 
obvious. Mosses grow more scantily on basalt, since it is distinctly 
seen that the majority of the tufts are situated in, or proceed from, 





Fig. 31. Tuff cleft (Bleiksågil) near Barkarstadr. 


a fissure, while tuff surfaces are often covered with a continuous 
moss-carpet, composed of many different species. It is also a fact 
of great importance that the water which flows down from above, 
and atmospheric humidity, are available to the mosses of basalt 
surfaees only as long as the water is flowing or is deposited, while 
the tuff is able to absorb water through its porose surface, and thus 
has a constant supply of necessary moisture for the vegetation. 
Owing to the slight power of resistance of the tuff against the ero- 


606 A. HESSELBO 








Fig. 32. Side of tuff-cleft near Barkarstadr. At the foot of the cleft, among others 
Mnium undulatum grew abundantly. On the sides there grew Hypnum filicinum (at the 
top, to the right), Fegatella conica (in the middle and to the left), Philonotis fontana, 


Mniobryum albicans and many other mosses. 


sive action of waler, many more clefts and caves with damp sur- 
faces are met with here than on the basalt, and these provide ideal 
habitats for a great number of mosses. 


Tuff deposits of volcanic origin are found in many places in 


THE BRYOPHYTA OF ICELAND 607 


Iceland, but the reason why in the following only the Bryophyte 
vegetation of South Iceland is mentioned, is because it is only this 
part of Iceland that has been somewhat thoroughly investigated. 

Fljétshlid. North of the great river Markarfljåt, the country 
rises to a height of 200—600 metres. The slope stretching down- 
wards to the Markarfljøt is trenched by a great number of smaller 
streams, many of which flow at the bottom of deep, narrow clefts 
in the tufflayers. 

One of the longest and deepest clefts is Bleiksågil near Bar- 
karstadr (Fig. 31). This cleft has almost. everywhere vertical or 
overhanging sides, 100 or more metres in height, and is so narrow 
at the top that in some places goats can jump across it. It is 
therefore quite inaccessible for the greater part of its length, hence 
only ils lower part has been investigated. The sides were there 
usually covered with enormous mats of mosses of which the prin- 
cipal species were Hypnum falcatum, H. filicimnum, H. molluscum, 
Brachythecium rivulare and Philonotis fontana, all of which were 
developed to an unusually luxuriant degree. H. falcatum and H. 
filicinum were in many places thickly covered wilh ripe capsules. 

In this moss carpet a great number of other Bryophyta were 
found, partly intermixed with each other, partly in larger or smaller 
tufts or growths. The following species were observed: Philonotis 
tomentella, Bryum æneum, B. pallens, B. ventricosum, B. inclinatum, 
Mniobryum albicans, Mnium serratum, M. punctatum, Anomobryum 
filiforme, Encalypta ciliata, Didymodon rubellus, Barbula cylindrica, 
Distichium montanum, Hymenostylium curvirostre, Anæctangium com- 
pactum, Amphidium Mougeottii, A. lapponicum, Haplozia riparia. H. 
cordifolia, Lophozia quinquedentata, L. Miilleri, Scapania irrigua, 
Blephorostoma trichophyllaum, Madotheca Cordæana, Radula compla- 
nata, Metzgeria furcata and Marchantia polymorpha. At the base 
of the rock-side grew, among others, Mnium undulatum and Thui- 
dium tamariscinum. Åt the extreme end of the cleft, where the sun 
could shine on the rock faces, fruiting specimens of Preissia commu- 
tata often grew in abundance. The walls in the numerous caves 
were often covered with Eurhynchium Swartzii and Fegatella conica, 
in company with many, or a few, of the species found on the sides 
of the cleft, which, however, on the whole, did not thrive well in 
the scanty light. In many places Fegalella formed also a belt along 
the basal part of the rock-sides, especially where they projected. 
On the roof of the caves Blepharostoma trichophyllum and Ambly- 


608 A. HESSELBO 


slegium Sprucei formed thin, dark-green coverings, and. where the 
water flowed down the roof and walls Thamnium alopecurum oc- 
curred as pure growths. 

Å similar vegetation was found everywhere in clefts and caves, 
and where the conditions pertaining to light and moisture were 
more varying, the number of the species was still greater than in 
the vegetation described above. This was for instance the case in 
the deep and broad Flokastadagil near Breidabolstadr. There, 
on the damp walls, almost the same species were growing as in 





Fig. 33. Bryoziphium norvegicum on one of the sides of Flokastadagil. 


Bleiksågil. AÅA high, dry, perpendicular rock-face was for a consi- 
derable distance entirely -covered with a shining, dark-green mat of 
Bryoxiphium norvegicum (Fig. 33), which has hardly been found 
elsewhere in such enormous quantities. In other places Grimmia 
torquata covered the rock-sides and blocks with its irregularly-rounded 
cushions, and Hypnum palustre was very common everywhere at 
the water's edge. Neckera complanata was common in dry ditches. 

At the side of a small waterfall, Klitnafoss, there was a rather 
large cave, down one side of which some of the water from the 
waterfall was running. There Thamnium alopecurum grew abun- 
dantly, while the drier walls of the cave were covered with Eurhyn- 
chium Swartzii and Mnium orthorrhynchum. On the ground at the 


THE BRYOPHYTA OF ICELAND 609 


bottom of the cave there grew Timmia austriaca, Bryum ventricosum, 
Brachythecium rivulare, Thuidium delicatulum and Marchantia poly- 
morpha, and at the entrance to the cave where the water was drip- 
ping down, Mnium undulatum occurred. The upper part of the 
walls and the roof were covered with Blepharostoma trichophyllum, 
Amblystegium Sprucei, Metzgeria furcata and Radula complanata, and 
in the better-lighted part around the entrance, the cave was covered 
with Homalothecium sericeum and Orthothecium intricatum. 

The Southern Slope of Eyjafjall consists partly of basalt 
and partly of tufflayers. The numerous rivers which issue from 
the Jokull usually flow at the bottom of deep, narrow clefts in the 
tuff-layers. 

Holtså, which is a glacier-river of rather considerable size, 
flows for almost its entire course, from the Jékull down to the low 
land, at the bottom of deep clefts which are accessible only in a 
few places, the water usually rising from the bottom up to the 
perpendicular rock-wall. The sides åre, as a rule, rather damp, 
and the water from the slopes above often flows down the rock- 
faces... The Bryophyte vegetation is extremely luxuriant, and the 
sides of the clefts are, for considerable distances, covered with enor- 
mous carpets of mosses and liverworts. The following species 
covered the fallen blocks and débris at the base of the cleft-sides: 
Hylocomium spp. (especially H. squarrosum and H. proliferum), Hyp- 
num stellatum, Acrocladium cuspidatum, Climacium dendroides, Eu- 
rhynchium Stockesii, Thuidium tamariscinum, Mnium undulatum, M. 
punctatum, Timmia austriaca, Marchantia polymorpha, etc. In the 
numerous open caves and under overhanging rocks, a characteristic 
vegetation of Marchantiaceæ was found. Preissia commutata (fr.) and 
Fegatella conica covered the sides with metre-broad mats, and Re- 
boulia hemisphærica occurred abundantly in several places, and was 
also fruiling. Here Eurhynchium Swartzii also was at home, espe- 
cially on the less damp rock-faces, while Thamnium alopecurum 
preferred spots where the water was flowing down. Where stream- 
lets were running from above down the rock-sides, there grew 
Rhynchostegium rusciforme, Brachythecium rivulare, Haplozia cordi- 
folia and, lowermost, Thamnium, while the more or less damp rock- 
faces were covered with a variegated carpet of mosses and liver- 
worts, unusually rich in species and composed of Hypnum filicinum 
(in masses), H. falcatum (in masses), H. molluscum (in masses), Philo- 
nolis tomentella, P. fontana (in abundance), Mnium serratum, M. stel- 


610 A. HESSELBO 


lare, M. orthorrhynchum, Bryum æneum, B. inclinatum, B. pallescens, 
B. ventricosum, Mniobryum albicans (common), Pohlia cruda, Plagio- 
bryum Zierii, Anomobryum filiforme (in one place in fr.), Amphidium 
Mougeottii, A. lapponicum, Anæctangium compactum, Hymenostylium 
curvirostre, Blindia acuta, Ditrichum flexicaule, Distichium montanum, 
Barbula cylindrica, Didymodon rubellus, Tortella tortuosa, T. fragilis, 
Grimmia apocarpa, Myurella julacea, Haplozia riparia, H. atrovireus, 
Madotheca Cordæana, Lejeunea serpyllifolia, Radula complanata, Metz- 
geria furcata, Chiloscyphus polyanthus, Lophocolea cuspidata (scanty), 
Scapania irrigua, Lophozia quinquedentata, L. Miilleri and ÅAneura 
pinguis.. The moss-carpet covering dry rock-faces and large blocks 
was chiefly composed of Hylocomium loreum (also in fr.), H. proli- 
ferum, Hypnum uncinatum, H. hamulosum and H. callichroum. In a 
dry cave above the cleft there grew Neckera complanata, Brachythe- 
cium populeum, B. velutinum, Plagiothecium denticulatum, P. depressum, 
Isothecium tenuinerve, Mnium cuspidatum, and on the roof of the 
cave there grew Amblystegium Sprueei and Blepharostoma. 

The southern side of Drångshlidarfjall and the small neigh- 
bouring group of mountains, Hrutafjall (Fig. 34), consist, in a 
great measure, of dry, lofty faces of tuff-rock, as much as 200 metres 
high, the base of which is moss-grown. One of the commonest 
species here is Leucodon sciuroides var. morensis which covered the 
rock-face for a considerable distance upwards with its extensive and 
often richly-fruiting mats. The following species were also com- 
mon: Homalothecium sericeum, Pterigynandrum filiforme, Hypnum 
revolutum and Grimmia torquata. At the foot of the rock-faces there 
grew Isothecium myurum (in abundance), Orthotrichum rupestre, 
Didymodon rubellus, Barbula cylindrica, Bryum capillare, Tortella 
tortuosa, Encalypta ciliata, Neckera complanata, Myurella julacea and 
Metzgeria furcata, and where the water was oozing through cracks 
and crevices the rock-face was covered with Blindia acuta inter- 
mixed with Hypnum molluscum, H. palustre and the rare Anomo- 
bryum concinnaltum. 

Among the large blocks of tuff on a small Urd there grew, 
among others, Mnium undulatum, Eurhynchium piliferam, Rhyncho- 
steginm murale (abundantly), Amblystegium Juratzkanum, Leskea 
nervosa and Lejeunea cavifolia. 

The walls in the interior of a ralher dry, open cave, were 
covered with Preissia commutata, Fegatella conica, Marchantia poly- 
morpha, Distichium montanum, Bryoxiphium norvegicum, Anæclangium 


THE BRYOPHYTA OF ICELAND 611 


compactum (all in abundunce), Reboulia hemisphærica, Mnium or- 
thorrhynchum, Funaria hygrometrica, Bryum æneum, B. inclinatum, 
Orthothecium intricatum, Blepharostoma trichophyllum and, where it 
was somewhat damper, with Eurhynchium Swartzii. 








Fig. 34. Hritafell south of Eyjafjallajokull. The perpendicular face of tuff-rock in the 
centre, is about 200 metres high, and, at the foot is covered with extensive moss carpets, 


whilst caves occur in many places. 


AÅ very interesting kind of vegetation was found on the dry 
faces of a small isolated rock, Drångshellir. There grew Leucodon 
sciuroides var. morensis, Homalothecium sericeum, Leskea nervosa, 
Grimmia Doniana, Barbula muralis, B. obtusifolia, Orthotrichum ano- 
malum and O. rupestre. Half of these species consist of xerophi- 
lous southern forms, which have their northern limit in South Ice- 
land, and are there met with only in specially protected localities. 
Drångshlid, which has a height of 478 metres, protects this locality 


612 ÅA. HESSELBO 


from the cold, north winds, without, however, preventing the sun 
from warming the tuff-faces. 


The Bryophyte Vegetation of the Lava-fields. 


Å very great part of Iceland is covered with lava. According 
to Thoroddsen (1914, p. 219) the post-glacial lava-fields of Iceland 
cover an area of about 11,200 square km. or almost "/9 of the entire 
area of the island. It is, however, only a very small part of this 
vast area which has been an object of bryological investigation. 
Gronlund (1874, p. 136 and 1890) was ihe first to mention the 
vegetation of the lava-fields, in his description of the Rhacomitrium 
heath as important for the further development of the plant covering. 
Ostenfeld (1898, p. 246) has given a description of the lava-fields 
of the peninsula of Reykjanes without, however, entering more 
closely into the subject of the Bryophyte vegetation. Helgi Jonsson 
(1900, p. 89) has given a very detailed account of the vegetation of 
Budahraun and Eldhraun in West Iceland, also with regard to the 
Bryophyta, and the same author, in his description of the vegetation 
of South Iceland, mentions the Bryophyte vegetation of several lava- 
fields, especially modern ones. Lastly, A. Wegener in 1912 made 
a few collections on the high-lying Odådahraun (600—1200 metres 
above sea-level). 

Besides a small part of Myvatnshraun, around the farm Rey- 
kjablid, which was very superficially investigated owing to the 
inclemency of the weather, my own investigations include only the 
lava-fields of SW. Iceland, especially Hafnarfjardarrhraun, Thingvalla- 
hraun and Svinahraun around Kolvidarhol. 

The lava-field is no single formation, but, on the contrary, in- 
cludes a great number of different formations, ranging from the dry 
rock-flats and the Rhacomitrium heath, through the heather moor 
and grassland to the bog formations in the deepest hollows, and 
the associations of damp rocks in clefts. When it is nevertheless 
described here in a separate section, as by Helgi Jonsson, the 
reason is that -the vegetation of the lava-fields of the lowlands every- 
where presents so many common features and peculiarities that we 
may be justified in describing the Bryophyte vegetation occurring 
there. 

According to the nature of the surface, the lava-fields may be 
classified into two groups, viz. block-lava (Icelandic Apalhraun) and 
sheet-lava (Icelandic Helluhraun). The former consists of blocks of 


THE BRYOPHYTA OF ICELAND 613 


lava thrown together into irregular heaps, while sheet-lava consists 
of large, irregularly broken surfaces, which are formed by a process 
of slow solidification, and are furrowed by innumerable clefts and 
fissures, both narrow and broad. Here vesicular hollows are also 
found, produced by the moisture of the substratum which has formed 
large steam-vesicles in the molten lava, and after solidification the 





Fig. 35. Lava-field in the district of Borgarfjordur. The lava is covered to a depth of 
one foot with a layer of Rhacomitrium hypnoides. Projecting parts of the lava are covered 
with crustaceous lichens. Here and there a tuft of Festuca ovina. Birch coppice in 


the background. 


roof of the vesicle has collapsed, and has formed a platform on 


the floor of the cavity. Such vesicles in the lava — which are 
characterized by very luxuriant vegetation, since they afford excellent 
shelter for the plants, and are inaccessible to sheep — may differ 


rather greatly in depth, according to the thickness of the lava-sheet. 
In Budahraun, according to Jonsson, their depth averages from 
2—6 metres, but is usually somewhat above 2 metres, and in 
Hafnarfjardarrhraun the depth was also usually some 2 metres. The 
lava-fields are often furrowed by deep clefts or cracks, frequently 
of considerable length and with perpendicular sides. This is, for 
instance, the case in Thingvallabhraun where the best-known cleft, 
Almannagjå, has a length of almost 15 kilometres. 


614 A. HESSELBO 


The surface of the lava-field is very dry, because the water can 
easily drain away through the cracks, therefore, the vegetation there 
is decidedly xerophilous. As a rule, the Rhacomitrium heath (R. 
hypnoides) —— in which other Bryophyta, such as Dicranum sco- 
parium, Hylocomium proliferum and Pltilidium ciliare, occur only 
extremely scantily — covers the greater part of the lava-fields, and 
there arrives at its fullest development, so that extensive areas may 
be found covered with foot-deep, soft, greyisli carpets which hide 
all irregularities of the surface. The importance of the moss-covering 
for the further development of the plant-covering, parlly by binding 
the drifting sand, partly by forming soil, has been demonstrated by 
Gronlund and Helgi Jonsson. 

The chief reason for the extensive distribution of the moss-heath 
must undoubtedly be sought in the nature of the surface. This is, 
as a rule, highly vesicular, and contains numerous small holes and 
cavities, in which the plants find good conditions for taking root. 
Therefore, many of these cavities are filled up with small moss- 
cushions (Fig. 36), which, from thence, extend over the rock-surfaces 
and gradually merge into one another (Helgi Jånsson, 1900, p. 83). 

The tops of the lava-cones and the protruding blocks are not 
covered with a continuous moss-carpet, but usually with scattered 
cushions of mosses, liverworts and lichens. The most frequent 
species there are Rhacomitrium heterostichum, R. fasciculare, Grimmia 
apocarpa, G. funalis, Andreæa petrophila, Hypnum revolutum, Ho- 

malothecium sericeum, Pterigynandrum filiforme, Dicranoweisia cris- 
” pula, and occasionally Gymnomitrium coralloides, G. concinnatum, 
Dicranum fulvellum, Rhacomitrium sudeticum, Polytrichum pilosum, 
besides which there are several other species. Thus, near Hafnar- 
fjordur (and partly also in Bådahbraun), in addition to the majority 
of the species mentioned above, there occurred Orthotrichum rupestre, 
O. Sturmii, Grimmia patens, Leucodon sciuroides var. morensis, Isothe- 
cum tenuinerve and Frullania Tamarisci. Near Thingvellir, where 
the surface of the lava is very dry, there grew upon the lava-cones 
many crustaceous lichens, Rhacomitrium fasciculare (in abundance), 
Grimmia apocarpa, Andreæa petrophila, Polytrichum pilosum and, 
here and there, some Gymnomitrium coralloides and Dicranum ful- 
vellum; and, where some soil had accumulated, Ceratodon purpureus 
and Pohlia nutans grew scantily. In the depressions Rhacomitrium 
hypnoides was gradually replaced by R. canescens, heather moor and 
grassland, and in the deepest depressions small patches of bog and 


THE BRYOPHYTA OF ICELAND 615 


swamp vegetation may occur. The Rhacomitrium heaths are gra- 
dually transformed into heather moors and birch coppices through 
the decaying of the mosses and their forming humus. But all these 








Fig. 36. The surface of a lava block with small moss-cushions (Andreæa, 


Grimmia spp. and Gymnomitrium coralloides). 


formations will not be treated of more fully here as ihe mosses 
occurring in them have been mentioned in previous sections. Very 
extensively distributed and very characteristic is the Rhacomitrium- 
canescens formation which often replaces R. hypnoides in flat depres- 
sions where the ground is somewhat damp, and where some sand 
has accumulated. It is often abundantly mixed with other Bryophyta 
such as Hylocomium spp., Hypnum uncinatum, Dicranum scoparium, 
Polytrichum alpinum and species of Lophozia. 


616 A. HESSELBO 


Lava-clefts. The caves, clefts and cavities which occur every- 
where in the lava-fields contain a. Bryophyte vegetation very rich 
in species. In the broader clefts and in the lava-vesicles (Helgi 
Jønsson's ”"Herb-cavities”, 1900, p. 90) the bottom is often covered 








Fig. 37. Lava-cleft near Thingvellir (Almannagjå). The lava-blocks are 


quite covered with Rhacomitrium hypnoides. 


with ferns and flowering plants, and especially in deeper clefts, or 
where the bottom is covered with loose blocks, there is also found 
a continuous carpet of mosses and liverworts in which Hylocomium 
spp. (H. proliferum, H. parietinum, H. loreum, H. squarrosum and 
H. triquetrum) together with species of Rhacomitrium, Hypnum and 
Dicranum play a prominent part, and are usually much interwoven 
with liverworts such as Lophozia lycopodioides, L. quinquedentata, 
L. Kunzeana, Cephalozia spp., Alicularia scalaris, Ptilidium ciliare, etc. 


THE BRYOPHYTA OF ICELAND 617 


Very frequently a large form of Hypnum callichroum, with regularly 
two-rowed ramification, is the most abundant constituent of the 
vegetation, especially in somewhat damp and shady clefts. Eurhyn- 
chium piliferum is often met with very abundantly, especially where 





Fig. 38. Lava-cleft near Thingvellir. At the bottom there is a very luxuriant vegetation 


of ferns, Geranium silvaticum, etc., and in the crevices large moss-cushions occur everywhere. 


the bottom is grass-covered, and several other southern forms, for 
instance Eurhynchium Stockesii and Isothecium myurum, likewise 
have a preference for the sheltered lava-clefts. 

In the very deep lava-clefts the conditions are somewhat dif- 
ferent from the aforesaid. Where the cleft is broad enough to allow 
the sun to shine on the bottom during a part of the day, a luxuriant 
covering of ferns, flowering plants or mosses is found (Fig. 38); but 
where the cleft is so narrow that the light is strongly subdued, the 


The Botany of Iceland. Vol. I, part Il. 40 


618 A. HESSELBO 


higher plants and the Hylocomium mats do not thrive. The snow 
lies here till far into the summer, and in many cases the tempera- 
ture hardly ever rises above freezing point. The bottom of these 
clefts is therefore either quite bare of vegetation, or covered only 
with mosses. Åt the bottom of narrow clefts (150—20 metres or more 
in depth) near Thingvellir, where snow and ice were still to be 
found even at the end of July, Anthelieta occurred on a damp sub- 
stratum as they do on flats irrigated by melting snow on mountain 
heights. Polytrichum sexangulare and Pohlia commutata grew abund- 
antly here as on rocky flats. The plants were considerably higher 
and more slender than on the rocky flats, and only slightly tomen- 
tose. The same was the case with Conostomum boreale and Mniobryum 
albicans, which likewise occurred in several clefts. The Bryophyte 
vegetation occurring there is affected both by the low temperature 
and also the deficient light and total absence of wind, which all 
combined contribute to produce a stronger longitudinal growth and 
a slighter development of rhizoids. Several other species, which 
have their main distribution on mountain heights, and are rare in 
or quite absent from the lowlands, are widely distributed about the 
lava-fields. In almost all the lava-fields of South-west Iceland, 
Pleuroclada albescens var. islandica, Dicranum molle, D. Blyttii and 
D. Starckei are commonly found in clefts, not only in those that 
are narrow, but especially in the broad moss-grown clefts, where 
Dicranum spp. form large tufts on the blocks of lava. 

On the sides of the clefts and of the fallen blocks, a great 
many mosses grow. Diplophyllum albicans is a character-plant of 
the lava-fields of SW. and W. Iceland; it is extremely common there, 
but is very rare and scanty in other localities. On the vertical, dry 
sides of clefts the mosses usually grow in large rounded cushions. 
The commonest species are Tortella tortuosa, ÅAnæctangium com- 
pactum, Amphidium Mougeottii, Grimmia torquata and G. funalis, but 
many other of the species occurring on rocks and on the ground 
are always found both on the walls and also on, and among, the 
blocks. The Bryophyte vegetation of lava-clefts has the greatest 
resemblance to that found on the Urd, but some of the most light- 
loving species are absent, for instance Orthotrichum, several Grimmia 
and Rhacomitrium spp. and Dicranoweisia crispula, while, on the 
other hand, others are found which belong to damp clefts or which 
need more shelter. Mesophilous forms are in the majority, whilst 
hygrophilous forms are, as a rule, entirely absent. 


THE BRYOPHYTA OF ICELAND 619 


In the following pages some examples of the Bryophyte vege- 
tation of lava-clefts will be described. 

Thingvallahraun. The lava-clefts (Icelandic Gjå) there, are 
peculiar on account of their unusual depth (as much as 20—30 
metres). Several of those situated nearest to the lake of Thingvellir 
have deep water at the bottom. 

The Bryophyte carpet at the bottom of the clefts is composed 
of Hylocomium loreum, H. proliferum, H. parietinum, H. squarrosum, 
Hypnum uncinalum, H. callichroum, H. stramineum, H. molluscum, 
Eurhynchium piliferum, Antitrichia curtipendula, Polytrichum alpinum, 
Rhacomitrium hypnoides, R. canescens, Dicranum Blyttii, D. Starckei, 
D. molle, Sphagnum teres, S$. Girgensohnii, Lophozia ventricosa, L. 
quinquedentata, L. Kunzeana, L. alpestris, Ptilidium ciliare and Sca- 
pania irrigua. On the sides of the clefts there grew especially An- 
æclangium compactum, Grimmia torquata, G. funalis, Gymnostomum 
rupestre, Amphidium Mougeottii and A. lapponicum. Intermixed with 
these, and in cushions on the ledges, in the crevices and among 
the blocks there grew: Hypnum cupressiforme, Isothecium tenuinerve, 
Plagiothecium pulchellum, P. denticulatum, P. silvaticum, Amblystegium 
Sprucei, Neckera complanata, Schistidium gracile, Rhacomitrium sude- 
ticum, Sælania cæsia, Ditrichum flexicaule, Tortella tortuosa, Fissidens 
osmundoides, F. bryoides (rare), Pohlia cruda, P. commutata, Bartramia 
ityphylla, Conostomum boreale, Philonotis tomentella, Mnium orthor- 
rhynchum, Bryum pallens, Plagiobryum Zierii, Distichium montanum, 
Diplophyllum albicans (in abundance), Lophozia heterocolpos, Lejeunea 
cavifolia, Blepharostoma trichophyllum, Alicularia scalaris, Å. geo- 
scypha, Anthelia Juratzkana, Scapania subalpina, Pleuroclada albescens 
var. islandica, Cephalozia bicuspidata, Eucalyx subellipticus, Gymno- 
mitrium concinnatum, Metzgeria furcata, Radula complanata, Aneura 
pinguis and Preissia commutdata. 

The above list is very long, but is nevertheless hardly complete, 
since some of the common chomophytes are not enumerated in it, 
and, at any rate, several of them were undoubtedly forgotten, while 
the list was being made on the spot, but it gives a good idea of 
the exceedingly great number of species which are found in these 
localities. 

Hafnarfjardarhraun. Almost the same species are found 
there as in Thingvaliahraun, but the situation close to the sea-side 
makes its influence felt. Rhacomitrium heterostichum and R. fasci- 
culare are exceedingly common, and in several places Ulota maritima 

40” 


620 A. HESSELBO 


occurs on lava-blocks and Trichostomum littorale on the sides of the 
clefts. Scapania dentata was frequent on the ground at the bottom 
of shady, damp clefts. Reboulia hemisphærica and Frullania fragili- 
folia were found in a single spot on the walls of caves. 

On the ground at the bottom of deep, dark caves, shade-forms 
are often found, especially of Hepaticæ, with greatly elongated shoots 
and expanded, widely scattered leaves. Here, it was especially the 
following species which were common : Cephalozia bicuspidata, Pleuro- 
clada albescens var. islandica, Alicularia scalaris and Blepharostoma 
trichophyllum, occasionally also Preissia commutata and in a single 
spot Ditrichum homomallum and Oligotrichum hercynicum. 

Svinahraun is situated about 30 km. south-east of Reykjavik, 
and about 250—300 metres above sea-level. The part which has 
been investigated is situated in the neighbourhood of the farm Kol- 
vidarhol, and is of block-lava consisting of blocks and flags piled 
up into wild-looking heaps, some 10 metres high, with numerous 
clefts and caves. The top part is covered with an unusually thick 
and luxuriant mat of Rhacomitrium hypnoides from which other 
mosses and flowering plants are almost absent. On the walls there 
grew Amphidium Mougeottii, Tortella tortuosa and Anæctangium com- 
pactum. In caves and clefts there was a Bryophyte carpet consisting 
of Hypnum uncinatum, H. callichroum, Hylocomium loreum, H. proli- 
ferum, H. parietinum, Lophozia quinquedentata, L. lycopodioides, L. 
Flærckei, Plagiochila asplenioides and Ptilidium ciliare. Diplophyllum 
albicans was extremely common everywhere. In addition to the 
above the following were found more or less abundantly: Polytrichum 
alpinum, Diphyscium sessile, Bartramia ityphylla, Conostomum boreale, 
Mnium orthorrhynchum, Pohlia cruda, P. commutata, Mniobryum albi- 
cans, Bryum elegans, Amphidium lapponicum, Rhacomitrium sudeticum, 
Schistidium gracile, Fissidens osmundoides, Dicranum scoparium, Di- 
stichium montanum, Sphagnum teres, Lophozia alpestris, L. quadriloba, 
Alicularia scalaris, Anthelia Juratzkana and Blepharostoma tricho- 
phyllum. 

It will be seen that the lava-fields here described greatly re- 
semble each other, and that it is essentially the same species which. 
constitute the bulk of the vegetation in all of them. Svinahraun 
is situated on the ridge of hills which extends along the whole 
length of the peninsula of Reykjanes, and the climate there is ex- 
tremely cold and damp, which is proved by the fact that it is the 
Rhacomitrium heath which forms the last stage in the development 


THE BRYOPHYTA OF ICELAND 621 


of the vegetation of the flats, and that many of the species of the 
low land are absent. 

Unfortunately there was no opportunity of investigating the 
lava-fields of West Iceland more closely, since unfavourable atmos- 
pheric conditions with rain and fog, combined with a want of time, 
made a longer stay there impossible, but the conditions appear to 
be essentially the same as in South-west Iceland. 

In his description of Budahraun Helgi Jonsson has also men- 
tioned soine mosses, and, as far as is possible to judge from the 
lists which, however, are no doubt incomplete, the conditions there 
are exactly like those in Hafnarfjardarhraun. There also coastal 
species are found, for instance Ulota maritima, and southern species 
such as Eurhynchium Stockesii and Lejeunea cavifolia, and the 
dominant species are quite similar to those in the lava-fields of 
South-west Iceland. 

In North Iceland the Bryophyte vegetation of the lava-fields has 
an essentially different composition from that in West and. South- 
west Iceland. All the lava-fields there are situated at a distance 
from the coast, and at a higher level above the sea, the vegetation 
is therefore decidedly xerophilous both on exposed surfaces and in 
clefts and crevices. The immense lava-flats at a great height above 
sea-level, are deserts almost entirely void of vegetation; they have, 
however, been very little investigated. There are to hand only a 
few collections from Koch's tour across Odådahraun in 1912, and 
these consist only of Tortula ruralis, Ceratodon purpureus and Di- 
cranoweisia crispula, which grew here and there upon the lava. 

The only lava-field which has been investigated is Myvatns- 
hraun, which is about 160 years old, and upon which the farm 
Reykjahlid stands. It was described by Gronlun d in 1890 and does 
not appear to have undergone any great change since that time. 
The Rhacomitrium heath occurred only scantily in the depressions, 
while the lava-cones were bare of vegetation or covered with scat- 
tered moss-cushions and crustaceous lichens. In the intervening 
space between the farm and the lake the following species were 
observed: Upon the blocks and on the lava were found Tortula 
ruralis, Ceratodon purpureus, Rhacomitrium hypnoides, Grimmia Don- 
iana, Schistidium confertum, Dicranoweisia crispula, Polytrichum pi- 
losum and Hypnum revolutum, all of which were very common. In 
sandy soil, especially in depressions, there grew Desmatodon latifolius 
and Rhacomitrium canescens. In dark caves and clefts Lophozia 


622 A. HESSELBO 


lycopodioides was usually the only species found and it formed there 
extensive mats. On the ground among the blocks and on the sides 
of these blocks as also on those of broad clefts there grew Brachy- 
thecium reflexum, Lescuræa filamentosa, Eurhynchium diversifolium 
(scantily), Polytrichum alpinum, Timmia austriaca, Bartramia ityphylla, 
Encalypta rhabdocarpa, Bryum elegans, B. inclinatum, Schistidium 
apocarpum, Didymodon rubellus and Plagiochila asplenioides. 

The above is a decidedly xerophilous flora from which not only 
all the mesophilous and hygrophilous forms from SW. Iceland, but 
also the lowland or southern forms proper, are entirely absent. 

In several places in this lava-field the heat of the substratum 
makes itself felt in deep caves and clefts. In a warm, damp cave 
where the temperature was about 25? (outside 49-59) there grew 
Fissidens osmundoides, Plagiothecium denticulatum, Calypogeia Tri- 
chomanis, Plagiochila asplenioides and Sphagnum rubellum. 

The development of the Bryophyte vegetation of the lava-fields 
has been very little investigated. The Bryophyta together with the 
lichens are the first plants which appear. The surface of the lava 
is very rough, consisting of small round cavities (lava vesicles) in 
which the spores find favourable conditions for germination, and 
form small rounded cushions which adhere very closely to the sub- 
stratum and, if conditions are favourable, extend over the entire 
surface of the lava as a continuous carpet. It is, however, only in 
the most expøsed parts of the lava-field, that the mosses are con- 
fined to the vesicles; this is not the case in the clefts. 

Jénsson (1905, pp. 55 and 56) has given a description of Kra- 
katindshraun near Hekla, a lava-field about 23 years old. The 
mosses grew there only in tiny, scattered cushions upon the lava, 
without anywhere forming continuous carpets; down in the clefts 
ihey grew somewhat more abundantly. There the following species 
were found: Bartramia ityphylla, Pohlia cruda, P. commutata, Bryum 
pallens, Bryum spp., Ceratodon purpureus, Dicranoweisia crispula (the 
commonest species), Rhacomitrium hypnoides, R. canescens, Scapania 
curta and Lophozia alpestris. The further development is exceedingly 
slow. On the flats Rhacomitrium hypnoides will usually grow over 
all the species and suppress them, but the formation of a continuous 
Rhacomitrium heath appears to require a very long time, in many 
cases, centuries, and the further development of the moss-heath into 
heather moor, birch coppice or other formations, through the decay 
of the mosses and their forming soil, undoubtedly takes place even 
more slowly. 


THE BRYOPHYTA OF ICELAND 623 


IAÆFHE BRYOPHYTEVEGETATION OF MOUNTAIN HEIGATS 


In "Vegetationen paa Snæfellsnes” (Helgi Jønsson, 1900) the 
plant formations of mountain heights are classified into three groups, 
viz., Rocky flat, Grimmia heath and Mountain bogs. Of these the 
Grimmia heath (Rhacomitrium heath) has already been described in 
a previous section. 


Rocky Flat. 


By Rocky Flat Jonsson understands that part of the country 
which is situated above the upper limit of the heather moor (and 
the birch), with the exception of the mountain bogs and the Rha- 
comitrium heath. Its surface varies greatly, and consists sometimes 
of gravelly or clayey flats, sometimes of horizontal or sloping stony 
tracts, bare rocks or talus of débris, all of which, however, have 
one characteristic in common, viz., that higher plants do not form 
any continuous carpét, but grow scattered singly or in small societies 
in the most sheltered localities. The lower limit of the rocky flat 
varies greatly according to the local conditions. It generally begins 
at an altitude of about 300—400 metres, but often typical rocky-flat 
formations are met with at far lower levels, in NW. Iceland, for 
instance, as far down as to the sea-level. 

Bryophyta play a very prominent part in the vegetation of the 
rocky flat, and are often dominant there, or form special Bryophyte 
associations. 

The dry, gravelly flats are poor in plants, and are often quite 
devoid of vegetation. Of mosses, as a rule, only Rhacomitrium hyp- 
noides is met with in scattered cushions, especially around some- 
what large stones, where it can hold fast, and find some shelter. 
In somewhat damper and less exposed localities Rhacomitrium heaths 
are developed as in the lava-fields of the low land, but whilst the 
mosses in the low land gradually make room for other plant asso- 
ciations by accumulating the blowing sand around them, and by 
forming humus, this is not the case on the rocky flat. Here flowering 
plants occur extremely scantily in the Rhacomitrium heath, which 
therefore forms the final stage in the development of the plant- 
covering, and is not replaced by other formations. 

On large stones on the rocky flat there grew jet-black cushions 
of Andreæa petrophila, Dicranoweisia crispula, Rhacomitrium fascicu- 
lare, R. sudeticum and Schistidium apocarpum. 


624 A. HESSELBO 


On somewhat damper gravelly ground, especially where large 
blocks lie scattered and afford shelter, the Bryophyte vegetation be- 
comes richer, so that here we sometimes find a continuous carpet 
of mosses and liverworts, through which the tops of the stones 
protrude. This carpet of mosses and liverworts is usually composed 
of a larger or smaller number of the following species: Hylocomium 
proliferum, Hypnum uncinatum, Brachythecium reflexum, Lescuræa 
Breidleri, Rhacomitrium hypnoides, R. fasciculare, R. sudeticum, R, 
canescens, Schistidium apocarpum, $. gracile, Bryum ventricosum, Pohlia 
commutata, P. cucullala, P. gracilis, Philonotis fontana, Conostomum 
boreale, Dicranum Blyttii, D. molle, D. Starckei, D. congestum, Poly- 
trichum sexangulare, Oligotrichum hercynicum, Lophozia alpestris, L. 
quinquedentata, L. ventricosa, L. lycopodioides, L. Flærckei, L. quadri- 
loba, Alicularia scalaris, A. geoscypha, Pleuroclada albescens, Anthelia 
Juratzkana and Gymnomitrium concinnatum besides some rare or 
more casual species, for instance Eurhynchium diversifolium, Brachy- 
thecium glaciale, Aulacomnium turgidum, Bryum elegans, Bartramia 
ityphylla, Pohlia Ludwigii, Didymodon rubellus, D. rufus, Desmatodon 
latifolius, Distichinm montanum, etc. Usually several species grow 
intermixed, but the moss carpet may also be formed by a single 
species or a few. At elevations above 500—600 metres Rhacomitrium 
sudeticum often forms extensive growths on gravelly flats. Flowering 
plants are entirely absent, but lichens (Cetraria islandica, Cladonia 
spp. and Stereocaulon condensatum) may occur abundantly in this 
moss carpet. In the more low-lying parts of the rocky flat Rha- 
comitrium sudeticum vrarely occurs abundantly, but is replaced by 
Schistidium gracile and Schistidium apocarpum, which form low, 
blackish-brown or reddish-brown mats. Lescuræa Breidleri is a com- 
mon species, especially in NW. Iceland, where it is in many places 
the most abundant constituent of the vegetation on stony slopes. 
Conostomum boreale is also common, especially in NW. Iceland 
where, on the dry gravelly flats of the mountain heights, the inter- 
vening spaces between the stones are filled up with its compact 
tufts. Brachythecium glaciale in company with Hypnum stramineum 
covered the damp areas of the rocky flat near Isafjordur, at a height 
of 450—500 metres. In the channels, in which the water is drained 
away during the melting of the snow, the stones are often entirely 
covered with Mniobryum albicans var. glacialis, and in damp spots 
and near springs Philonotis fontana forms light-green cushions, as 
it does near the "Dy” in the more low-lying tracts. In the following 


THE BRYOPHYTA OF ICELAND 625 


pages some examples of the rocky-flat vegetation will be given, it 
is, however, only in NW., SW. and in a few districts of N. and E. 
Iceland, that this vegetation has been somewhat closely investigated, 
whilst the whole of the interior high land is quite unknown as far 
as bryology is concerned. 

1. South Iceland. Barkarstadr, dry stony flat al an alti- 
tude of 350 metres: Scattered cushions of Schistidium apocarpum v. 
rufescens, Rhacomitrium hypnoides, R. fasciculare, Dicranoweisia cris- 
pula and Andreæa petrophila. 

2. Gravelly flat on Esja, at an altitude of about 550 metres: 
Moss carpet, partly torn up by the wind, composed of Hypnum 
uncinatum and Schistidium apocarpum, scattered in which occur 
rocky-flat plants such as Silene acaulis and Saxifraga oppositifolia. 

3. Akureyri, rather dry gravelly flat, at an altitude of about 
900 metres: Moss carpet composed of Rhacomitrium hypnoides, Hyp- 
num uncinatum, Hylocomium proliferum, Camptothecium nitens, Aula- 
comnium turgidum, Dicranum congestum, Didymodon rufus, Eurhyn- 
chium diversifolium, Lophozia quinquedentata and L. quadriloba. 
Flowering plants entirely absent. On a dry slope at an elevation 
of about 770 metres there grew Lescuræa Breidleri, Brachythecium 
glaciale, B. reflecum, Dicranum Starckei, Lophozia lycopodioides and 
L. alpestris, and on damp gravelly flats near the snow line there 
grew Pohlia gracilis (abundantly), P. cucullata, P. commutata, Poly- 
trichum sexangulare, Oncophorus virens, Dicranum Starckei, Lophozia 
alpestris, L. ventricosa, L. quinquedentata, L. quadriloba, Pleuroclada 
albescens and Anthelia Juratzkana. 

Salix herbacea-Sibbaldia Vegetation (Helgi Jonsson, 
1900, p. 33) occurs everywhere in damp depressions and on slopes 
where the subsoil consists of a layer of clay. This vegetation has 
iis main distribution from 300—400 metres to about 700 metres, 
and here, close to the snow line, it is replaced by pure moss-com- 
munities, in which Salix herbacea occurs but scantily and at last 
disappears entirely. The moss community consists of a low, dense 
carpet of mosses and liverworts, interwoven with creeping stems of 
Salix herbacea, so that only the tips of the shoots protrude above 
the carpet. The most abundant constituent of this moss carpet is 
usually a low-growing form of Hypnum uncinatum or of Dicranum 
Starckei in association with Rhacomitrium canescens, Dicranum molle, 
Conostomum boreale, Polytrichum sexangulare, Anthelia Juratzkana, 
Pleuroclada albescens, species of Lophozia, and sometimes other species. 


626 A. HESSELBO 


ÅAnthelia flats (Helgi Jonsson, 1900). In damp parts of the 
rocky flat, especially where the snow-water from the melting snow- 
flats spreads out over the ground, extensive flats of a greyish-black 
or bluish-black colour are very commonly met with, and these are 
principally formed of Anthelia Juratzkana. Interspersed in this 
Anthelietum occur several other Bryophyta of which the most 
frequent are Alicularia scalaris, A. geoscypha and Lophozia alpestris, 
while scattered plants of Polytrichum sexangulare, Oligotrichum her- 
cynicum and Pohlia gracilis protrude here and there. Pleuroclada 
albescens is also occasionally met with. 

Although the Anthelia vegetation also occurs in more low-lying 
tracts, yet it has its main distribution near the snow line, and must 
therefore be reckoned to the moss associations of the Snow region. 
There are only a few Bryophyta which occur in the Snow region. 
At the boundary between the Anthelia-flats and the drier gravelly 
flats Dicranum falcatum occurs in semiglobular cushions, densely 
matted with rhizoids, but it lies so loosely upon the gravel that 
one can lift up the whole cushion. Near Dyrafjéordur Gymnomitrium 
varians grew in a similar manner to Anthelia, close to the melting 
snow-flats.. Dicranum Starckei, D. Blyttii, Pohlia cucullata, P. com- 
mutata, Oligotrichum hercynicum and Polytrichum sexangulare are 
also common near the snow line but, with the exception of a few 
species (Dicranoweisia, Andreæa and Rhacomitrium spp.) growing on 
rocks, the list is thereby exhausted. 


Mountain Bogs 


are extensively distributed in great parts of Iceland, but have not 
yet been more closely investigated. The vegetation in the boggy 
depressions often consists of a low, black mat of Hypnaceæ, mostly 
Hypnum revolvens, H. exannulatum, H. sarmentosum and H. strami- 
neum; but the moss covering may also contain other species and, 
in composition, somewhat resemble the bog vegetation of the low 
land, although the species are fewer in number. 

Near AÅkureyri boggy flats, situated 500—600 metres above 
sea-level, were partly covered with Hypneta and partly with Cin- 
clidium stygium with scattered cushions of Sphagnum and species 
of Lophozia. On a wet flat, at an altitude of about 600 metres, 
there grew Polytrichum commune (in abundance), Hypnum sarmen- 
tosum, H. stramineum, H. Lindbergii, H. uncinatum, Dicranum Starckei, 
Sphagnum teres, S. Girgensohnii, Meesea trichoides, Bryum ventricosum, 


c— 


(Ul 


THE BRYOPHYTA OF ICELAND 


munoyopns umgmuosvyy UNA Jey & st ozoy)y puno1g-ojpprur oy1 ur JyoT au) og 
10990 1170970 / Umunddig pue Vole yqVUuV souojs oy) uoom4øq punoag oy1 uo puwe 'suorysno 428|q aArsuo)x0 ur sm401g 
[940] Bos oA0OqgU somnow Q0G moqe umpzoljesg deou Je ÅÆyooy 


wmoyapns wmsmutomyy souojs zogaey oyu1 UuQ 


"GE 


Q 


olg 





S 


mia longipe 


O 


Dissodon splachnoides, Dichodontium pellucidum, Aongstr 


and Lophozia Kunzeana. 


On a tract of knolly bog- 


land, 530 metres above sea-level, there grew Hypnum turgescens, H. 


Barkarstadr (South. Iceland). 


628 A. HESSELBO 


revolvens, H. exannulatum, H. Richardsonii, H. sarmentosum, H. poly- 
gamum, Hylocomium squarrosum, Oncophorus Wahlenbergii, Dicho- 
dontium pellucidum, Aongstræmia longipes, Bryum ventricosum, Pohlia 
gracilis, Dissodon splachnoides, Dicranum congestum (in the Sphagnum- 
tufts), Søohagnum teres, Alicularia scalaris, Scapania irrigua and An- 
thelia Juratzkana. 

Jud som a botanical point of view, the interior high land 
has be very superficially investigated. Ås regards the mosses, only 
a few collections are to hand, which were made by A. Wegener 
during Koch's tour through the island in 1912. But as regards 
almost all these collections, conditions pertaining to soil and height 
above sea-level have not been more closely notified. 

Hvannalindir (altitude 656 metres): Hypnum stellatum, H. re- 
volvens, Philonotis fontana, Pohlia commutata (fr.), Distichinm mon- 
tanum, Splachnum vasculosum and ÅAongstræmia longipes. 

Eyolfsfjall (June 22nd): Aulacomnium palustre, Philonotis to- 
mentella, Bartramia ityphylla, Bryum ventricosum, Cinclidium stygium, 
Timmia norvegica, Mnium affine var. integrifolium, Pohlia cruda, Di- 
stichium montanum, Dichodontium pellucidum, Hypnum sarmentosum, 
Amblystegium Sprucei and Anthelia Juratzkana. 

Esjufjall. Rhacomitrium canescens, R. hypnoides, Distichium 
montanum, Philonotis tomentella, Mnium orthorrhynchum, Pohlia cruda, 
Bryum spp., Hypnum uncinatum, H. revolutum and Anthelia Juratz- 
kana. 

Wet Bogs and River banks; June 22nd (without more parti- 
cular specification of the locality). Hypnum exannulatum, H. Kneiffii, 
Aulacomnium palustre, Philonotis fontana, P. tomentella, Mniobryum 
albicans var. glacialis, Pohlia commutata, Distichium montanum, Schis- 
tidium gracile and Aongstræmia longipes. 

These lists almost exclusively contain species which are of 
common occurrence everywhere. 





FREE ERCOMLONEN FS 
OFTE BRYOPHYTE FEORÆSE 


IR consequence of the geographical situation and climate of the 

island, the vegetation of Iceland has a composition which cor- 

responds most nearly to that of Scandinavia and South Greenland. 

With the exception of the species found only in Iceland, there is 

only one species (Bryoxiphium norvegicum) which has not been 

found in the other parts of Europe, but which has its home in 

North America, and one species (Tortula obtusifolia) which I do not 

think has been found in Scandinavia. The other species are all 

common to Iceland and Scandinavia. At present it is not possible 
to compare the Bryophyte flora of Iceland and Greenland, partly 
because there is no collective account of the distribution of the 
species found in Greenland, and partly because the older Greenland 
collections and lists of species greatly need revision. 

The Bryophyta found in Iceland can be arranged in four geo- 
graphical groups (Arnell and Jensen, 1910, p. 238): — 

1. Ubiquitous species, which are almost equally distributed 
over the whole of North Europe as far as north of the Arctic 
Circle. 

Meridional species, which are most frequent in Southern 

Scandinavia and far rarer in Northern Scandinavia. 

3. Boreal species, which are more frequent in Northern than 
in Southern Scandinavia, and have their main distribution 
below the tree-limit. 

4. Alpine species, which have their main distribution above 


9 


the tree-limit. 

In Table II and in the following Tables the plant-geographical 
character of the species is denoted by the initial letters u, m, b and 
a, which are prefixed to the name of the species. Icel. indicates 
that the plant is known only from Iceland. As already mentioned, 
the species found only on warm soil — almost all of which belong 


630 A. HESSELBO 


to the meridional group — are not included in the present or fol- 
lowing sections. 


Table I. The Plant-geographical Groups. 




















| u m b a Icel. 
É ze INSEE = re 
Hepaticæ SIE eee erne 28 18 TOR 22 
Sphasnales ES SEE SE ol 2 | 4 | 
Maven e  eE DENN re EA KS EEN 6 
Total 132 En ER 6 











Of greatest interest in this connection is the comparatively large 
group of meridional species, which comprise about "7/6 of all the 
species. According to their distribution in the different districts of 
Iceland they can be divided as follows: — 

a. Found only in the Southern part of Iceland (South Iceland 
and most nearly adjacent parts of South-east and South-west Iceland). 


Reboulia hemisphærica. 
Fegatella conica. 
Fossombronia Dumortieri. 
Aneura multifida. 
Marsupella Funckii. 
Jamesoniella autumnalis. 
Lophozia excisa. 
Lophocolea cuspidata. 
Lepidozia setacea. 
Frullania fragilifolia. 
Lejeunea cavifolia. 
Sphagnum papillosum. 
Weisia crispata. 

HE viriduld 
Trichostomum littorale. 
Barbula unguiculata 

var. cuspidata. 
Tortula muralis. 
— mucronifolia. 


b. Most common in South Iceland, rarer in the other parts of 


the island. 


Madotheca Cordæana. 
Barbula cylindrica. 
Mnium serratum. 
Catharinea undulata. 
Neckera complanata. 


Hedwigia albicans. 
Orthotrichum anomalum. 
— saxatile. 
= cupulatum. 
Mnium undulatum. 
Leucodon sciuroides. 
AÅnomodon viticulosus. 
Thuidium tamariscinum. 
= Philiberti. 
Isothecium myurum. 
Scleropodium purum. 
Eurhynchium piliferum. 
== Swartzii. 
= Stockesii. 
Rhynchostegium murale. 
Plagiothecium depressum. 
== elegans. 
Amblystegium fluviatile. 
Hypnum molluscum. 


Thuidium delicatulum. 
Thamnium alopecurum. 
Rhynchostegium rusciforme. 
Camptothecium lutescens. 
Hypnum commutatum. 


THE BRYOPHYTA OF ICELAND 631 


c. Found only in a single locality, or in a few localities, in 
the different paris of Iceland. 


Haplozia crenulata. Bryum bimum. 

Lophozia barbata. Mnium cuspidatum. 
Lophocolea minor. Eee Stellare: 
Diplophyllum obtusifolium. Cylindrothecium concinnum. 
Sphagnum angustifolium. Eurhynchium strigosum. 
Dicranum Bonjeani. Amblystegium serpens. 
Trematodon ambiguus. = Juratzkanum. 
Fissidens bryoides. — trichopodium. 
Leptodontium flexifolium. Hypnum Sommerfeltii. 
Barbula fallax. = Sendtneri. 


d. Frequent everywhere, or in large parts of Iceland. 


Radula complanata. Common in South, East and West iceland, 
absent from North Iceland. 

Rhacomitrium aciculare. Very common all over Iceland. 

Mnium hornum. Common, especially in South Iceland, rarer 
in North Iceland. 

Pogonatum nanum. Common in several parts of West and 
South-west Iceland. 

Metzgeria furcata. Common over the whole of Iceland. 


The majority of the meridional species (a = 37 species and b = 
10 species) have a decidedly southerly distribution. Of the species 
belonging to group c only Lophozia barbata, Diplophyllum obtusi- 
folium and Bryum bimum have been found, each in a separate 
locality in East Iceland. The other habitats are equally distributed 
over South, West and North Iceland. 

Table I shows that there is an almost equal number of Boreal 
and Alpine species and a somewhat larger number of Ubiquists, . 
but in order to obtain a really comprehensive view of these groups, 
showing which has the greatest importance with regard to the com- 
position of the vegetation, it is also necessary to investigate how 
many species with a somewhat high degree of frequency are con- 
tained in each group. 

If all the species are included, the frequency of which — 
throughout a larger part of Iceland (besides South Ieeland) — is 
expressed by the figure 3 (see p. 644) or upwards, the number of 
species contained in each group will be as follows: — 


632 


" A., HESSELBO 


Table II. Species with the Frequency of 3 or upwards throughout 
a larger part of Iceland. 






































| u m b a 
= gær ! 2 É 
Hepaticæ ENS SANSER RSS | 15 3 7 9 
Sphagnales AE ENE ESS RR 4 
Mas N er rer er NE ORE RS | 55 (ØR SE Mer SA 
SEERE REE ED Er 
| 





Group 1. 


=Marchantia polymorpha. 
”Aneura pinguis. 
=Pellia Neesiana. 
=Alicularia scalaris. 
=Lophozia quinquedentata. 
== ventricosa. 
— Mulleri. 
æPlagiochila asplenioides. 


Sphagnum Girgensohnii. 
< — teres. 

— Warnstorffii. 
— rubellum. 


Gymnostomum rupestre. 
Ånæctangium compactum. 
=Dicranum scoparium. 

= congestum. 
Ceratodon purpureus. 
”Ditrichum flexicaule. 
"Didymodon rubellus. 
=Tortella tortuosa. 
Tortula subulata. 

— ruralis. 

Schistidium maritimum. 

— apocarpum. 

— gracile. 
Rhacomitrium heterostichum. 

— canescens. 
== hypnoides. 

Amphidium Mougeottii. 
Ulota maritima. 
Orthotrichum rupestre. 
Encalypta rhabdocarpa. 


+ 








Ubiquitous species. 


æCephalozia bicuspidata. 

£ — Hampeana. 
=Blepharostoma trichophyllum. 
=Ptilidium ciliare 

Scapania dentata. 

Fy be undulata. 

"Frullania Tamarisci. 


Funaria hygrometrica. 
=Leptobryum pyriforme. 
=Pohlia cruda. 

”"Bryum inclinatum. 


= argenteum. 

— pallens: 
£ —… ventricosum. 
=Mnium affine. 

ES eeerr 
SE Ep unetatum 


Meesea triquetra. 
=Aulacomnium palustre. 
=Bartramia ityphylla. 
=Philonotis fontana. 
Pogonatum urnigerum. 
Polytrichum juniperinum. 
Diphyscium sessile. 
Fontinalis antipyretica. 
Antitrichia curtipendula. 
=Pterigynandrum filiforme. 
æClimacium dendroides. 
Brachythecium albicans. 
Plagiothecium Ræseanum. 
”Hypnum stellatum. 

3 — uncinatum. 


THE BRYOPHYTA 


”Hypnum exannulatum. 
z filicinum. 
cupressiforme. 
Lindbergii. 
stramineum. 


0) 


Groupk? 
Metzgeria furcata. 
Madotheca Cordæana. 
Radula complanata. 


£Rhacomitrium aciculare. 


OF ICELAND 633 
"Acrocladium cuspidatum. 
=Scorpidium scorpioides. 
=Hylocomium proliferum. 

z parietinum. 
squarrosum. 


i: 


Meridional species. 


Mnium hornun. 
Catharinea undulata. 
Camptothecium lutescens. 


Group 3. Boreal species. 


=Alicularia geoscypha. 
=Lophozia lycopodioides. 
5 Kunzeana. 
=Chiloscyphus polyanthus. 


"Cephalozia pleniceps. 
"Scapania irrigua. 
z curta. 





”Andreæa petrophila. 
=Dichodontium pellucidum. 
=£Oncophorus Wahlenbergii. 
G virens. 
Dicranella squarrosa. 
crispa. 
Dicranum Blyttii. 
Fissidens osmundoides. 
=Blindia acuta. 
=Distichium montanum. 
Tortella fragilis. 

Grimmia torquata. 
=Rhacomitrium sudeticum. 
5 fasciculare. 
£Amphidium lapponicum. 
Encalypta ciliata. 
Dissodon splachnoides. 
Splachnum sphæricum. 
vasculosum. 
=Pohlia gracilis. 
=Mniobryum albicans. 
Bryum purpurascens. 
lacustre. 

affine. 

cirratum. 


kal 


EJ 


Bryum pallescens. 
Duvalii. 
=Mnium cinclidioides. 
=Paludella squarrosa. 
=Meesea trichoides. 
=Catoscopium nigritum. 
=Philonotis tomentella. 
=Timmia austriaca. 
Polytrichum strictum. 
gracile. 
=Myurella julacea. 
tenerrima. 
Lescuræa filamentosa. 
Thuidium lanatum. 
=Camptothecium nitens. 
Brachythecium reflexum. 
rivulare. 
Plagiothecium pulchellum. 
="Hypnum polygamum. 

- revolvens. 
Kneiffii. 
decipiens. 
falcatum. 

SR ØVE giganteum. 
Hylocomium rugosum. 


x 


Eg 


i: 


Group 4. Alpine species. 


=”Gymnomitrium concinnatum. 
=Haplozia cordifolia. 
atrovirens. 
Lophozia quadriloba. 
alpestris. 


E5 


The Botany of Iceland. Vol. I, part II. 


Pleuroclada albescens. 
Anthelia julacea. 
Juratzkana. 
"Scapania subalpina. 


5 


41 


634 


=D)icranoweisia crispula. 


Aongstræmia longipes. 
Dicranum falcatum. 

: — Starckei. 
— molle. 
Desmatodon latifolius. 
=Schistidium rivulare. 

Grimmia Doniana. 

£ — funalis. 
Tetraplodon bryoides. 
=£Pohlia commutata. 
Bryum arcticum. 


The species to which an asterisk has been prefixed have every- 
where, or throughout the larger part of Iceland, the degree of fre- 
quency expressed by figure 4 (see p. 644), and consequently they 
constitute the bulk of the vegetation. Groups 1 and 3 inctude con- 
jointly 75 (41 + 34) species, while group 4 has 15, and group 2 only 


A. HESSELBO 


=Mnium orthorrhynchum. 
Conostomum boreale. 
Philonotis seriata. 
Psilopilum lævigatum. 

== hercynicum. 
=Polytrichum alpinum. 

— sexangulare. 
Amblystegium Sprucei. 
="Hypnum revolutum. 


— alpestre. 
£ — ochraceum. 
E — sarmentosum. 


one species which is very common everywhere. 





VBE AÅLETEIETUDINALSDISTRIBUTEON 
OF'EHESSPEGTES: 


HERE are only a few species which are equally common in the 

low land and on mountain heights. The majority of them are 
either Lowland forms, which rapidly decrease in frequency above a 
certain altitude and at last disappear completely, or Alpine forms 
which have their main distribution at greater heights above sea- 
level and decrease in frequency downwards. In, countries with a 
continental climate it is, as a rule, not difficult to draw a some- 
what sharp limit for the altitudinal distribution of plants, but the 
conditions for survey are far less favourable in a country like Ice- 
land, where a cold and damp insular climate prevails in the coastal 
districts. On account of the low summer temperature, the majority 
of the Alpine species can thrive as far down as the low land, and 
conversely many of the lowland mosses will be able to establish 
colonies in favourable localities, which are situated essentially higher 
than their usual limit of growth. In South Iceland, for instance, 
I have seen a southern slope, at an altitude of above 500 metres, 
which was covered with species of Hylocomium together with other 
species belonging to the same formation. 

Ås regards Iceland a fairly distinct boundary line may be drawn 
between the Lowland and Highland formations by taking the upper 
limit of the heather moor (and of the birch) as a basis for the 
classification (Helgi Jå6nsson, 1895—1899—1900); it is then seen 
that the upper limit of growth of a very great number of Bryophyta 
nearly coincides with the upper limit of the heather moor. The 
upper limit of the heather moor varies somewhat in the different 
parts of Iceland, and is also in a high degree dependent on local 
conditions such as shelter, direction of exposure, and conditions per- 
taining to moisture. In East Iceland the boundary line lies, as a 
rule, at about 300 metres, and the same is the case in the greater 
part of South-west, West and North Iceland. Around Myvatn, ac- 

41= 





636 A. HESSELBO 


cording to Thoroddsen's statement, the birch coppices ascend to 
550 metres. In North-west Iceland the birch coppices in Dyrafjérdur 
ascend to about 270 metres, and around Isafjardardjup probably 
not very far above 200 metres. In South Iceland the upper limit 
of the heather moor generally lies at an altitude of 300—350 metres, 
but in sheltered valleys both birch coppices and heather moor may 
be met with at an altitude of above 500 metres. On the whole, 
the upper limit of the heather moor may, however, be reckoned to 
lie, on an average, at an altitude of about 300—350 metres. 

All that is situated above this limit is in the following designa- 
ted the Alpine region, since the willow region which occurs in the 
mountains of Scandinavia and Central Europe is not typically de- 
veloped in Iceland; not, at any rate, in the coastal districts. The 
reason for this must in a great measure be sought in the geological 
structure of the country. The gradually-ascending or terrace-formed 
land near to the coast rises to a height of 200— 300 metres, and 
from here the mountains almost everywhere shoot abruptly upwards 
to a height of 600—700 metres, right up to the plateau on the 
mountain summit, from which again a few peaks or rock-masses 
protrude. This leaves very little room for vegetation between 300 and 
600 metres, with the exception of the valleys, which are, as a rule, 
very narrow. This is very decidedly noticeable in North-west and 
East Iceland, where almost everywhere along the coasts, from a 
quite narrow coast-land, the mountains rise abruptly to an almost 
constant height of 600 metres, so that only at the head of the val- 
leys do terraces occur at various levels as far upwards as the 
mountain heights. In North Iceland, especially towards the east, 
the country rises more gradually up towards the interior high land, 
and there — at any rate somewhat inland — willow coppices and 
willow swamps occur also, but their Bryophyte vegetation has been 
very little investigated. 

Taking the above as a basis the vegetation may be classified 
into zones, according to altitude. 

A. The Lowland region which reaches to the upper limit of 
the heather moor and the birch. 

B. The Mountain region which includes slopes and flats to 
a height of about 600 metres. 

C. Alpine Heights (the Snow region) to which is reckoned 
all that is situated higher than about 600—700 metres. The 
vegetation there is very poor in species, and is strongly affected 


THE BRYOPHYTA OF ICELAND 637 


by the severe climate and the snow, which remains over ex- 

tensive tracts till late in the summer, and saturates the soil 

with ice-cold snow-water. 

In the following table the first column consists of the species 
which have been found in the Lowland region, the second column 
the species which have been found in the Mountain region, and 
the third the species which have been found in the Snow region. 
Å cross (+) indicates that the species has its main distribution in 
the region in question; a dash (—) indicates that the species in 
question decreases in frequency and extends to a more or less high 
level (or low level) in the region in question, and a dot (-) indicates, 
that the species has. been found only in a few places close to the 
boundary of the region to which it belongs, or that it has, on the 
whole, been found so scantily that no opinion can be formed with 
regard to its distribution within the different altitudinal regions. 


Table III. The Distribution of the Bryophyta in the different 
Altitudinal Regions. 

















E|s|g gg & 
2 |e|S ISls|S 
£Ull|s KEE 
BK SÅES Alikællke 
SØE Elske 
Aliceulariat'scalaris teter + + | == 
Hepaticæ. — geoseyphassmeenrer re Es are 
Sauteriatalpina rss sees . Eucalyx subellipticus....... + .- 
Reboulia hemisphærica ..... — Haplozia"cerenulata mee dL 
FEimbriarias pilosas ss . — sphærocarpa ...... > a RR 
Eesatellakcomeassrrre eee — — cordifolasrerseee + — 
Preissia commutata 45 2443: + — Pipåriasr sst FE] 
Marchantia polymorpha .. + — — atrovirenss re + | — 
Aneura multilidas ÆBLE — — pumila SES 
MOTU ON SE FA SELER gere —+ Jamesoniella autumnalis.... | 
Pella Neesran as + Sphenolobus minutus..... 
Metzseria furcatase sone — — polis . 
BlasTak pusle tret — Lophozia quinquedentata ... ++ | + 
Fossombronia Dumortieri,.. | - — lycopodioides . + | —|. 
Gymnomitrium coralloides.. + | — — Flærckersseesrk + '—+ 
— concinnatum  — + — — quadrilobaneres ++ 
— varians..... . — Kunzeanasssrsre ++! — 
—= revolutum .. "| - = barbariske —+ | 
Marsupella Funckii....….... al He — ven tricosateneer + + + 
— emarginata ..... SÅEDE — Wenzel seed + .- 


— aquaticars seere . — alpesmisk serne + — — 


638 


A. HESSELBO 


Table III. The Distribution of the Bryophyta in the different 
Altitudinal Regions (continued). 

















É É rE É 
HER ESIlES (ESKE ES 
Sls!s Is's!s 
's/7|2 |s| Te 
|=|S' 2 SEE 
| =D 19 | | DD |M | 
| | 
Eophoziakexersa treere SAG SAR Frullania Tamarisci ........ | | 
SER SLS SE du esser se (HESS snes) 
— Mullen 038 88 + — Madotheca Cordæana....... — 
— Hornschuchiana... + |— Eejeuneacavilobhasvsreen — | 
— heterocolpossse | — | | 
Gymnocolea inflata. ........ . Sphagnales. ER 
Plagiochila asplenioides..... + — Sphagnum medium....…... — 
Leptoscyphus anomalus..... | - — papillosum + 
Lophocolea cuspidata....... — — imundatum ses — 

— Minor sus SER + — Gravetikss 3.3 | 
Chiloscyphus polyanthus ... —+ — compactum...... AES 
Harpanthus Flotowianus…. "— — tres SENSE ++ 
Cephalozia bicuspidata ..... — — squarrosum ..... Fl | 

— ambiguaeserneee LE — fimbriatum ..... ||| — 

— pleniceps mee + | — — Girgensohnii..... + | — 

Ba r— medias. . | = Russowii... . 
Pleuroclada albescens....... | — + — Warnstorffii ..... + |— 

— var. islandica ... "+ — rubellum 538 -— 
Cephaloziella Hampeana ..../ + — — acutifolium...... — 

— rubel ser + -— — subnitens serene + 
Odontoschisma elongatum... "+ — Lindbergii....… . 

— denudatum .. | - — ripariumnms seer + 
— Macouni..... en — angustifolium... | + | 
Eepidozia=setaceass seere . 
Blepharostoma trichophyllum + —+ Musci veri, | 
Anthelia-julacea seere + — Andreæa petrophia seere dL UdE UL 
— … Juratzkana... .|—|+|—+ | Gymnostomum rupestre ..../+ | | 
PtilidiumSellkare esser "+ — - | Hymenostylium curvirostre . |—+ 
Diplophyllum albicans..... + | Ånæctangium compactum..."—+ 
= obtusifolium.. | - Weisia ;crispata sanne + 

Scapania subalpina......... Ea Re viridula SES SER + 

== remotarsserse —"Wimmeriana 3 334 AM 

— irrigua NSSS SAR + — Dicranoweisia crispula...... =- + 

== uliginosa SES —+ — compacta….. | + | 

— paåludosa 3: soen — Cynodontium polycarpum..." - 

== dentata Lr see — Dichodontium pellucidum,.. +" — 

— undulat Eee + — ÅAongstræmia longipes ...... + + 

— OURE KS ASERNES + — Oncophorus virens SE "+ + — 

— Bartlinsn seerne ve == Wahlenbergii…. ++ — 
Radulayeomplanatans seeren — Dieranellasquarros ars HL JE! 
Frullaniaudilatata seven . — Schreberr NDR ml 








THE BRYOPHYTA OF ICELAND 


639 


Table III. The Distribution of the Bryophyta in the different 
Altitudinal Regions (continued). 








| 












































HEE |E g|E 
"1518 Z|o|8 
'5'8|S 3!S|s 
ieL, = [lse 8] bel er et É em if 
Dicranellat erisp au RES ES HEE Torterra sis meer | TE — 
— cerviculata 7445. +- | Barbula unguiculata 338 +| 
— subulataksreereee +! | aar SNE En 
Dicranum fulvellum .....…. +  eylindric ae FE 
— Andersonii. 3% +| Eem ad opl asseer + 
— faleatum seere — |+- | Desmatodon latifolius ....…. RS 
— Bly RER -|+ + — cernuusserre ES 
= Sandet neo orkc BE EEN or ora ob ts ifo harer NE 
— Moler EOS — + — ub ST er SEERE + | 
— Bonjeanisses ser + Er subulatarsesseer +" 
— angustumn sense, + | — mur onion 
— majussmessessee + 4 URAS SNERRE + — 
— SCOpPAarUumE ene i ke == ST Faciphyla sner i 
— fuscescenssss. 5 +) — Schistidium maritimum .…. "+ 
— congestum 2... + + — — apocarpum kirke + +. 
— spadiceumsmsstee + | — sracile rss + + — 
== elongatum 1.0 LEE ba = confertum...... + 
Campylopus Schimperi ..... + — alpicolassmese LEGE 
Trematodon ambiguus ...... + — — v. rivularis|-H | 
Fissidens bryoides ......... + Grimmia commutata ....... ER 
— osmundoides...... + — Donianas ES ER ERE 
— decipienssrsemsese . — alpestriss Seere —+ 
— adlanthoides sr — — OVat aser nn + 
Blindiatacutaerse ESS +! — — ncunvaserese . | 
Ceratodon purpureus ....... ++ — — batens 4 rene —+. 
Ditrichumstortile MESSER == — funakisees SSER | + — 
— nivalesssssse: SE) — tforquatas ses SSSeR "+ | 
= homomallum ....!+| | Rhacomitrium sudeticum … | + + — 
— flexicaule SES + — heterostichum (+ —| 
Sælania cesla SNS se + — microcarpum. | - | 
Distichium montanum...... ++ — canescens....|+ + | + 
— inclinatum...... + -—- hypnoides…... + ++ 
Bryoxiphium norvegicum + | — fasciculare ... "+ +) —+ 
Pottras Hem EER … + | = aciculare….../+| - 
SR Jati fo akser noe : Hedwigia albicans.......... "+ 
Didymodon rubellus........ ri Amphidium lapponicum .... + 
— Fufus sees dd . — Mougeottii ..... — 
Leptodontium flexifolium ...| - Ulotasp hyle + | 
Trichostomum littorale ..... Orthotrichum anomalum....|—+| 
Fortellasinclmata eee ssese . — saxatile 55330. + 
—  tortuosa +|— — cupulatum ...|+| | 





640 


AÅA. HESSELBO 


Table III. The Distribution of the Bryophyta in the different 
Altitudinal Regions (continued). 














ålle |te Ege 
be) | bra so en | ble) 
Elske ek 
SEE SEE 
Orthotrichum rupestre…..… |—+ | Bryuminliginosumseeerrere + 
— Sturmiieke + FR Ha ARS Er re |. 
— Killiasnser + za Væneume 53. RAE JET — 
— Bly | > UTIDE gab ao ss . 
E lævigatum.... | - — ohne sides — 
Encalypta: ciliata ASS — 3" Heirratu msn RÅ — | 
— rhabdocarpa ..... + |— HR intermedium Re — 
= Conor me — pallescens ERE + 
Dissodon splachnoides..... "+ +. — … subrotundum ...- . 
Tetraplodon bryoides...... I+!| | —, capillare. ;.. J.E er J 
Splachnum sphæricum..... +! .- HF cæspiticium FEER + 
— vasculosum ..... "+ | + | == comens er SENER . 
Funaria hygrometrica ..... | + | | elegans Seer +|— 
Leptobryum pyriforme ...….. |+| | —… argenteum sider 7. 
Anomobryum flliforme ...… "+ RE neodamense ss | 
= concinnum ...! + RE Duval SEERE flg 
PlagiobryumZieriiskrkenee lap == "pallens se HH FÆNSEE SP | 2] 
— demissumss tænk | HE ventricosum FS + + - 
Pobliakacuminatarkrereer || | SEANAD GUT TEE Eg BO dDD +! | 
=—HHpolymorphake eee | =H| —T Fpendulum eee E= 
He er ER En re bre Brownie | 
SEER FANS SEE SEEREN + |— | Mnium hornum Eee + 
ru ONS aars Bb tal SN) + + —  orthorrhynchum .... "+ |— 
EU LW ESSEN | — —H rr serratum serene |" 
—HEcommutata seer — + + RE spinosum ser | + 
HE grace lis SSR EERER '—|—+ + rr undukatum RER "hære | 
Roth rener + | —… cuspidatum.........- "+ 
SE ten ufo EJE | smed FN REE + 
== gran difloraseeeer fr | Raft ner SES + — 
=proligerak REE | o == iSeligeri VSSE + 
Mniobryum albicans........ +++ FRE Sstellare ss ERE — 
Bryum purpurascens ...... "+ - —Hcinclidroides Fre + —. 
—RFlaeustre rear ERNE | + punter 5 (ae | 
Kar SEERE IE —  subglobosum........ + | 
—  archangelicum ..... I | Ginclidium?'stysum tre + + 
— Jorgensenli HAMRE 2 Paludella squarrosa......... + — 
ED nun I+| Amblyodon dealbatus....... ' 
— Bebus une | + Meesea" trichoides RE RErE + — 
Elsa din SES ER rigt stra SEE ER + 
—. Grænlundii…......:; "ER, Catoscopium nigritum ..... re 
—H HE calophylum RS Conostomum boreale ....... — ++ 






































THE BRYOPHYTA OF ICELAND 


Table III. The Distribution of the Bryophyta in the different 
Altitudinal Regions (continucd). 














E|g|E (Elg |E 
Sie FEE 
aleollle Fi KS ES 
SEE | |= 
|| | 
Aulacomnium palustre...... (SF sr Pterigynandrum filiforme … |—+ | 
— tursidumssrg UL dL Lescuræadecipiens rr ; ! | 
Bartramia ityphylla ........ + — — — radicosa mere HE 
Plagiopus Eder mee milke — filamentosa vs | + — 
Philonotis"fontana sees. — + + — Breidleri SE | -|+ | — 
— Arnelliissekiliee (læ — patens sær eje, 
— seriatar: ALEN — — Heterocladium ea + 
— tomentella ........ 1+/+-| - | Thuidium tamariscinum .… |—+ | 
Timmia norvegica 2. IÆ! . | — delicatulum ...... 14 
re RR aUS IEA ASE oe Eee, — Philiber see | — 
Catharinea undulata........ == — abietinum ts 03 "+ 
Oligotrichum hercynicum.../— —+ | — — lanatum see "re |— 
Psilopilum lævigatum....... + | Orthothecium rufescens..... "Ez: | 
Pogonatum polytrichoides... +. | == intricatum.... | + 
—  dentatum var. minus! . | | — chryseum . | + — | 
= urnigerum ses + Cylindrothecium concinnum. |= | 
Polytrichum alpinum ...... | ++ "— | Climacium dendroides ...… '+| 
= formosum ..... ' Isothecjum myurum........ + 
— grace sier — | — tenunerven re (HERE 
== sexangulare .... || — "+ | Homalothecium sericeum ... + —| 
råt = | | Camptothecium lutescens ... | | 
var. tenellum. |—+ | — nitens (JE dL 
= pilifer um AS | — Brachythecium Mildeanum.. | — | 
—= juniperinum ... (+ — = salebrosum.. || — | 
== strictum see + |— == collinum ...| - 
— commune... + — = populeum... + | 
— Swartzi ere "+ —= velutinum ..|+| | 
Diphyscium sessile 1.12. + + SE glaciale..... 1 ++ 
Fontinalis antipyretica.….... FEE) = reflexum sv. 1—] EN 
— islandica,. 7: = longipilum..| .| | 
= lonsifoiaree eg — glareosum ……. ||! | 
RE flinlensis rer er — albicans.….. 4 
— androgyne. = erythrorrhizon |+ | 
Leucodon sciuroides........ + FE; rivulare..... + | — 
Antitrichia curtipendula ..….. |—+ = latifolium... | ER 
Neckera complanata 48424 + Seleropodium purum ....... | | 
Myunelasjulacea SSR — Eurhynchium strigosum .….. "+ 
— tenerrimassmree — = diversifolium . | . . 
eskeninervosass sees + = cirrosum ss 14 
— st catenulatar si ilde. | = piliferum 5. "+ 
AÅnomodon viticulosus ...... — | = Swartzii...... 1 | 























AA. HESSELBO 


Table III. The Distribution of the Bryophyta in the different 
Altitudinal Regions (continued). 

















' Above 600 m. 





S!is's ERE= 
og o | bi= og 3 
SENSE 2) | 
SHE BE 
| 
REE SES AR TØR, . |) ENN ae eV res ENKE - å 
Eurhynchium Stockesii..... I+| Hypuum flicinum RR "+ — 
Rhynchostegium murale.... | + — Curvicaule Ske || —=—1 +] 
— rusciforme : — — decipiens "SE. — 
Thamnium alopecurum..... "+ — commutatum...... + 
Plagiothecium silvaticum .… |— — — v. fal- 
— Ræseanum …. |+- catum + 
— denticulatum. "— — mollnscunsske — 
— pulchellum .. —+ — imponens NE — 
— depressum ... + — Bamberger se . 
— elegans . — revolutum sms + — 
Amblystegium Sprucei...... + — — cupressiforme ..... — 
— fluviatile 252 . — hamulosum ....... - — 
— Serpens sr, "+ = callicbroum SSR + 
— Juratzkanum. + — Lindbergs "+ — 
— littorale rer mm — palusre rer —+ 
— compactum .. + — arcticumss sære . 
== salinum .….... + — alpestre SEERE + 
— trichopodium. —+ — alpinumskkee — 
Hypnum Sommerfeltii...... . — ochraceum ÆS3s — 
— chrysophyllum .... + — cordifolium 2.2] + 
— protensum STE — — Richardson + — 
— stellatum ss ÆRE + + — — giganteum ..... … |" — 
— polygamum ....... —+ — stramineum RAS + + 
— intermedium . ae En — sarmentosum ..... Ii 
— revolvenskerrsere + + = frifanium SEE EA 
— uncinatum see + + — — turgescenstreee | + — 
— Sendtneri 224 BSK Acrocladium cuspidatum.... |—+ 
— KnoeitimSSSE +. Scorpidium scorpioides ..... +) - 
— exannulatum ...... + + Hylocomium proliferum .... (+ —+ 
— — var. pur-| — pyrenaicum ...! - 
purascens. + /—+ — parietinum.... re 
— — var. Rotæ.|!+- — loreum rer … || + 
— — var. ser- — triquetrum .... |— 
ratum. | - — squarrosum ...|+ — 
— Fiuitans tos SEN: — = rugosum isse + 


v. falcatum. 





Table I shows that of the 424 species enumerated in it (the 
habitat of Sphenolobus saæxicola is not known) 416 occur in the low 
land, and that only 8 species (Gymnomitrium varians, Pleuroclada 


THE BRYOPHYTA OF ICELAND 643 


albescens, Scapania remota, Ditrichum nivale, Pohlia polymorpha, P. 
Ludwigii and Brachythecium glaciale) have not been found below 
the upper limit of the birch. 
In the Mountain region the number of species is as follows: — 
42 Hepaticæ 
5 Sphagna 
134 Musci veri 


181 Total species. 

Of these, however, 10 Hepaticæ and 31 Musci veri have been 
found only in the lowest part, at the boundary towards the Low- 
land region, whereby the number of species which in reality be- 
long to the Mountain region, is reduced to 140. Of these, 2 He- 
paticæ, viz. Gymnomitrium varians and Scapania remota and 53 Musci 
veri, viz. Ditrichum nivale, Pohlia polymorpha and P. Ludwigii have 
been found only in this region. 

In the Snow region the number of species is as follows: — 
16 Hepaticæ, 42 Musci veri, total 58, of which, 6 Hepaticæ and 10 
Musci have been found only in a few localities and cannot be re- 
garded as belonging to this region. In all, 42 species remain, of 
which, however, only 8 Hepaticæ and 20 Musci veri are fairly fre- 
quent, while the others have been found only in a few localities. 
The majority of the species belong to those which are of equal 
frequency at all altitudes, and only 4 species (Pleuroclada albescens, 
Dicranum falcatum, Pohlia cucullata: and Polytrichum sexangulare) 
can be designated true Snow-region species. 

The number of species decreases rapidly as the height above 
sea-level increases, and even at the upper limit of the heather moor 
the number falls to about 40 2%/, of the total number of species. 
Above a height of about 600 metres the number scarcely reaches 
1559 skorfallstherspecies! 


NOTRE HORIZONTÅA ESDISERIBURON 
OFF FEFEFSPECTES: 


R order to obtain a correct notion of the importance of each plant- 
species within a floral district it is necessary to know not only 
their distribution in the different parts of this district, but also their 
relative abundance (number of individuals) in comparison with other 
species. The following Table gives a summary of all the Bryophyta 
found in Iceland, with the exception of the species growing in warm 
soil, since, as regards these species, it is chiefly the heat of the soil 
and not the climatic conditions which determine their distribution. 
The frequency within the different districts is indicated by the figures 
1—4, so that 1 indicates rare or very scantily occurring species; 2 
those that occur here and there or only in a comparatively few 
localities and not in abundance; 3 frequent, but as a rule playing 
no important part in the moss-covering, or occurring only locally 
in abundance; and 4 the common and abundantly occurring species. 

The majority of the species are not equally distributed in all the 
different altitudinal regions, and the frequency is therefore reckoned 
relatively to the region in which each species is most widely distri- 
buted. Ås regards species which grow only in quite definite areas 
(for instance littoral species) the frequency is reckoned separately. 

The figures in the table are, however, in many cases somewhat 
doubtful, since many districts are too superficially investigated to 
allow one to form an opinion as to the frequency of the less com- 
monly occurring species. This is the case, for instance, as regards. 
the majority of the districts of East Iceland situated above an alti- 
tude of 300 metres, because in June, when I travelled through this 
part of the country, the mountains were still for a great part snow- 
covered, which in connection with continual fog made investiga- 
tion almost impossible. 

It must also be noted that the table is based only on the 


THE BRYOPHYTA OF ICELAND i 645 


coastal districts, and on the low land with the neighbouring heights. 
The high land of the interior, which is about 90,000 km”. in dimen- 
sion and, usually, 600—1000 metres in height, is from a botanical 
point of view practically unknown. Outside the Jåkulls it consists 
almost everywhere of gravelly flats which are practically bare of 
vegetation. According to Thoroddsen, however, there are in some 
places extensive mountain-bogs, but the Bryophyte vegetation of 
these is quite unknown. 


. Table IV. The Frequency of the Species in the different Districts 
of Iceland. 











[Dy N. NW. W. S: 
Icel: "Tee" "Icel. Klee HFed: 








Hepaticæ. 
ae Santerria alpint. essere] Ea ofte lg 

m | Reboulia hemisphærica ...…............ 37 
ate En DE Aria pos ae ee eee erne ms 1ke 15 

m | Eegakelkakeonie ass res ene År: 4 
El KR Brest. eO EA EA RE ESS ESS SE DE 1 2 de 
u | Marchantia polymorpha ........... Ade im 47 47 47 4" 
11) HR ANSER DISSE Sae DSE RE EST ERER 4 4 4= 4 47 

m EU Ear OSSE SST Sr 1 
u EU URO TS ae SEE 1 1 1 
US IEEE ANES Tan Se RE En Ree 4 AT 4 4 

was Mebzse rr bar ea Ea re Era 3 2 3 3 + 
us he Blasiay pusle ES ERE 2 1 

m EFossomberonia Dumo£rtieri 23.23. 1 
a | Gymnomitrium corallieides ........... 3 2, 3 2 
a — concinnaftum 733824. Er AE AE 47 47 
a — MARA BS es Sr onen dene 1 1 
a — FEVONE TEE le Sr SL 1 
u | Marsupella emarginata….... RENEE 1 2 
u = AHA GARE as rare es eyete å i! 

m — IS ILD SD RE DEEPER SEND ERE 1 1 
ud HVALER Lara Scars SEE eee AS AS 4 El 4 
b — BeoseYDhAES NE rs sa jaeerele mes 47 4 47 4" 47 
ANE Eucalyssubellipheustasess eeanee 2 ig på 1 1= 
b Haplozia'sphærecarpå 22242. v8e. SED 1 
a — Cordt ohan es Seere 4 47 4" 4 AS 
b ri para eres ag SNE 37 
a — ALKO EBST SEE eee Eee ejen es 35 BE 3: ag 3 
b == Ba RSS TESTS NER Eg 
m — EPE FNS TE SE NESS 1 1 
me Jamesoniellasantumnalis se say 1 


64( 


) 


A. HESSELBO 


Table IV. The Frequency of the Species in the different Districts 


of Iceland (continued). 



































E: NMENIW: Wi IESS: 
| Icel. | Icel. | Icel. | Icel. | -Icel. 
|| 

Sphenolobusimmmutus sk | 1 1 1 
— SAXO OLA FS PENSEL: 
— poltus ere FREE | li 
Lophozia quinquedentata ............ | 4" 4" 4 4 
— lIycopodiordes rt rReRERRR | 4 4 4 4 3 
— Flere ne RER 4 2 
— QUadclobar ss SSR ESS ETS SEE 3 2 2 REDE 
— Kunzeamasee ener ere 4 4 4 al JP mdt 
— barbatassinr derne Rn es 1 2 1 
— VER IDICOS ARE RER Ree 3 3 47 3 3 
Wen el SL tree | FæG 1 ig 
— alpestnis st eee | 4 4 ASE SS Øe4 
— OXCISAL EN SR SENERE SE | | SETE 
— SCHULZ ii HERRERNE 1 1 iv | 
—— Muller SES ER mer | 3 3 3 3 | 3 
— Hornschuchiana ere |Lg9 lg 1 URE 
— heterocolpos ........ RER 1 1 1 1 
Gymnocolea sin data eres NSSS 1 
Plagiochila asplenioides.....…......... 4 4 4 4 4 
Leptoseyphusfanomalus senere (et 
Lophocoleafenspid tamme | 2 
— Minor LE ES eN 1 1 
Chiloscyphus polyanthus…............ 4 4 + 4 + 
HarpanthustElotowianus mee LEES 4 2 
Ceplialoziakbicus pi data rer 4% | 4" 47 4" 47 
— ambiguarss seeren | leg 
— plenice ps sr enn 3 ege 4" 3% 2 
— MEDIA Kes Er ERE. Nie 1= 
Pleurocladatalbes cer see een ERR SNE MES 4 2 
= — var. islandica... UaeT 2 
Cephaloziella Hampeana ….….....:.…... 4" 4 4" 4 4 
— Tube ASERNE ER 1 el 2 1 1 
Odontoschisma elongatum...... ..... 2 1 3 1 
— denudatom er ener 1 
— Macounir serene BA  beby] 1 
Eepidozia setaceammeeesne Eee | 1 
Blepharostoma trichophyllum........ ALA gs 4 4 4" 
Chandonanthus setiformis?...... 
Anthelia' jula cesser ses ae Så Se BE 3% SE 
ENA BYTE LES SE ao da don dede SEA Eu Is AS 4" 47 
Piilidiumtelrane me eres ASER 4 4 4 
Diplophylkum tab icars eee 3 3 
— obtusifolium FNS EEK 1 1 


THE BRYOPHYTA OF ICELAND 647 


Table IV. The Frequency of the Species in the different Districts 
of Iceland (continued). 





KERES KEN: NW. W. S. 
Icel,. | Icel. | Icel. | Icel. ""Icel. 

















| 
a Scapa des ba sa REE IN ÆRE? ER DÆRE AE 47 4" 
a | = FOO SE ele arne ets jers le re | as 
b | — ITD UAE NE AE RENS SENER REE: CA GARN I 4 4F 
b — Ulreinos aL SS SE SIS KE TØR) | 3 
b | — paludosass ERE REE | | TE be 
u | — (entakt ST EAR | Eben kalt Kg BG MBE 
u | — (are ad FSR or Eee Es ES enge 87 Bed KE SAN ek ST Be 47 47 
b — ENE ea ER SE ra Et | 47 4 sg 
a | == Barks ER SESSERESE | 1 
si (tadnlas com plandtas ser SS ISS E.SE 3 ole 3 BE 
NE DT Er ED ENE 2 2 3 4 
re || REE ENE EN ER Se se RE 1 
u | = arm anis les 4 4 i 4 4 
m | RE fra silfo ae eee RR | Mar 
mi Eejeunea canon: sees ee | 1 3 

| 
Sphagnales. | 

bi HE Sphagnum medium SEE SELER 1 1572 
m — papillosum es nere RG | Sa RRS 
u = undt eee eNRER | ter 
u == Gravetiks isse neserÅ ds | | (LSE FISSE) | 
b = Compac hun seN SES ESSEG | | 1 
u = Ferestre REE ns eres 4 4 | 4 4 4 
u — STUAFTOSUumM ELERS: 1 3 
u = Da 9 59 2 1.95 12 7 BERN ERE SEERNE ER Tr AO SE 2 1 
n — Girgensobnii rss REE RS 23 3 35) SEERNES 
hb — BUSSON ES EDER etat | 1 
u | — Warnstorthi FE Sar 3 3 3 3 3 
u — Fab SEENDE 3 3 3 SR 3 
u — EU ON ONES SST Ta HE ES 3 
u — Subaenss DS SS REESE 1 2 2 
b == nr dere sen seee | 1 
u — Pipin Sr EVER RE | tr 23 
m — angustifolmmee EEN: | bl 


Musci veri. 


Nr Eee EDIT EET ry 4 | 47 
KN Gymnostom um rupestre rs ELLE Be ber sbaE > | ØRRED MERE 
b | . Hymenostylium curvirostre.......... | | 1 | | J KYESR Ma BE 
u Anæctangium compactum............ [va SÆT ae 
m | Weisia crispata ..............22414420 1 SA SE 
m | RTE 8] CY Eee SEER Sr rs | | rs: 





a | EWS sne ke så is 


648 


A.-HESSELBO 


Table IV.' The Frequency of the Species in the different Districts 


of Iceland (continued). 














ES JANS NNWA HRWSR ES: 
Ice Ælcels Ice her cel reel 
| 
a Dicranoweisianeris pulsen AS AS A= AES NESAS 
a — Compac ta sen SER 1 | 
u Cynodontium polycarpon ............ ik 
b Dichodontium pellucidum.........…. 4" AA RAS 5 BREDE 
a Aongstræmiatlongipes eee 3 BEEN en] PAR NT 2 
b Oncophorus Wahlenbergii............ AG: 4 4 4 4 
b | = NIFENS 3 He eee Er 47 Ass ASE HRRAS | 47 
b Dicranellats ag mabr os are een 2 Sr 4 2 1 
u | — Sehr eD SEES SST ERE i 1 | 
b — CIS PAR Ane re Re rr 4 4" 47 ASE UREN 
b = SU ua bree e EEE 1 1 løser 
u | — Ceres seere 2 2 1 
an Dicr annum ulve um SNEEN | 3 
al — Anders OEEEE E | NE3E 
a — faleatum ser 2t VE 327 | [92 SE 
b == Bly ti se EN NDR SE 1: je 3% Sl LIGE 
a EN ES OG MURERE NS He SEN 24) GEN BÅS 
a — MO ESS ESR ii ike 4" SE TE 
m — Bonj sanse REESE il i DE 
b — anus bumser ses 3) i LE SED il 
u — MajuSs ERNE SE SS SE Sats : ON ET] | 
u — SCOpPanumeerEre Er eer SEES 4 4 47 ASERNE SE 
u — fuscescens ESSENS 47 il 
u | — Congestum rss re d 1 EN s8) 2 
a | — SsPadiee um 2 1 het 
a | — elongatum ke see 2 2 2 
a Gampylopus"Schim perkere 1 1 (va 9 1 
m Tremafodontambiguus ERE REESE 1 
m Fissidensibryordes eee iz 1: 
b — osmundordes eee 3 3 3 BE 3 
u — adlantholdes eN SRESN SE 1 22 
u — (decipiens sy EEN SE 5 i 
b7|» kBlindiacuta ESS SET AE SEE AT SAGE | 54 REALE BEAS 
u | Ceratodon purpureus .….............. DE BØ BE BLD BOE 
u Ditrichum tore re ener il | 
a == Mivale EEN lelele« 1 
Uu — homomalumsmmærrsssser 1 1 PS NE] 
u — flexicaule SS SEA ASE 4 4 ASER RER: 
a SEA OB ST Eee 2 25 37 DE 
b Distichium sm on tan unsere NER AS: 4" At 4 47 
u | — inclnmatum see RENSE 1= 25 TEN 
a | Bryoxiphium norvegicum............ | 3 
u7|KBottiad Heim ege Ree SNE RE TØNE Pe: EET SE DE 


























THE BRYOPHYTA OF ICELAND 


649 


Table IV. The Frequency of the Species in the different Districts 
of Iceland (continued). 
































N. NW. W. S? 

| 1 Icel. |. Icel. | Icel. | Icel. | Icel. 
a Potato has ER SEE SEERE 1 
uw Didymodonrtrubelas AA JELREN ER 47 AS AG NE SÆR NL 
b = 1 RS re re EL DEERE 1 i! 1 | 
m Leptodontium flexifolium ............ (ni | | 
m Eriehostomumslittorale se ERE 1 1 
ut FE Kortellatin en atase  ESELTEEE TEN | 
u SEER FORE OS AN EDER ER er 4 4 AEG NEVÆ 4 
b fra sli s eve sas Tr SE ev IRIS SE 3 3 3 Bakers 
m Barbula unguiculata var. cuspidata ... 1 1 
m = LE SER ES SEE SEE ere 1 1 
m SE SEE ERE 1 Babel 37 
a — rem adobe 1 IE oo 
a Desmatodontatifolmns ære REE 3 PS dr DE 
a — CEED USE re ere eee IAN) 
a Kortularobrusito las Eee | ig 
m IE TE FSR REE TERESE SEERE ks 
u | ES RS eee ere nd DE 3 age 35 BE 
m SR muerto ESS E ig 
u SEALS ERE areas SR verde, DE år BE 2 2 
a Er acip ya SNE, SER RED RR: | 1= 
u SIS mar IEEE 32 3% SE BE DE 
u — apocar punter ere 35 3: 3 Båg 35 
u -—- BEA een (KS ÆS AT 45 4 ÆRE 
b — Con fertum ss SIDES: |Vss bg BE 
a — ran] ES er SES Es | | 1 
al] — — var. rivularis..... | AN NÆS 47 47 3 
b Grimmiat commit taken SEERE li ere Ål tapet 
2 — Bonar see nesker ass ES 57 MENS is 
a | — alpestrisere ae r 1 (Sv 
b | — ONE ER DE SE SEE 2 DN) | 
al — NE Er Er i | | 
u = Batenss SEE REESE 1 1 
a — ben alS RE SDEVS ARD RESG 4" 47 47 4E 4F 
b — borg rat Sele trelee 2 1 2 3 4 
b Rhacomitrium sudeticum ............ 4 Sk AS BE 30 
b — HALE KOTE hu ons 1 DERES SEES 1 
u — heterostichum......... 3 FYN 2 4 4 
u — CAHEeSCEnSE ert eee 48 GÆR URAS 4 4 
u | == Hypnordes tres | an hg | 4 4 
b | — faseiculare tre KÆR 32 SYÆN FEG' hg 4 
m — ACICUIAFEr snore erne ze 42 4" | 4" 4F 
m Hedwigiatalbicansns JENS NEDE: | 1 
b | Amphidium lapponicum ............. 4" 4 is 4 Aa 

The Botany of Iceland. Vol. I, part II. 49 


650 


Table IV. 


A. HESSELBO 


The Frequency of the Species in 
of Iceland (continued). 


the different Districts 





| 
| 
I 
|| 














| 
FÆR EN: We RES: 
| Icel. | Icel. | Icel. ! Icel. | Icel 
| | | 
Br SE gg | 
u AmphidiumsMougeosireeesreeeesrer Væg 2 3 4 4 
un KUL Ea p hy ant ae SENSE 3 3 3 3 3 
m Orthotrichum anomalum............. | ib 
m | — saxatile ene ER Nr: | 1 
m — Cup Ulam FE ERER ' ig 
u — runerne se SE SENE SEER 3 3 SE 
u — SUT IE  S ES SENSE lig "vens 
a — Killiasii eres i ER 2 
a er Bytte Son SERR DE SY LSTN ME KDRTS 
a = lævisatumssersersseee 2 ile ube 
b Encalyp tar era tuen ES (5: BE ler 8 
u — rhabdocar parere 2 | SE SER HER SE æg 
u — Contor kasse ASSENS | sen 
b Dissodon"splachnordes ere eee 18 i SE lee ile 
a Fetraplodonfbryond es ere seernes DE DE DE Dj DE 
b Splelmumes p hære ure 38 SE De DE TEINREDE 
b — Vase os ME ar SE BE 3- FBE 
u Runarrakhysrome mia 35 SE Så ore || 83 
u Eeptobryum pyrtor mere AS 4" BERLT E 4% 
a Anomobryumkfiliform emner 2 13 SE 
a — COII UMS EET RER | 1 
i Dael omuen ØGE odd nes one EN oDE uk er 1 OMEN REDE 
a | — demissum eee læ 
Pohlhatacumin ate eee | is 2 
a ED OL OT PARRE EREREER | 1jER 
u SE ord ENES SEES BE Aron HE 45; 4 Ås ASE 
u ED VUE VINTER OSTE ERE RE BRIDE terra ls ilke SE BAG) (aA 30 
ES DN i EN > ENG SER TERESE SOREN ESKE ER ESS | | il GE | | 
a FT HE dwi SES  E RS RE ES DRE SØN | is | 
a rr KOOL S EN  n e AeR | 4 ARE RE 
b == dgraeilis er er KS SER DS se I et Er SE MED 7 
b => BROEN SS ze NÆR BRENNER: | 1 | | 
b SE Sl Eee 2 2 ea 
b pro ger ASE El E | | Jl 
b == eran don SEE ER Er | | 12 
b Mniobryumfalbreans eee AE 47 47 As de 
b Bryup r Uraseess 2 BE DE REM KG 
b == lacus tret SEE SURE ANSER 27 BE ME As DE 
a ER SES SE Eee sr ng i 
a | — i Harchangereun rer | | alkg ale 13 
u 7 nemt um IR HEER ER SØS HDRA SE 4 AE SA NDDNAE 
a EA ST SE SEERE 0 ERE |. ox als 
Icel RE FIS ATT am SE gr RR SER | le 

















THE BRYOPHYTA OF ICELAND 


651 


Table IV. The Frequency of the Species in the different Districts 


of Iceland (continued). 















































| 
EEN ENNA As: 
ÆT cel st Rrce LE ElcelS STcels cer 
b Bryumscalophyllum ss SEERE RENGE |SæR1 
u R uliginosum INSSE VER SERR | DS lig 
Icel. | SE Gr æn dies REE SER | hi: 
b ES EVE SAGERNE ss ER SER ESS ER alle | 
a | KODE TEE EET SETE Re EGE 2 1 il | ibg BE 
m DIN UNE SE ne GR Ree Eg | 
b SE AFTER EER FLER INSEE SENSE ae ES Sl sk Bå= 3: BE 
b | EEN DD ØRER ES RER REDDER ERE er PE 2 SR Påg ikke 
u SE Santerme drum HS SDS ore ile ilkg 
b SR pallese ens oe Snes || 32 SE 3 32 AF 
al  subrotunduns AES SEEST 1 
u ADA SEE eee 1 i i 2 
u  <cæspitiolume aser eee BE USD ES NDS flig 
a | ODENSES abe bon beo Do Go ra 1 
u | SEEST NE 1 AE ENE EE  E RR |lgS 3 3 3 3 
a | — … neodamense var. ovatum ..... | 1 
D | ED Sl DYNEN MOLS VERS BE ERE RØRE Farens se slet | 40 rs 4 4 4 
u | SE palle ns es ERE Se Nee ere LE 3 3 3 4 
u | ventil cost me ERR es | ØE: 4 eg 4% 4" 
a | St Fare ton Se ERR SE RER less 3E SE ES DSE 
u pen dulu ms SS ae | lg ig DE 
a OSS Sne Doro eo bro Glare | lg lig 
a | RENT Or gense se REE ESSEN | 16 
0 NEN KOSTEN So 00 so arge or SETE 3 2 5) 3 4 
a —R Forthorrbync um SSL NE 4 då 4 4 
m | ES OD PS UI Er Re eee | i DE Så 
ue FEE SPIN OS UN ER EDER ore eee | 1 
m | and ul fu ER REESE | aå 
m | > cuspidatums Al serende 1 1 
b DU re ea ar 1 
us) EU NNE ES øne ED ne SERENE SEE SEE 4 4 4 4 4 
u | =HREKS STS ERIE ae RE ere GORE RENE 3 2 2 4 
m | RE Stean een Ro Sens 1 ilkg had il 
b | Rend older ESSEN ER | 4 - SR 4 
u RED un etat um SENER SE "EG EE0 HEY es 4" AE 4 
b HEE Subslobosumkneere rese ng | | Pir leg 1 
b Cm clid ramt sty gum ENE EN NS | AES RGS Bk 3 30 
DN ER Paludelaksg arr os ar ER SSR | Fe) hger,: 3 3 2 
b Amblyo done deal aus rese | æg 
b Meesea tvichoides ES syd NS ERE (FRE Sy | 4% ÆE AS 4 
u | ES TU EEEAEE n reeeemedeNet es 3 3 2 2) 2 
b Gatoscop rum kni grun se EEN AES Uni 4E Dig 47 DE 
al | Gonostomumt boreale ASSER BE 3 AF ES E FE 


A. HESSELBO 


Table IV. The Frequency of the Species in the different Districts 





of Iceland (continued). 








| Es UN UNW Lowe MAS: 
| | Icel. | Icel. | Icel. | Icel. |! Icel. 
u Aulacomniumspalus mes SEES 4 4 | ASE GEA 4 
a — tursidum ss >, 2 2 l i 
| Bartram ra rtyphydla Free AS; 4 4 AS AS 
ba EPlasiopus kædes SSR SEERE Er "ler 5 it= 
u Philono tis fora are ere 4 = 4% 4" AS ASE 
hb — Fald hf) USE bed SØDE SSI SBS See na | i 1 al 
a — SE i ENE REESE SISSE SEERE FEE, 2 BE "SR Er 1 
b — forment elser | gx ASE RES 4 At 4 
a Fimmiarnorvegieat ER EEG 1 æt) 1 
DS re aus SNE aen eee a ere 4 ÅSE 2 4 4 
m Ga than eau due Pis 1 |SRSE 4 
a Oligotrichum hercynicum...…........….. AR DE: files | Peg BE 
a Psilopilumlævigatum eee 35 SEES SER SE 3= 
m Pogonatumipolytricholdes seeren | DE 2 
a — dentatum var. minus..... I's 
u | — urnigern meeen 3% Se BEN 3 35 
ad UR Polytrichbum alpin um kr SEERNE 4x AE  LARER RE 5< AS 
m | — formosum seere | i 
b | — Sraciler sa NNE BE BE SEE MRE38 SE 
a — serang ul are renee eee 2 3å Mrk 2 3= 
u — Piliferum EEN lg DE at SES DES EROS 25 
u — juniperm um er Æ3> 3 | 3 ES = BE 
h == Sme bun NR LE SR 2 SE EET 52 2 
u — Com mune ERE RR 2 SETE NS 82 DE D 
b — Saba SERIENS ERENSS Rk 
b — — var. nigrescens .. vs 
u Diphysciumfsess lee eee 3 be IN) 38 Bø 
u Fontinalis antipyre henses 3 3 | 3 3 3 
Icel — FAN IC FE ERR SEEREN, 1 | 
Icel — Lonsikolra rese SS PÆg 
Icel. == thulensis rese re il 
b | — androgyne eee ERR ig ik 
mi euncodonsciuroides FEER 1 1 2 
u Antitricbrascortipenduasrtkeeseeer 2 2 2 3 3 
m Neckerarcomplanats SES | 1 2 
b Myurellatjulace as NSSS LÅSE 4 4 4 4 4 
b | == apIc Ulla aser eee Er 3 SEES 3 2 
bå es eros |lgen ! 1 
u ER Fe afen later SEE ENE ERE ) (id 
m Amomodomkvr tre nlos SEE Er | 1 
u Pterigynandrum filiforme ............ 4 4 Mm dl 4 
a Leseur eamdeciplens ss ERE il 1 
a — radicosarss Seere referater 1 





























THE BRYOPHYTA OF ICELAND 


653 


Table IV. The Frequency of the Species in the different Districts 


of Iceland (continued). 


























E: N. NW. | W. S: 
(tlcer| Fedtet" "Teds KTeee 
b eserræattilamentosats . ERE Re 3 3 47 3 3 
a | — Bredden ES FK SOS E 1 2 3 1 
b | — Patenssymrs mee 1 j lGd PE 2 jl 
b Heterocladium squarrosulum......... 1 il 1 1 
m Kande tam ars ci  SSSERE 3 
m — deliea tun SES FSR il 2 3 
m | — Philibertis SF SKEER il 3 
b — lanatum reen se NS SI EG: 4f 45 DE 2 ile 
a Orthothecium rufescens ………...…...…. 1 
a — mtricatunmtss ss ERE ss 1 2, 
a = ChryseumnsANSS SE SEE 4 4 2 2 
m Cylindrothecium concinnum ......... 1 1 
u | Climacium dendroides ………..….......…. 4 4 4 AG 4 
ms HI solhedu mm yuru MESSSSESESSEE 1 2 
b | — tenumerves SÆæSeRE 1 1 3 3 
ul Homalotheciumsericeum ESME DER 4 4 4 47 
mi ME Camptotbeaum ntescens sees ES 2 2 2 z 3 
b: | — milten se Ra SE 47 4 47 47 BE 
ur UN Brachytheciums Mildeanum SE SEE il 1 
w| — salebrosumn skeer 2 2 2 2 Pr 
a = Column SS TEE SE SS il 
Icel. — Fonsikomumes Es il 
HR | — populeum SEE ES 1 
u | — velutinumirt 33835 2 
a | — Places SNrSEaa 1 1 1 
b"| — reen rer se BE 3% Så 35 BE 
Bd == plareos mn SEE 1 1 1 
u — Albi ans ERE RE 3 3 3 3 3 
b. | — erythrorrbizon ss sss,. 1 1 
b | — Livia syer SKE SER ser 4 4 4 4 4 
b | — latifokunmnmssst SØER I! 
m | SCLeL OP OTTE PEERS ER ENE 3 
mUMSEurbynchiunmstrigosum SSD, i! 
m | — — var. præcox.. 1 1 1 
a | — diversikolummres SILK 1 
a | — CILTOS HER 132] SISSE Sr 1 
m — pilen ter SENDER: 2 3 
m | — SEE VA KE ESEE ESS. SES ERE 3 
m | — Skockesie, es ae 2 
m Rhynchostegium murale ..........…. ifsg 
m — ruscilorme…s isse: i 1 2 Bj 
b — —  v. atlanticum 2 1 2 
mi Ebam nun kal peer um EL SSSRERNSER 1 3 


654 


A. HESSELBO 


Table IV. The Frequency of the Species in the different Districts 


of Iceland (continued). 

































































| EN Nw wil es 
| Icel Icel Ice RICE Ælcel 
| 
u Plagiotheciumsikvaticun seere | ml | 1 
u — Ræseanum seere 3 3 3 BTS. 3 
u | — denticulatum sees | 2 1 1 1 
b — pulehel um eee "RESEN BESES IT EA ESS RE 
m — depressumer ere ii | 1 
m — elegans SSR | 1 
a AmblystegsiumFsprucer Ses sene 15553 3 3 3 3 
m — fluviatile SERENE | | 1 
m — serpens: za Se RE | æn iz 1 
m — Juratzkan un ere | UeES 155 
b — Littorale SES SEER tre 1 
hb | — compachum seere AR Sl 
b | — Sal unsere SER EER | I 
m | == trichopodium .. | 1 
mi EH ypoaumtSommerfeltii sr | |egt 1 
u | — chrysophyllumreeeeeer | 1 | 1 1 1 
u — protensum BESS ER Er |. 2 [5 2, 
u | — stella bumser herre SEER | ASA 4 A's 4 4 
b — polygamum rese 4% 4 BE 4% 3 
u — interm edium ses 2 3 2 GE Kran 
b | ze revolver seere RER 4 SER SAG eg AE ES 
u — Fe 4% ASE EN 4 ASE EÆS: 
m — ven diner se DERS iL ES 
b Er NS II SE ER SEE EOS ER 3 FBE 2 2 2 
u — exannulatum eee eee | Ms 4" 4 3" 
a | — — varrRotæe sier > | 1 
u — fluitans see ENE SE Er« | 4) 1 
a | — vans falcatum str | 2g 
u | ERE licin un SENSE RER SE | op HI fB FED FEER SAMS AA seks 
a za curvicatle So RSS REE | DE il 
b — decipiens Senere "KØGE: 4 1 
m | =— commutatunkresrserksrnerr "gan KT NER ak RR2 ) 3 
b NE ES BERNER OL Ug | 42 AES NA Ale RAS 
m — mollus cum sees NEÆL BENE 
bå — imponens sees esse VRE 1 
a | — Bamberger se ES. | 1 
a — revolutunss esse (4 4 4 EEN KR 5 
u — cCupressiformesskes ste L.9 IT R2 GØREN RF: 
a — hamulosum srt ASSER | TØNLEN 1 DE LNG 
a — calliehroumns rese læ ad BE 2 3 
u = Eindbers ise RR | 4 | 4 4 Me 
u | — Palustr er DSN NE SDS | alig | PER DSR EHESE 
a AO NE as ED Ea VE | | 15 æn 





THE BRYOPHYTA OF ICELAND 655 


Table IV. The Frequency of the Species in the different Districts 
of Iceland (continued). 


























| | 
| EEN ENE | wil ks: 
| Icel. | Icel. | Icel. | Icel. | Icel 
Å | | 
= | 
a Elypnumtalpestrese ER Re ILERBrs | 35 BE DESIRE DS 
ag | — APN RR HEE | SS RESE, 62 | 
a | — OCDR ACE UN MERE ER Sr | 4 hæs SR 4 4 + 
u — Cordt E SEDSEEREE | bug: ile | 
b — Richards oner | 25 AR 16% get 
b | — giganteu mere RØN Ear | AE REE |: hst ES Sae NED fa 
u | — skramme SERENE REE FE: AR NÆ 4 "ol ES Æ 
a >= sam entos um see (1.53 | SON RS 3 3 
b — tran um SØS SER es | I hg 9] | | 
a — furgescensks ten ea Tr Ia ET SL oRSÅ | areal 
u Acczocladiumfcuspidatum skere se | AE GES STATEN SSSRAGE SS ANA SS 
u Scorpidiumsscorploides RESEN 4 EO sg 43 KE EMSÆ 3 
u Hylocomiumiprolifer um FSR ER RE | 4 
b — pyrenaicum es KÆRE | "et: 
u — parietinumsrereee EEN mee | 2 | 3 3 3 
u — Loren SEE ERE | | 2 3 3% 
u == trigusetrum skere | 2 | 1 2 3 3 
u = SQUarrosum SSI SE Eee | 4 AS EA: 4 4 
b — BUSOS UNE S REE PLÆRS m3 | SE HS 3 5) 
| | 








Of the 93 Hepaticæ, 20 Sphagna and 326 Musci veri found in 
Iceland, 7 Hepaticæ, 3 Sphagna and 4 Musci veri (besides 3 varieties) 
have been found only on warm soil. The other 425 species (87 
Hepaticæ, 17 Sphagna and 322 Musci veri) enumerated in the above 
table can be arranged according to frequency in the following 
groups (Table V): — 

a. Dominant or very common species, the frequency of which 
is expressed by the figure 4 in all parts of Iceland or by 3 
in only a few parts. 

b. Frequent species, the frequency of which is expressed by the 
figure 3 in the whole, or at any rate in the greater part, of 
Iceland. 

c. Species of which the frequency is expressed by the figures 1 
å 2, or which have a higher degree of frequency only in a 
single district of Iceland. 


656 A. HESSELBO 























Table V. 
— == = == 
Group a | Group b Group c 
ra få SES SE | FE= dk 2 
Hepaticæ 5 SES ÆSEES 23 | 14 49 
SphasnassSEEEER | 1 | 3 13 
Musi veri ESS AEE | 64 | 82 | 176 
Total | kv tee NEN 238 
20.7 2/0 23.5 0 | 55.8 2/0 





Iceland, according to the climatic conditions, may be divided 
into 5 districts, which division, also as regards the Land vegetation, 
coincides well with that employed by Helgi Jonsson (1912, p. 5) 
for the Marine vegetation along the coasts. 

East Iceland, from Eystri Horn in the South-east to Langa- 
nes in the North-east. 

North Iceland, from Langanes to Cape Horn in the North- 
west. 

North-west Iceland, from Cape Horn to Låtrabjarg. 

West Iceland, from Låtrabjarg to Reykjanes. 

South Iceland, from the western point of the peninsula of 
Reykjanes to Eystri Horn. 

The boundary lines between these districts are in most places 
formed hy ridges or high-lying mountain plateaus. In the South- 
east, Løénsheidi (height about 400 metres) and Hofsjoåkul form the 
boundary between South and East Iceland. West Iceland is sepa- 
rated from South Iceland by the mountain-chain which stretches 
throughout the peninsula of Reykjanes, the lofty mountain of Hengill, 
the lake of Thingvalla and the mountains situated north of a line 
drawn from Thingvellir to Geysir. Between North and East Iceland 
there is no sharp boundary line, either as regards climate or geo- 
graphy. South-east Iceland (from Seydisfjéårdur southwards) has a 
very damp climate with many foggy days (near Berufjérdur on an 
average 171 annually), while the western part of North Iceland (the 
districts about Eyjafjérdur, Skagafjérdur and Hunafl6i) has a greater 
difference between the summer and the winter temperature, con- 
siderably less precipitation and comparatively few foggy days. North- 
west Iceland, which comprises the country around the deep, narrow 


THE BRYOPHYTA OF ICELAND GHSØ 


fjords which open out towards the north-west, lies rather isolated, 
separated as it is from North and West Iceland by Jåkulls and 
lofty mountain-ridges. The boundary between North and West Ice- 
land is rather undefined. Several species which are common in 
West Iceland, but rare in or absent from North Iceland, are met 
with right in around Hunafléi. Here the range of hills east of 
Blondudalur appears to form a natural boundary, since Campto- 
thecium lutescens and Rhacomitrium heterostichum, for instance, have 
not been found east of this line, whilst other species common in 
North Iceland, for instance Orthothecium chryseum, Hypnum decipiens 
and Thuidium lanatum, are rare west of it. 

Of the 87 Hepaticæ, 17 Sphagna and 322 Musci veri enumerated 
in Table II, 30 Hepaticæ, 4 Sphagna and 108 Musci veri (or col- 
lectively 33 %/,), are almost equally frequent in all the districts of 
Iceland, the frequency being (with a few exceptions) 3—4. The rest 
of the species can be grouped as follows: — 

1. Species with a mainly Southern distribution. 

2. » » >» » Northern and Eastern distribution. 

3 » SE > » Western distribution. 

4. Species which occur sporadically in several of the districts of 
Iceland. 


1. Species with a mainly Southern Distribution. 


a. Found only in South Iceland (the figure denotes frequency). 


Meridional. Seleropodmumfpurunseee ED 
É ; : . Eurhynehume svarer SED) 
Kebouliat hemisphærica ss ki 3 3 Rg z É 
g -— Stoekesmssr genre 2 
Hesatellagconre arsen d % 
z Fe Rhynchostegium murale ....... 1 
Fossombronia Dumortieri...... 1 So : ) 
; Plagiotheciumfelesansmeene 1 
Aner am ulk fla eee i 
5 É = depressunsserer i 
Jamesoniella autumnalis..... asked 
Eophoziat excs se Fee eEns 1 5 
Å Boreal. 
Eophocolear'cuspidatak serene 1 
Lepidozia setacea..... fr REE (AH a plot eri para eereeer 3 
ANE TSA ae SES phagnumeom pa em 1 
TORRES SEE 1 Pohlia grandiflora Setaleet skele SNS 2 
rs mucronifolia........ HPA mm bhyste sum san 1 
TE EINGENS Ure se eD DØDE 1 Hylocomiums pyrennicun ke 1 
<OrthotrichumFanomaum ss SS nnen 
— saae Ses | SS ADIDIE: 
== CupulatumsessE fæ Scapaniak Barti nose 1 
Main un dut un ERE SED jer anni kuvee 3 
Anomodonsvitreulosus ss Bryoxiphiumnorveg cum er 3 
Rudme fam arsen dm ESS ET or tu ob sko rr 1 


658 A. HESSELBO 


Ånomobryum concinnum ...... 1 Icelandierspecies 
7 O g rå ØRNEN SSR IK sS es re = - - = 
Bryum Jårgensenii.... E Brachythecium longipilum ..... b: 
Ubiquists. Kontinalsætnulensis seere 1 
Brachythecium populeum...... 1 = Tonsitolreerreere æ 
= velutinum rese 1 


b. Found in South Iceland and in the most nearly adjacent parts 
of West and East Iceland. 


SR WIDE. SYFWÆ3E 
Meri'dional. Hypnum im oluscurn rese 

Marsupella Funckii ..… 1 1. Thuidium Philiberti..... om l 
Frullania fragilifolia .... 1. 1 
Eejeuneaf serpyllifolraereres el Boreal. 
Sphagnum papillosum... 2 2 Fontinalis androgyna.... 1 1 
Weissia crispatarrs eee rest 
Barbula , unguiculata var. Alpine. 

cuspidata SISSE 1 st ; sk 
DE ED ER littoraler rd Dicranum Andersonii ... 3 1 
Leucodon sciuroides .. PER | SUSE RLSS 
Isothecumsmyunrumesseese 2] Ubiquists. 
Eurhynchium piliferum.. 3 2 Grimmiapa lenses ist 


c.. Most frequent in South Iceland, rarer in the other districts of Iceland. 


STEWSÆNWEÆNEDE S. W. NW. N. E. 
Meridional. Grim mia torgu ta seere EE2EE RD: 
zadula complanata . 3 3 JERSEER DP Fagtob ryu meer 22 
Madotheca Cordæana 4 3 2RED, 
Barbula cylindsicaseen See 1 f Alpine. 
Mnium hornum . AP SERIES 
rr serratum Fe. JP 3 2 1 Bryum Oak bo 01554732 3 1 1 1 VA 
Catharinea undulata. 4 3 12 
Neckera complanata. 2 1 Ubiquists. 
Thuidium delicatulum 3 2 1 
3 i 9 9 
Rhynchostegium rus- OPR ERer, MB ER å > 1 SER 
Ciform reen SPAND | men Es E Ea 2 1 1 
Thamnium  alopecu- ir RE SER SÅS 
Eg P SEE Mnium. Seligeri. ÅRRE), ASS 
RESTEN SØREME : 2 Antitrichia curtipen- 
Hypnum commutatum 3 2 2 2 2 
VI dula seen 3730 
Boreal. Hypnum' palustre 573 2 aud! 
Sphagnum medium... 2 1 1 — … Ccupressiforme 4 3 % 2 
Hymenostylium  cur- Hylocomium loreum. 3 3 2 
VILS re sen 3 1 Fl URE SBE DT] Sa) RÅ dl 2 


2. Species with a mainly Northern Distribution. 


With the exception of the southernmost part of East Iceland 
— almost as far as Djupivogur — and the westernmost part of 
North Iceland — as far as Bléndudalur — there is a very great 


THE BRYOPHYTA OF ICELAND 659 


resemblance between the Bryophyte vegetation of East and North 
Iceland, therefore they will partially be mentioned collectively in 
the following lists. 


a. Found only in North Iceland. 


Meridional. Alpine. 
Sphagnum angustifolium....... JENS auterra al pinde es ere 1 
Leptodontium flexifolium ...... RS capania remote ERR 1 
Amblystegium trichopodium..….. 1. "Weisia Wimmeriana ;:....:1713 1 
Dicranoweisia compacta ....... 1 
Borcal. Pottiazlatifolrar FN ERE SEER 1 
Haplozia pumila... 1 4 Desmatodon cernuus.......... 1 
Pollia proligerar DE TUDSER 1 Bryum COMES CS NESS 1 
ED ESSEN KEEL SER SEER FRR Obs En dys USS FEE SES 1 
EN e 1 Eurhynchium diversifolium .... 1 
Moiim So SE Eescuræarrigescens Een i 
Amblyodon .dealbatus: SÆR 1 É 
Brachythecium latifolium ...... 1 Icelandic. 


Bryumtislandrenm sees rese 1 


b. Found only in East Iceland. 


Meridional. Bryum neodamense var. ovatum. 1 
Brevum bint mA SAS AE SE sg EkaHEn: BER HESS HERE 1 
Grim msn eur va PE 1 
Boreal. Ubiquists. 
Leptoscyphus anomalus ....... 1. Cynodontium polycarpum...... 1 
Bryum calophyllum........... Torbe ane na kar 1 
Alpine. Icelandic 
Dire males ESS KS 1FFontinalis islands ore 1 


c. Found only in North and East Iceland. 


N. E. N. E. 
Meridional. Plastopust dere lt 
Diplophyllum obtusifolium .. 1 1. Hypnum trifarium.......... od 
Boreak Alpine. 
Schistidium confertum...... 2 1 OrthotrichumKillaso ER ser 
Grimmia;ovatar tisse des 2 2 


d. Most widely distributed in East and North Iceland. 


E. N.NW. W.S. E. N.NW.W. 

Boreal Poblia tenuifohas sæ BER 1 
Dicranum congestum.. 4 4 ? 3 2 Bryum purpuråscens.. 2 3 2 2 1 
Distichium inclinatum. 1 2 1 lacus re FH DESEE PO NH IKA 
Dissodon splachnoides. 1 4 3 1 1  Paludella squarrosa... 4 4 3 3 2 


660 A. HESSELBO 


E. N. NW,W. S. E. N.NW.W. 
Cinclidium stygium ... 44333 All pine: 
Thuidium lanatum . . 44221. Aulacomnium turgidum 2 2271 
Orthothecium chryseum 4 4 2 2 
Hypnum decipiens.... 4 4 1 Ubiquists. 
— Richardson see2 1 Meesealtriquetrameeere SE 522 
3. Species with a mainly Western Distribution. 
a. Found only in North-west Iceland. 
Boreal. Mortulakacip hy agterste 


Haploziaysphærocarp FE SSR Ey pnu me iluitanssvartfalea arme 
Polytri cum Sari ses 


— varme seensesseeeel Ubiquists. 
Marsupellakaquaticasessmeeeer 
Alpine. Gymnocoleasin dataene 
Gymnomitriumrevolutum sreseeRes phasnum Grave her 
Sphenolobuskp olmus rese 1 = Lipatium ss 
Cephalozia ambigua..... eee Eesker Catena rr ene 


b. Found only in West Iceland. 


Meridional. Pohlianpolym or phase 
Trematodon ambiguus..... .……. 1. Pogonatum dentatum var. minus. 
Polytrichumsformosumn seere 1 
Pogonatum polytrichoides...... 2 Ubiquists. 

Fissidenstdeciprens serene 
Alpine: Encalyptakcontorta seere 
Plasiobryunkdemnissunserrer 1 


c. Most widely distributed in North-west Iceland. 


E. N.NW.W. S. E. N. NW.W. 
Boreal. Philonotisfseratustsere SE 4E2, 
Lophozia Flærckei.... 4 2 Oligotrichum  hercyni- 
HarpanthusFlotowianus 1 1 4 2 CUS SSR FEE PE DERANED: 
Scapania uliginosa.... 1 3 Polytrichum sexangulare 3 2 4 3 
Dicranellafisquarros ar EP ES escur ædt Breidlersere ss 2ÆS 
Bryum ki cinr stum 2EPÆS EPE 
Ubiquists. 
Alpine. Djijeranum majuskeeer Gl 
Pleuroclada "albescens 33 TABE 2 = fuscescens... 4 1 
Pohliatcucullata ser 12 Pohliasnutans rss eres] 
d. Most widely distributed in the western part of Iceland. 
E. N.NW.W. S. E. N.NW.W. 
Ubiquists. Dicranum elongatum .. PARSP NEP 
Dicranella cerviculata.. 1 2 2 Isothecium tenuinerve . 155) 
Boreal. Meridional. 
DicranumsBlytti FEE SES Camptotheciumlutescens 2223 


Barbue 1 


DR 


[55] 


THE BRYOPHYTA OF ICELAND 661 


Table VI. Collective Survey of the Number of Species 
in the different Districts of Iceland. 









































| 
Plant-geographical character m b | a | u | 5.2 
| — | El) | E 2 
| | | | | SA 
Frequency | 12 SEER BLE 2 ESS ÆRE 28832 | 1-2 | 3-4 > 
| | | 
FEE > TI = FIT EG | 
SOUEDEE Ice lan drer I 39 16 31 41 |" 30 | 36 23 | Tor 1297 
Found only in South Iceland | 26 GR ER SE KARET | HEE ASK MURE SE RES RR 51 
| li | | 
Bret ledn, 38 does asrE (Er 245 52 650 51 K3 9 830 530 RG SE Roo 
Found only in East Iceland 1 |ES> | 4 |E2A) 9 
Northrend FLE ES ES Et | 483 | 30 | 38 | 68 | 287 
Found only in North Iceland | 3 ml | | 10 20 
Found only in N. and E. | | | | | 
ICSlan der REPARERE hæs SEN eet | 6 
North-west Iceland .…...… za BR53E 1980 Er 9 H557ÆR ESTER 525 | 61 | 222 
Found only in North-west | | | | | | 
end SNERRE 358 | 1538 HERE FL SS Sr Vere 
Westikcelandker, rr ne Ios Es 375 Fag 305 957 33 76 logo 
Found only in West Iceland 3 | | 3 2 | 8 
| | | | 

















By comparing the Tables and Lists of species contained in the 
present section it is seen that: — 

The Meridional species belong mainly to the southern part 
of Iceland, in the south-western part of which they arrive at their 
maximum, both as regards number and frequency, and from thence 
decrease in number and frequency towards north-west and east. In 
south-east Iceland there is a tolerably sharp limit for their distri- 
bution, whilst a great number of species are to be found — but 
with decreasing frequency — even throughout the whole of West 
Iceland. The respective percentages of the entire number of species 
are tastfollowstr south Iceland" 199/$7East Iceland 6"9/8-ÆN orth 
Iceland 9 2/,; North-west Iceland 5 ?/, and West Iceland 12 2/,. A 
comparatively great number of species have been found in a single 
locality, or in a few localities, in North Iceland, especially around 
Eyjafjéordur and eastwards in the valleys towards Myvatn, which may 
probably be due to the higher summer temperature of these districts. 


662 A. HESSELBO 


The Boreal Group, over the whole of Iceland — with the 
exception of South Iceland — constitutes the same percentage of the 
vegetation (S. Icel. 24 %,; E. Icel. 307,5 N. Cell ENE es 
and W. Icel. 30 2/,), and the great majority of the species belong to 
those which are common everywhere (56 species are common to 
all the districts of Iceland, with a frequency of 3—4, corresponding 
to about 60—75 9/.). 

The Alpine Group likewise shows almost the same percent- 
age in all the” districts (SS Tcel 25 YE TE: Iceleo5 "SØN Feel OS 
NW. Icel. 26 /, and W. Icel. 20 2/,) of which about half the number 
(some 30 species) are common to all the districts. 

The Ubiquitous Group comprises chiefly species which have 
a great distribution in all parts of Iceland (S. Icel. 34 %,; E. Icel. 
39 92/,: N. Icel. 38 %/,; NW: Icel. 39 2%, and. W. Icel: 38. 2/8), 73 0£ 
which (67—85 ?/.,) are common to all the districts, with a frequency 
of 3—4. 

The composition of the vegetation as regards percentage is thus 
very nearly the same everywhere as regards the three last groups, 
whilst, with regard to the meridional species, the country may be 
divided into a north-eastern and a south-western part. 


The North-eastern part comprises East and North Iceland 
as far as Blåndudalur. The meridional species play a very slight 
role in the vegetalion here. Two species only, Metzgeria furcata and 
Rhacomitrium aciculare, are common everywhere in this part, and 
five species, viz. Radula complanata, Madotheca Cordæana, Catha- 
rinea undulata, Mnium hornum and Hypnum commutatum, are some- 
what frequent in East Iceland, but are absent from, or are rare in, 
North Iceland. 

Some few, principally Boreal and Alpine species, are charac- 
teristic of North and East Iceland. 

Hypnum decipiens is very common from Seydisfjårdur to Blåndu- 
dalur, but is absent from, or is rare in, the other parts of Iceland. 

Orthothecium chryseum is one of the most commonly occurring 
species in the whole of East and North Iceland, but is absent from 
South Iceland, and is rather rare west of Hunafl6i. 

: Grimmia ovata has been found only in East and North Iceland. 
Philonotis tomentella is very common in meadow-ground in North 
and East Iceland, from Seydisfjårdur; but is less frequent on damp 
rocks in other parts of Iceland. 


THE BRYOPHYTA OF ICELAND 663 


Cinclidium stygium, Meesea triquetra, Paludella squarrosa, Thuidium 
lanatum, Hypnum exannulatum, H. Richardsonii, H. sarmentosum and 
H. alpestre occur far more abundantly in the northern half of Ice- 
land than in the southern, and are often the most abundant con- 
-stituents of the vegetation. 

Dicranum congestum, which is one of the most commonly oc- 
-Curring species in North and East Iceland and cccur there in 
numerous forms, is far more rare in South and West Iceland, and 
is evidently absent from North-west Iceland. 

A rather considerable number of species, which are most fre- 
quent in South and West Iceland, decrease in frequency throughout 
East Iceland, and are absent from, or are rarer in, North Iceland, 
for instance Scapania dentata, Anæctangium compactum, Grimmia 
torquata, Rhacomitrium heterostichum, R. fasciculare, Amphidium Mou- 
geottii, Mnium Seligeri, Diphyscium sessile, Hypnum filicinum, H. cu- 
pressiforme, Hylocomium parietinum and H. triquetrum. 

South Iceland. The vegetation is primarily characterized 
by the presence of numerous meridional species, of which those 
with a frequency of 3—4 constitute an essential part of the moss- 
<overing. 

Fegatella conica and Reboulia hemisphærica in company with 
Preissia commutata and Marchantia polymorpha form special Mar- 
chantiaceæ-associations on damp shady tuff-faces. Lejeunea cavifolia, 
Madotheca Cardæana, Barbula cylindrica, Mnium serratum, M. undu- 
latum, Catharinea undulata, Thuidium tamariscinum, T. delicatulum, 
T. Philibertii, Eurhynchium piliferum, E. Swartzii, Rhynchostegium 
rusciforme, Scleropodium purum, Thamnium alopecurum, Hypnum 
molluscum and H. commutatum are the most frequently occurring 
species in SW. Iceland. Of the boreal Bryophyta Haplozia riparia, 
Hymenostylium curviroslre, Grimmia torquata and Plagiobryum Zierii 
are most widely distributed in South Iceland; but this is no doubt 
-chiefly due to the fact that a suitable substratum (tuff) abounds. 

The Alpine species Dicranum Andersonii, D. fulvellam and Bry- 
-—oxiphium norvegicum are common in the tuff districts of South Ice- 
land, but have not been found in other parts of the country. 

Several ubiquists occur more frequently and abundantly in South 
Iceland ihan in the other districts, for instance Mnium Seligeri, Anti- 
trichia curtipendula, Hypnum palustre, H. cupressiforme, Hylocomium 
loreum, H. triquetrum and H. parietinum. These species ascend only 
exceptionally above the limit of the birch. 


664 A. HESSELBO 


The difference in the climate is also apparent from the fact 
that species, which in North and East Iceland are common as far 
down as the low land, in South Iceland are first met with in 
abundance at a higher altitude. This is for instance the case with 
Schistidinm rivulare, Oncophorus Wahlenbergii, O. virens and Dicra- 
num congestum which åre not common until at a height of 200— 
300 metres. The Bryophyte vegetation in the higher-lying bogs (200 
—3500 metres) also corresponds closely with the bog vegetation of 
the lowlands of North and East Iceland, while the bog vegeta- 
tion of South Iceland differs somewhat considerably in character 
(pas oe 

North-west Iceland. The Bryophyte vegetation differs in 
several points from that of the other districts. From a narrow belt 
of coastal land the country rises abruptly to a height of about 500 
metres, so that there is only room for the boggy tracts, so common 
in the other districts of Iceland, in those valleys which, from the 
head of the fjords, ascend towards the high land. Here the number 
of species is essentially smaller than in the other districts of Ice- 
land, since many of the common species, as well as of the rare 
ones, are absent, on the other hand, however, there are some few 
species which are widely distributed in North-west Iceland, but are 
absent from, or rarer in the other districts. This, among others, 
is the case with regard to several of the species belonging to the 
heath formation. 

Dicranum fuscescens, D. majus and D. molle are very common 
in NW. Iceland and, in association with other species (especially 
Hepaticæ), form Dicranum heaths (p. 591): a formation which is 
otherwise rare in Iceland. Pohlia nutans and Bryum cirratum are 
also common, especially in the birch coppices. Polytrichum piliferum 
is tolerably frequent and has been found in fruit in several places. 

Lophozia Flærkei is very common as far upwards as about 
500 metres above sea-level. It has, however, only been found in a 
lava-field in SW. Iceland. 

There are also several hygrophilous species which are charac- 
teristic of NW. Iceland: Sphagnum squarrosum is frequent on damp 
slopes and $. riparium is rather common in pools. On inundated 
ground Scapania uliginosa, S$. paludosa, Harpanthus Flotowianus and 
Philonotis seriata are very common and often form — either sepa- 
rately or collectively — the bulk of the vegetation. Hypnum fluitans 
var. falcatum occurred abundantly north of Isafjérdur. 


THE BRYOPHYTA OF ICELAND 665 


The rocky flat is very widely extended in NW. Iceland, since 
it comprises all the flats and plateaus above the mountain slopes, 
and on the north side of the mountains descends as far downwards 
as to the sea-level. Several rocky-flat plants are therefore more 
widely distributed here than in the other districts of Iceland. Oligo- 
trichum hercynicum, Rhacomitrium sudeticum, Conostomum boreale, 
Lescuræa Breidleri, Gymnomitrium concinnatum and Pleuroclada al- 
bescens are very common. Sphenolobus politus, Haplozia sphæro- 
carpa, Gymnomitrium revolutum and G. varians have been found 
only on the rocky flats of NW. Iceland. 

Lescuræa filamentosa is very common and found fruiting on 
detached blocks and on rocks, and L. patens is also frequent and 
likewise sets fruit. 

Some few species, which are widely distributed elsewhere in 
Iceland, are absent here, for instance Scapania dentata, Radula com- 
planata, Gymnostomum rupestre, Campylopus Schimperi, Sælania cæsia, 
Mnium Seligeri, Hypnum cupressiforme and Hypnum palustre, whilst 
others are rare, for instance Preissia commutata, Leptobryum pyri- 
forme, Catoscopium nigritum, Timmia austriaca and Hypnum falcatum. 
The comparatively abundant occurrence of species of Sphagnum, 
and the lesser frequency of the last-mentioned lime-loving species, 
appear to indicate that the soil here is less calcareous than in the 
other districts of Iceland. 

The Bryophyte vegetation of NW. Iceland has, on the whole, 
a more decidedly xerophilous and Arctic character than that of the 
rest of Iceland. 

West Iceland has, with the exception of the southern part 
as far as Borgarfjordur, been very superficially investigated and, 
apart from a few scattered collections made by Helgi Jonsson, 
especially in Buådahraun on Snæfellsnes, and my own collections 
near Stykkisholmur, the whole stretch of coast north of Hvalfjordur 
and around Breidifjordur is, from a bryological point of view, quite 
unknown. 

Several of the species characteristic of South Iceland occur here 
also, partially decreasing in frequency throughout West Iceland, for 
instance Thuidium delicatulum, Hylocomium loreum, Eurhynchium 
piliferaum and (in the South-west) Madotheca Cordæana. Some species 
have a decidedly westerly distribution in Iceland, being of almost 
equal frequency in West and South-west Iceland and a few also in 


The Botany of Iceland. Vol. I, part II. 43 


666 A. HESSELBO: THE BRYOPHYTA OF ICELAND 


the North-west, whilst they åre absent from, or rare in, the other 
districts of the country. To this group belong: Dicranella cerviculata, 
Dicranum Blyttii, D. elongatum (western part of N. Iceland around 
Hunafl6éi; NW. and W. Iceland), Ditrichum homomallum (common in 
the South-west), Isothecium tenuinerve, Rhynchostegium rusciforme, 
Hypnum hamulosum and Hylocomium parietinum. Finally, Diplo- 
phyllum albicans is characteristic of all the lava-fields of West and 
South-west Iceland. 








BIBLIOGRAPHY. 


- Ås to the older records respecting the Flora of Iceland (Zoéga, Gliemann, 
Hooker, Hjaltalin, Vahl, Hornemann, Lindsay, Caroll and others) 
reference may be made to the full mention of these authors by Gréånlund 
in Botan. Tidsskr., Vol. 4, 1870—71, p. 147 and Vol. 7 (2. R., 3. Bd.), 1873, p. 1. 


Årnell, H. W. (1892), Lebermoosstudien im nårdlichen Norwegen. Jønkåping. 

Arnell, H. W. and Jensen, C. (1907), Die Moose des Sarekgebietes. Naturw. 
Unters. des Sarekgebietes in Schwed.-Lappland, III, Stockholm. 

Braithwaite, The British Moss-Flora. London, vol. I, 1880 - 1887; vol. II, 1888 
—1895; vol. III, 1903—09. 

Flora danica. Icones plantarum in regnis Daniae et Norvegiae nascentium ad 
illustrandam Floram danicam ediderunt G.C. Oeder, O. F. Muller, M. Vahl, J.W. 
Hornemann, S. Drejer, F. M. Liebmann, Joh. Lange, Fasc. 1—50. Hafniae, 1761 
—1880. 

Gronlund, Chr. (1870—71), Bidrag til Oplysning om Islands Flora. Bot. Tidsskr., 
Bd. 4, pp. 147—172. 

— (1873), Bidrag til Oplysning om Islands Flora. 2. Hepaticæ og Musci. Bot. 
Tidsskr., Bd. 7, pp. 1—26. ; 

— (1877), Islandske Naturforhold med særligt Hensyn til Mosvæxtens Betydning 
for Landskabet. Tidsskr. for populære Fremst. af Naturvidensk., 5 R., IV, 
pp. 321—356. 

— (1881), Islands Flora. Kjøbenhavn. 

— (1884), Karakteristik af Plantevæxten paa Island sammenlignet med Floraen i 
flere andre Lande. Den Naturhistoriske Forenings Festskrift, Nr. 1, Kjøben- 
havn, pp. 107—145. 

— (1885), Afsluttende Bidrag til Oplysning om Islands Flora, Musci, Hepaticae, 
Lichenes. Bot. Tidsskr. XIV, pp. 1539—217. 

— (1895), Tillæg til Islands Kryptogamflora, indeholdende Lichenes, Hepaticae og 
Musci. Bot. Tidsskr. XX, pp. 90—115. 

Hagen, I. (1899—1901), Musci Norwegiae Borealis. Tromséå Museums Aarshefter 
21—22, 

—  Forarbejder til en norsk Lovmosflora I—XX. Det Kgl. Norske Videnskabers 
Selskabs Skrifter, Trondhjem 1908—1915. 

Jensen, C. (1897), Beretning om en Rejse til Færåerne i 1896. Bot. Tidsskr. XXI, 
pp. 157—219. 

— (1901), Bryophyta. Botany of the Færåes, vol. 1, Copenhagen, pp. 120—197. 

— (1915), Danmarks Mosser. I. Hepaticales, Anthocerotales, Sphagnales. Køben- 
havn og Kristiania. 

Jønsson, Helgi (1895), Studier over Øst-Islands Vegetation. Bot. Tidsskr. XX, 
pp. 17—89. 

— (1899), Floraen paa Snæfellsnes og Omegn. Bot. Tidsskr. XXII, pp. 169—207. 

43" 


668 BIBLIOGRAPHY 


Jønsson, Helgi (1900), Vegetationen paa Snæfellsnes. Vidensk. Medd. fra den 
Naturhist. Forening i Kjøbenhavn, pp. 15—97. 

— (1905), Vegetationen i Sydisland. Bot. Tidsskr. XXVII, pp. 1—82. 

— (1912), The Marine Algal Vegetation. The Botany of Iceland, Part 1,2. Copen- 
hagen and London. z 

Limpricht, K. G., Die Laubmoose in Rabenhorst Kryptogamenflora von Deutsch- 
land, Oesterreich und der Schweiz. Leipzig 1890—1904. 

Loeske, Leopold (1905—6), Kritische Bemerkungen uber einige Formen von 
Philonotis. — Hedwigia, Bd. 45, p. 100. 

— (1905—6), Kritische Ubersicht der europåischen Philonotis-Arten. Hedwigia, 
Bd. 45, p. 145. 

— (1911), Revision einiger Amblystegium aus dem Herbare Limpricht. Ung. 
botan. Blåtter, pp. 272—97. 

Muller, K., Die Lebermoose in Rabenhorsts Kryptogamenflora von Deutschland, 
Oesterreich und der Schweiz. Leipzig 1909—1915. a 

Ostenfeld, C. H. (1899), Skildringer af Vegetationen paa Island. Bot. Tidsskr. 
XXII, pp. 227—253. 

Stefånsson, St. (1896), Bemærkninger til Chr. Grénlund: Tillæg til Islands Kryp- 
togamflora indeholdende Lichenes, Musci, Hepaticae. Bot. Tidsskr. XX, pp. 399 
—402. 

Thorodsson, Th. (1914), An Account of the Physical Geography of Iceland, with 
Special Reference to the Plant-Life. The Botany of Iceland, Part I, 2. Copen- 
hagen and London. 

Warming, Eug. (1884), Om Grånlands Vegetation. Medd. om Grønland, XII, 
245 pp. 

— (1895), Plantesamfund. Grundtræk af den &kologiske Plantegeografi. Kjøbenhavn. 

Warnstorff, C., Kryptogamenflora der Mark Brandenburg, Bd. I, Moose. 1901 
—1903. 

Winther (1910), Beitråge zur Kentniss der Pohlia commutata, P. gracilis, P. cu- 
cullata und P. carinata. Hedwigia 49, p. 54. 


FISTFOFFHEFSSPECIES: 


Acrocladium cuspidatum (L.) Lindb. 541. 
Alicularia geoscypha De Not. 409. 
— scalaris (Schrad.) Corda 408. 
Amblyodon dealbatus (Dicks.) P. B. 491. 
Amblystegium aduncum (L.) Lindb. 531. 
chrysophyllum' (Brid.) Lindb. 530. 
cordifolium (Hedw.) de Not. 539. 
compactum (C. M.) Br. eur. 527. 
curvicaule Dicks. et James. 534. 
exannulatum de Not. 532. 
filicinum (L.) de Not. 533. 
fluitans (L.) de Not. 533. 
fluviatile (Sw.) Br. eur. 526. 
giganteum de Not. 540. 

glaucum (Lam.) Lindb. 534. 

P, decipiens Lindb. 534. 
intermedium Lindb. 531. 
Juratzkanum Schimp. 526. 
Kneiffii Br. eur. 532. 

littorale (C. Jens.) Hsb. 526. 
molle P, alpinum Lindb. 538. 
ochraceum Lindb. 539. 

palustre (Huds.) Lindb. 538. 

— polygamum Br. eur. 530. 

— protensum Lindb. 530. 

revolvens de Not. 531. 
Richardsonii Lindb. 540. 
rivulare (Sw.) Lindb. 538. 
sarmentosum de Not. 541. 
scorpioides (L.) Lindb. 542. 
salinum Bryhn. 528. 

Sendtneri (Schimp.) de Not. 531. 
serpens (L.) Br. eur. 526. 

— var. littoralis C. Jens. 526. 
Smithir (Sw.) Lindb. 538. 

Sprucei (Bruch) Br. eur. 526. 
stellatum Lindb. 530. 

stramineum de Not. 540. 
trichopodtum (Schultz) Br. eur. 528. 
trifarium de Not. 541. 

turgescens Lindb. 541. 


Amphidium lapponicum (Hedw.) Schimp. 
463. 

— Mougeottir (Br. eur.) Schimp. 463. 

Andreæa petrophila Ehrh. 434. 

Aneura latifrons Lindb. 405. 

— multifida (Lindb.) Dum. 404. 

— pinguis (L.) Dum. 404. 

Anisothecium crispum Lindb. 439. 

Anomobryum concinnum (Spr.) Lindb. 469. 

— filiforme (Dicks.) Husnot. 469. 

Anæctangium Mougeottii Lindb. 463. 

— l[apponicum Hedw. 463. 

Anomodon viticulosus (L.) H. et T., 510. 

Anthelia julacea (L.) Dum. 423. 

— Juratzkana (Limpr.) Trevis. 423. 

— mnivalis (Sw.) Limpr. ex p. 423. 

Anthoceros punctatus L. 429. 

Antitrichia curtipendula(Hedw.) Brid. 509. 

ÅAongstræmia longipes (Somf.) Br. eur. 437. 

Archidium phascoides Bridel. 434. 

AÅstrophyllum cinclidioides (Blytt) Lindb. 

490. 

cuspidatum (L.) Lindb. 489. 

hornum (L.) Lindb. 487. 

marginatum (Dicks.) Lindb. 487. 

medium Lindb. 488. 

orthorrhynchum Lindb. 487. 

pseudopuncetatum (B.S.) Lindb. 490. 

punctatum (L.) Lindb. 490. 

Seligert Lindb. 489. 

silvaticum Lindb. 488. 

spinosum Lindb. 488. 

stellare Lindb. 490. 

undulatum Lindb. 488. 

Aulacomnium palustre (L.).Schwågr. 493. 

— turgidum (Wahlb.) Lindb. 493. 


Barbula curvirostris Lindb. 435. 
— cylindrica (Tayl.) Schimp. 454. 
— fallax (Hedw.) 453. 

— — var. /ævifolia n. sp. 453. 


67 


0 


Barbula icmadophila Schimp. 455. 


rubella (Hoffm.) Mitt. 451. 
unguiculata (Huds.) Hedw, 453. 


— — var. cuspidata Braith. 453. 
Bartramia ityphylla (Haller) Brid. 494. 


ØÆderi (Gunn.) Limpr. 494. 


Blasia pusilla L. 407. 
Blepharaostoma trichophyllum (L.) Dum. 


422. 


Blindia acuta (Huds.) Br. eur. 448. 


Brachythecium albicans (Neck.) Br. eur. | 


52 

cirrosum Schimp. 523. 

collinum (Schleich.) Br. eur. 518. 
erythrorrhizon Br. eur. 521. 

glaciale Br. eur. 520. 

glareosum (Bruch.) Br. eur. 521. 
latifolium (Lindb.) Philib. 522. 
longipilum n. sp. 518. 

Mildeanum Schimp. 517. 

populeum (Hedw.) Br. eur. 520. | 
veflexum (Starcke) Br. eur. 520. 
rivulare (Bruch) Br. eur. 521. 
salebrosum (Hoffm.) Br. eur. 518. 
velutinum (L.) Br. eur. 520. 


Bryoxiphium norvegicum (Brid.) Mitten. 


450. 


Bryum affine (Bruch) Lindb. 482. 


archangelicum Br. eur. 475. 
arcticum (R. Br.) Br. eur. 486. 
argenteum L. 485. 

bimum Schreb. 482. 

Brownii Br. eur. 487. 
calophyllum R. Brown. 479. 
capillare L. 484. 

cirratum Hoppe et Hornsch. 483. 
comense Schimp. 484. 
cuspidatum Schimp. 482. 
cæspiticium L. 484. 

Duvalii Voit. 485. 

elegans N. v. Es. 484. 

fallax Milde. 480. 

Grønlundii n. sp. 479. 
inclinatum (Sw,) Br. eur. 475. 
intermedium (Ludw.) 483. 
islandicum n. sp. 478. 
Jørgensenii Kaurin. 475. 
Kaurimi Philib. 475. 

lacustre Blandow. 474. 
Neodamense Itzigs. var. ovatum (Jur.; 
485. 


LIST OF THE SPECIES 


Bryum æneum Blytt. 480. 
— pallens Sw. 485. 

— pallescens Schleich. 483. 
— pendulum (Hornsch.) 486. 
— purpurascens (R. Br.) 474. 
— retusum Hagen. 477. 

— rutilans Brid. 480. 

— subrotundum Brid. 483. 
— uliginosum (Bruch) 479. 
— ventricosum Dicks..486. 


Calypogeia Trichomanis (L.) Corda. 422. 

Camptothecium lutescens (Huds.) Br. eur. 
Sige 

— nitens (Schreb.) Schimp. 517. 

Campylopus flexuosus (L.) Brid. 447. 

— fragilis (Dicks.) Br. eur. 447. 

— Schimperi Milde 446, 

Catharinea undulata (L.) Web. 497. 

— — var. thermophila n. var. 498. 

Catoscopium nigritum (Hedw.) Brid. 492. 

Cephalozia ambigua Mass. 419. 

— bicuspidata (L.) Dum. 419. 

— media Lindb. 420. 

— pleniceps (Aust.) Lindb. 419. 

Cephaloziella Hampeana(Nees)Schiffn. 421. 

— rubella (Nees) Warnst. 421. 

Ceratodon purpureus (L.) Brid. 448. 

Chandonanthus setiformis (Ehrh.) Lindb. 
422. 

Chiloscyphus polyanthus (L.) Corda. 418. 

Chomocarpon quadratus(Scop.)Lindb. 403. 

Cinclidium stygium Sw. 491. 

Climacium dendroides (L.) W. et M. 516. 

Conostomum boreale Swartz. 492. 

— tetragonum (Vill.) Lindb. 492. 

Ctenidium molluscum (Hedw.)Schimp. 535. 

Cylindrothecium  concinnum (de Not.) 
Schimp. 516. 

Cynodontium polycarpum (Ehr.) Schimp. 
437. 


Desmatodon cernuus (Hubn.) Br. eur. 455. 

— latifolius (Hedw.) Br. eur. 455. 

Dichodontium pellucidum (L.)Schimp. 437. 

Dicranella cerviculata (Hedw.) Schimp. 
440. 

— crispa (Ehrh.) Schimp. 439. 

— Schreberi (Sw.) Schimp. 439. 

— secunda (Sw.) Lindb. 439. 

— squarrosa (Starke) Schimp. 438. 


LIST OF THE SPECIES 671 


Dicranella subulata (Sw.) Lindb. 439. | Eurhynchium Stockesii(Turn.) Br. eur. 523. 

Dicranoweisia compacta (Schleich.) | — strigosum (Hoffm.) Br. eur. 522. 
Schimp. 436. SE Var PK ÆCOT (Hedw.) 522. 

— crispula (Hedw.) Lindb. 436. | — Swartzrii (Turn.) Curnow. 450. 


Dicranum Andersonii(Wich,)Schimp. 440. | Eustichium norvegicum Br. eur. 450. 
— angustum Lindb. 443. 


— arcticum Schimp. 442. | Fegatella conica Corda. 403. 
— Blyttii Schimp. 441. Fimbriaria pilosa (Whlb.) Tayl. 403. 
— Bonjeani de Not. 443. Fissidens adianthoides (L.) Hedw. 448. 
— congestum Brid. 445. | — bryoides (L.) Hedw. 447. 
— — var. fleæicaule Brid. 445. — decipiens de Not. 448. 
— — — subspadiceum Arn. et Jens. 445.  — osmundoides (Sw.) Hedw. 447. 
— elongatum Schleich. 446. Fontinalis androgyna Ruthe. 508. 
— falcatum Hedw. 441. — antipyretica L. 506. 
— fulvellum (Dicks.) Sm. 440. — islandica Cardot. 506. 
— fuscescens Turn. 445. — longifolia C. Jensen. 506. 
— glaciale Berggr. 442. — thulensis C. Jensen. 508. 
— majus Smith, 444. Fossombronia Dumortieri (Hub.et Genth.) 
— molle Wils. 442. 406. 
— palustre Br. eur. 443. | Frullania dilatata Nees. 428. 
— Schisti (Gunn.) Lindb. 441. — fragilifolia Tayl. 428. 
— scoparium (L.) Hedw, 444. — Tamarisci Nees. 428. 
— spadiceum Zett. 446. Funaria hygrometrica L. 468. 
— Starckei Web. et Mohr. 442. | — obtusa (Dicks.) Lindb. 468. 
Didymodon rubellus (Hoffm.) Br. eur. 451. 
— rufus Lorentz. 452. | Grimmia alpestris Nees. 459. 
Diplophyllum albicans (L.) Dum. 423. | — alpicola Swartz. 458. 
— obtusifolium (Hook.) Dum. 424. | -— apocarpa Hedw. 457. 
Diphyscium sessile (Schmid.) Lindb. 506. | — commutata Hub. 459. 
Dissodon splachnoides (Thunb.) Grev. 467. " — Doniana Smith. 459. 
Distichium capillaceum Br. eur. 450. — funalis (Schwågr.) Schimp. 460. 
— inclinatum (Ehrh.) Br. eur. 450. — hypnoides (L.) Lindb. 462. 
— montannm (Lam.) Hagen. 450. — incurva Schwågr. 459. 
” Ditrichumflexicaule(Schleich.)Hampe.449.  — microcarpa (Gmel.) 460. 
— glaucescens (Hedw.) Hampe. 449. — ovata W. et M. 459. 
— homomallum (Hedw.) Hampe. 449. — ovalis Lindb. 459. 
— mivale (C. M.) Limpr., 448. — patens (Dicks.) Br. eur. 460. 
— tortile (Schrad.) Lindb. 448. — ramulosa Lindb. 461. 
Dryptodon patens Brid. 460. — torquata Hornsch. 461. 
å Gymnocolea inflata (Huds.) Dum. 417. 
Encalypta ciliata (Hedw.) Hoffm. 466. Gymnomitrium concinnatum (Ligthf.) 
— contorta (Wulf.) Lindb. 466. Corda. 407. 
— rhabdocarpa Schwgr. 466. - — corallioides Nees. 407. 
Entosthodon ericetorum (Bals. et de Not.) — revolutum (Nees) Phil, 408. 
Br. eur. 468. — varians (Lindb.) Schiffn. 407. 
Entodon orthocarpus Lindb. 516. Gymnostomum vrupestre Schleich. 435. 


Eucalyw subellipticus(Lindb.)Breidler. 409. 

Eurhynchium cirrosum (Schwgr.) Limpr. | Haplozia atrovirens (Schl.) Dum. 411. 
523. — cordifolia (Hook.) Dum. 410. 

— diversifolium (Schleich.) Br. eur. 522. | — crenulata (Sm.) Dum. 409. 

— vpiliferum (Schreb.) Br. eur. 523. — pumila (With.) Dum. 411. 


672 


Haplozia riparia (Tayl.) Dum. 411. 
— sphærocarpa (Hook.) Dum. 410. 
Harpanthus Flotowianus Nees. 418. 
Hedwigia albicans (Web.) Lindb. 462. 
— ciliata Ehrh. 462. 
Heterocladium dimorphum (Brid.) Br. eur. 
513: 
— squarrosulum (Voit) Lindb. 513. 
Hepatica conica Lindb. 403. 
Homalothecium sericeum (L.) Br. eur. 516. 
Hylocomium loreum (L.) Br. eur. 543. 
— parietinum (L.) Lindb. 542. 
— proliferum (Hedw.) Br. eur. 542. 
pyrenaicum (Spr.) Lindb. 542. 
rugosum (Ehrh.) de Not. 544. 
— Schreberi (Willd.) de Not. 542. 
splendens (Hedw.) Br. eur. 542. 
squarrosum (L.) Br. eur. 543. 
triquetrum (L.) Br. eur. 543. 
Hypnum albicans Neck. 521. 
— aduncum (L.) Lindb. 531. 
alpestre Sw. 538. 
alpinum Schimp. 538. 
arcticum Sommerf. 538. 
Bambergeri Schimp. 536. 
callichroum (Brid.) Br. eur. 537. 
chrysophyllum Brid. 530. 
— var. tenellum Schimp. 530. 
commutatum Hedw. 534. 
— var. falcatum (Brid.) 534. 
cordifoium Hedw. 539. 
cupressiforme L. 536. 
curvicaule Jur. 534. 
cuspidatum L. 541. 
decipiens (de Not.) 534. 
erythrorrhizon Hartm. 521. 
eæannulatum (Gumb.) 532. 
— var. purpurascens (Schimp.) 
— — Rotæ (de Not.) 532. 
— — serratus (Warnst.) 532. 
filicinum L. 533. 
fluitans L. 533. 
— var. falcatum Schimp. 533. 
giganteuwum Schimp. 540. 
glaciale Hartm. 520. 
glareosum Bruch. 521. 
hamulosum Br. eur. 536. 
imponens Hedw. 535. 
intermedtum Lindb. 531. 
Kneiffii (Br. eur.) Schimp. 532. 
latifolium Lindb. 522, 


532. 





LIST OF. THE SPECIES 


Hypnum Lindbergii Mitten. 537. 

lutescens Huds. 517. 

molluscum Hedw. 535. 

murale Neck. 524, 

ochraceum Turn. 539. 

— palustre Huds. 538. 

— piliferum Schreb. 523. 

— plumosum Huds. 518. 

— polygamum Wils. 530. 

— populewum Hedw. 520. 

— protensum Brid. 530. 

— purpurascens Limpr. 532. 

DUT UM IE S22: 

vevolutum (Mitt.) Lindb. 536. 

vevolvens Sw. 531. 

Richardsonii (Mitt.) L. et James. 540. 

rivulare Bruch. 521. 

vusciforme Neck. 524. 

sarmentosum (Wahlb.) 541. 

Schultzit Limpr. 533. 

Sendtneri Schimp. 531. 

Sommerfeltit Myrin. 530. 

stellatum Schreb. 530. 

Stockesii Turn. 523. 

stramineum Schreb. 530. : 

strigosum var. diversifolium Lindb. 522. 

Swarteinw Turn. 523. 

trifarium W. et M. 541. 

trichoides Neck. 517. 

turgescens Th. Jens. 541. 

uncinatum Hedw. 531. 

— — var. orthothecioides (Lindb.) 531. 

Hymenostylium curvirostre (Ehrh.) Lindb. 
435. 


Isopterygium elegans (Hook.) 525. 

— nitidum var. pulchellum Lindb. 525. 
Isothecium myurum (Pollich) Brid. 516. 
— var. piliferum C. Jens. 516. 

— tenuinerve Kindb. 516. 


Jamesoniella autumnalis (de Cand.) Steph. 
AJ ol re 
Jungermannia bantryensis Hook. 416. 


Leersia contorta Lindb. 466. 

— laciniata Hedw. 466. 

— rhabdocarpa Lindb. 466. 

Lejeunea cavifolia (Ehrh.) Lindb. 428. 
— serpyllifolia (Dicks.) Spr. 428. 
Lepidozia setacea (Web.) Mitten. 422. 


LIST OF THE SPECIES 


Leptobryum pyriforme (L.) Schimp. 469. 

Leptodontium flexifolium (Dicks.) Hampe. 
452. 

Leptoscyphus anomalus(Hook.) Lindb. 417. 


Lescuræa Breidleri (Kindb.) Arn. et Jen- | 


sen. 512, 
— decipiens (Limpr.) 510. 
— filamentosa (Dicks.) Lindb. 511. 
— patens Lindb. 513. 
— radicosa (Mitt.) Hagen. 511. 
— rigescens (Wils.) Br. eur. 511. 
Leskea catenulata (Brid.) Mitten. 510. 
— nervosa (Schwgr.) Myrin. 510. 
Leucodon sciuroides (L.) Schwgr. 
morensis (Schwgr.) 508. 
Lophocolea cuspidata Limpr. 418. 
— minor Nees. 418. 


var. 


Lophozia alpestris (Schleich.) Evans. 414. | 


— barbata (Schmid.) Dum. 414. 

— eæcisa (Dicks.) Dum. 415. 

— Flærckei (W. et M.) Schiffn. 413. 
— heterocolpos (Thed.) Howe. 416. 
— Hornschuchiana (Nees) Macoun. 416. 
—- Kunzeana (Hub.) Evans. 414. 

— lycopodioides (Wallr.) Cogn. 412. 
— Miilleri (Nees) Dum. 416. 

— quadriloba (Lindb.) Evans. 413. 
— gqgwinquedentata (Huds.) Cogn. 412. 
— Schultzii (Nees) Schiffner. 416. 
— ventricosa (Dicks.) Dum. 414. 

— Wenzelii (Nees) Steph. 415. 


Madotheca Cordæana (Hub.) Dum. 427. 

— rivularis (Dicks.) Nees. 427. 

Marchantia polymorpha L. 404. 

Marsilia Neesiana Lindb. 405. 

Marsupella aquatica (Lindb.) Schiffner. 
408. 

— emarginata (Ehrh.) Dum. 408. 

— Funckiri (W. et M.) Dum. 408. 

Meesea trichoides (L.) Spruce 491. 

— triquetra (L.) Aongstr. 492. 

— tristicha Br. eur. 492. 

— uliginosa Hedw. 491. 

Metzægeria furcata (L.) Lindb. 405. 

Mniobryum albicans (Wahlb.) Lindb. 473. 

— var. glacialis (Schleich.) 473. 

Mnium affine Blandow. 489. 

-— cinclidioides (Blytt.) Hub. 490. 

— cuspidatum (L.) Leyss. 488. 

— hornum L. 487. 





673 


Mnium medium Br. eur. 488. 

— orthorrhynchum Brid. 487. 

— punctatum (L.) Hedw. 490. 

— Seligeri Jur. 489. 

— serratum Schrad. 487. 

— spinosum (Voit) Schwgr. 488. 

— stellare Reich. 490. 

— subglobosum Br. eur. 490. 

— undulatum (L.) Weiss. 488. 
Mollia fragilis (Drum.) Lindb. 453. 
— inclinata (Hedw.) Lindb. 452. 
— littoralis (Mitt.) 452. 

— tortuosa (L.) Schranck. 452, 
Myurella apiculata (Hub.) Br. eur. 509. 
— julacea (Will.) Br. eur. 509. 

— tenerrima (Brid.) Lindb. 509. 


Nardia crenulata (Sm.) 409. 
— hæmatosticta Lindb. 409. 
— minor (Nees) AÅrnell. 409. 
Neckera complanata (L.) Hub. 509. 


Odontoschisma denudatum (Mart.) Dum. 
421. 

— elongatum (Lindb.) Evans. 421. 

— Sphagnt (Dicks.) Dum. 421. 

Oligotrichum glabratum Lindb. 500. 

— hercynicum (Ehrh.) Lam. 499. 

— incurvum (Huds.) Lindb. 499. 

— lævigatum (Wahlb.) Lindb. 500. 

Oncophorus virens (Sw.) Brid. 438. 

— var. serratus Br. eur. 438. 

— Wahlenbergiri Brid. 438. 

— var. Ø, compactus (Funch.) 438. 

— — elongatus Hagen. 438. 

Orthothecium chryseum (Schwågr.) Br. eur. 
515. 

— intricatum (Hartm.) Br. eur. 515. 

— rvufescens (Dicks.) Br. eur. 515. 

Orthotrichum anomalum Hedw. 464. 

— Blyttii Schimp. 465. 

— cupulatum Hoffm. 464. 

— Killiasii C. M. 465. 

— lævigatum Zett. 465. 

— rupestre Schleich. 464. 

— saæatile Schimp. 464. 

— Sturmii Hornsch. 4635. 


Paludella squarrosa (L.) Brid. 491. 
Pellia Neesiana (Gottsche) Limpr. 405. 
Philonotis alpicola Jur. 496. 


674 


Philonotis Arnellii Husnot. 495. 

— fontana (L.) Brid. 494. 

— seriata (Mitt.) Lindb. 496. 

— tomentella Mol. 496. 

Plagiobryum demissum (H. et H.) 470. 

— Zierii (Dicks.) Lindb. 469. 

Plagiochila asplenioides (L.) Dum. 417. 

Plagiopus (Ederi (Gunn.) Limpr. 494. 

Plagiothecium denticulatum (L.) Br. eur. 
525. 

— depressum (Bruch) Dixon. 525. 

— elegans (Hook.) Sull. 525. 

— pulchellum (Dicks.) Br. eur. 5235. 

— Ræseanum (Hampe) Br. eur. 524. 

— silvaticum (Huds.) Br. eur. 524. 

— — var. Ræsei (Hampe) Lindb. 524. 

Pleuroclada albescens (Hook.) Spruce. 420. 

Pleurozygodon æstivus (Hedw.) 435. 

Pogonatum dentatum (Menz.) Brid. 501. 

— var. minus (Wahlb.) Hagen. 501. 

— nanum (Schreb.) 500. 

— polytrichoides (L.) Brockm. 500. 

— subrotundum Lindb. 500. 

— urnigerum (L.) P. B. 501. 

Pohlia acuminata Hornsch. 470. 

— albicans (Wahlb.) Lindb. 473. 

— annotina (L.) Lindb. 472. 

— commutata (Schimp.) Lindb. 471. 

— cruda (L.) Lindb. 470. 

— cucullata (Schwågr.) Bruch. 471. 

— gracilis (Schleich.) Lindb. 472. 

— grandiflora H. Lindb. 473. 

— Ludwigii (Sprengel) Schimp. 471. 

— nutans (L.) Lindb. 471. 

— polymorpha H. et H. 470. 

— proligera Lindb. 473. 

— Rothii (Correns) Broth. 472. 

— tenuifolia (Schimp.) 473. 

— Weigelii (Schimp.) Lindb. 471. 

Polytrichum alpinum L. 501. 

— commune L. 505. 

— — var. fastigiatum (Lyl.) Wils. 505. 

— formosum Hedw. 501. 

— gracile Dicks. 501. 

— — var. anomalum (Milde) 502. 

— inconstans Hagen. 505. 

— juniperinum Willd, 504. 

— piliferum Schreb. 504. 

— pilosum Neck. 504. 

— sexzangulare Flærcke. 502. 

— — var. vulcanica C. Jens. 502. 





LIST OF THE SPECIES 


Polytr. sexang. var. tenellum n. var. 502. 

— strictum Banks. 505. 

— Swartzii Hartm. 505. 

— Swartzit var. nigreseens (Warnst.) Ha- 
gen. 505. 

— urnigerum EL. 501. 

Pottia Heimii (Hedw.) Br. eur. 451. 

— latifolia (Schwågr.) C. M. 451. 

Preissia commutata (L.) Nees. 403. 

Pseudoleskea atrovirens (Dicks.) Br. eur. 
511. 

— patens (Lindb.) Limpr. 513. 

— radicosa (Mitt.) Lindb. 511. 

Psilopilum arcticum Brid. 500. 

— lævigatum (Wahlb.) Lindb. 500. 

Pterigynandrum filiforme (Timm) Hedw. 
510. 

Ptilidium ciliare (L.) Hampe. 423. 

Ptychodium decipiens Limpr. 510. 

— oligocladum Limpr. 512. 

— Pfundtneri Limpr. 511. 


Radula complanata (Dum.) Gottsche. 427. 

Reboulia hemisphærica (L.) Raddi. 403. 

Rhacomitrium aciculare (L.) Brid, 462. 

— canescens (Weis) Brid. 461. 

— fasciculare (Schrad.) Brid. 462. 

— heterostichum (Hedw.) Brid. 461. 

— hypnoides (L.) Lindb. 462. 

— lanuginosum (Ehrh.) Brid. 462. 

— microcarpum Brid. 461. 

— sudeticum (Funck) Br. eur. 460. 

BRhynchostegium murale (Neck.) Br. eur. 
524. 

— rusciforme (Neck.) Br. eur. 524. 

Riccardia latifrons (Lindb.) 405. 

— multifida Lindb. 404. 

— pinguis (L.) Gr. 404. 

Riccia bifurcea Hoffm. 402. 

— ecrystallina L. 402. 

— sorocarpa Bischoff. 402. 


Sauteria alpina Nees. 403. 

Scapania Bartlingii (Hampe) Nees. 427. 
— Carestiæ de Not. 427. 

— curta (Mart.) Dum. 426. 

— — var. geniculata (Mass.) 427, 

— — — rosacea (Corda) 427. 

— dentata Dum. 425. 

— irrigua (Nees) Dum. 425. 

— paludosa C. M. 425. 


LIST OF THF SPECIES 


Scapania purpurascens (Hook.) Pcars. 425. 

vemota Kaalaas. 424. 

subalpina N. ab Es. 424. 

— uliginosa (Sw.) Dum. 425. 

— undulata (L.) Dum. 426. 

Schistidium alpicola (Sw.) Limpr. 458. 

var. eualpicola Loeske. 458. 

— rivularis (Brid.) 458. 

— apocarpum Br. eur. 457. 

subsp. confertum (Funck) Loeske. 

458. 

— — gracile (Schwgr.) Loeske. 458. 
— — vulgare (Chal.) Loeske 457 

Seleropodium purum (L.) Limpr. 522, 

Scorpidium scorpioides (L.) Limpr. 542. 

Sphagnum acutifolium Ehrh. 432. 

— angustifolium C. Jensen. 433. 

Austini Sull. 430. 

compactum de Cand. 431. 

cymbifolium (Ehrh.) Hedw. 430. 

Dusenii C Jens. 433. 

— fimbriatum Wils. 431. 

Girgensohnii Russow. 431. 

Gravetit Russow. 431. 

imbricatum (Hornsch.) 430. 

tnundatum Russow. 430. 

- Lindbergii Sch. 433. 

medium Limpr. 430. 

papillosum Lindb. 430. 

rigidum Schimp. 431. 

vriparium Aongstr. 433. 

rubellum Wils. 432. | 

Russowir Warnst. 432. 

squarrosum Crome. 431. 

— subnitens Russ. et Warnst. 432. 

teres (Schimp.) Aongstr. 431. 

— Warnstorffii Rnssow 432. 

Sphenolobus minutus (Crantz) Steph. 411. 

— politus (Nees) Steph. 412. | 

— sazicola (Schrad.) Steph. 412. 

Sphærocephalus palustris (L.) Lindb. 493. 

— turgidus (Whlb.) Schwågr. 493. 

Splachnum pedunculatum (Huds.) Lindb. | 
467. j 

— sphæricum (L.) Swartz. 467. 

— vasculosum L. 468. 

Stereodon arcuatus Lindb. 537. 

— Bambergeri (Schimp.) Lindb. 536. 





Stereodon callichrous Brid. 537. 
chryseus (Schwågr.) Mitt. 515. 
hamulosus Lindb. 536. 
imponens (Hedw.) Brid. 535. 
— revolutus Mitten. 536. 
rufescens (Dicks.) Mitt. 515. 
— subrufus (Wils.) Lindb. 515. 
Swartzia inclinata Ehrh. 450. 
montana (Lam.) Lindb. 450. 
Sælania cæsia (Vill.) Lindb. 449. 


Tayloria lingulata (Dicks.) Lindb. 467. 

Tetraplodon bryoides (Zoéæga) Lindb. 467. 

— mnioides (L.) Br. eur. 467. 

Thuidium abietinum (L.) Br. eur. 514. 

Blandowii (W. et M.) Br. eur. 515. 

— delicatulum (L.) Mitten. 514. 

lanatum (Strøm) Hagen. 515. 

Philiberti (Limpr.) 514. 

tamariscinum (Hedw.) Br. eur. 514. 

Timmia austriaca Hedw. 497. 

— norvegica Zett. 497. 

Tortella fragilis (Drumm.) Limpr. 453. 

— inclinata (Hedw.) Limpr. 452. 

tortuosa (L.) Limpr. 452. 

Tortula aciphylla (Br. eur.) Hartm. 457. 

cernua (Hubn.) Lindb. 455. 

latifolia (Hedw.) Lindb. 455. 

— mucronifolia Schwågr. 456. 

muralis (L.) Hedw. 456. 

norvegica (Web.) Whilb. 457. 

obtusifolia Schleich. 456. 

— ruralis (L.) Ehrh, 456. 

— subulata (L.) Hedw. 456. 

Trematodon ambiguus (Hedw.) Hornsch. 
447. 

Trichostomum littorale Mitt. 452. 





Ulota maritima C. M. et Kindb. 463. 
— phyllantha Brid. 463. 


Webera annotina (L.) Lindb. 463. 

— bulbifera Warnst. 473. 

— sessilis Lindb. 507. 

Weisia crispata (Br. germ.) Jur. 436. 
— maritima Britt. 463. 

— Wimmeriana (Sendtn.) Br. eur. 436. 
— viridula (L.) Hedw. 436. 





. 429 


434 
438 
463 
471 
473 
588 


ERENAMKAS 


add Geysir! 
should read "I. Musci veri". 
for "elongatum” read "elongatus'. 


from bottom for '""Weissia” read "Weisia''. 


6 for "Wiinsted” read "Wiinstedt'. 


for "tenuifolia” read "grandiflora”. 


for "Associations” read "Formations". 


CONFENT S: 


Page 

Ha trodne ron SE SE SSR SE RE ARENSE EN VESTSIDE Rene EEN ae ER SEEREN 397 
FAR BisEsorsbhe Bryne SERENE: 402 
IEEE DELSE EET TREERE ET EEN el Eee AE SE 402 

INS phagnales sr Seer EET ENE eN ea EET 430 
BILERNES AEV EEN EEN SEE ESSEN SEE RE TEE ra Eee Er 434 

LEI hSBry0 plny ter Coma ES SEE EET rer 545 
bre Eowlands bor mathon ss EET 546 

bier ErttorakkBbBryophyter vere bab TOD SEE Er ER ere ne 546 
EHydrophlorsBryophytes Eorma tror ser Ea Nee 547 


The Bryophyte Vegetation of Running Water and of Lakes.... 548 


The Bryophyte Vegetation of Inundated Gravelly Soil... 550 

The Bryophyte Vegetation of Muddy Soil near Springs (Dy) ... 551 
iiherBryopliytes Vegetation of og sy S Ol re RER 555 
TherBryophyterVvegetationforeDampsandy Soren 564 

chef Bryophyter Vegetation mear” Hot"Springs LEES BE ASKERErE 565 

MesopiilorstB yo phytcEr os eee 3584 
XerophiloussBryophyter EF ormationse (le aEs) eee eee 588 
ikherBryophytenvegetatronsotherRocks Er RR PEN SS RENNER 595 
bhetBryophytet vegetation ore uk Roeks renere 604 

ke Bryophyter Vesetations of the avast ds re eee 612 

NMN eSBryoplyter Vegetahontor MountansHejg hes Eee eee 623 

HER Ehe Components orker Bryophkyte Flora NESS ENE 629 
Kære tAlbitudmalsDystributjon ofthe" Species FS AS FN ER 635 
Værter Horizonte Distribution of ther S pe es e 644 
Bibljog kap bye SEE REN En ET SNE ERE ere ERE Eee Re 667 
HEistroktherspecies sees ra de renser ERE ENE RTE TRE Tr ERE eee ERE 669 


rr ENE EEN En ER Rnen me ab Sr 2 ERE Me RS KENSERAE RE RENE: 676 





FREBOTPÅNY 


OF: 


CELAND 


EDITED 
BY 
L. KOLDERUP ROSENVINGE 


AND 


EUG. WARMING 


PH. D., SC. D. 


VOL. I 


WITH ONE PLATE AND 82 FIGURES IN THE TEXT 


COPENHAGEN 
JERIEM ODS 


LONDON 
VJOHNEWHEED ONE CO: 
1912-1918 


PUBLISHED BY THE AID OF THE CARLSBERG FUND 


H.H. THIELES BOGTRYKKERI. KØBENHAVN 


CONTENTS 
PARE-E 
BREBN CE SEE EN SR er STER E arer sted Eee eee see III 
1. HexGi Jonsson: The Marine Algal Vegetation of Iceland. 1912 1 
2. Tx. THORODDSEN: An Account of the physical Geography of 


SEE orn 5 UI ØRENE ES HER ERE SEES DS EEN ENES NEED SKE 187 


PART II 1918. 
3. ERNsT ØstRuP: Marine Diatoms from the Coasts of Iceland 


fwithronesPlate) Perm d im TOLG) FE SST SE ME DSE 345 


ARR UGSHESSEEB Ohe bryophyta ok Iceland SEES ERE 395 





Særtryk af Botanisk Tidsskrift. 37. Binds 4. Hefte. 1922. 


LIBRARY 
NEW YORK 
BOTANICAL 

GARGEN 


On a new species of Furcraea Vent. 
from Nicaragua. 
By 
Johs. Boye Petersen. 


(With Plate IV.) 


F; the year 1848 an Agave-like plant was introduced into 
the Botanical Garden of Copenhagen by A. S. Ørsted, who had 
collected it in Nicaragua. During the time the plant was culti- 
vated in the hot-house as Agave sp., it did not flower until 1921 
and therefore a determination was impossible. Prof. W. Trelease, 
paying a visit to Copenhagen .in 1912, critically revised the collec- 
tion of Agave in the Botanical Garden and also saw the specimen 
mentioned. He was, however, unable to name the sterile plant, 
but supposed that it belonged to an undescribed species. 

In the year 1921 it was observed that the specimen was 
about to flower and the development of the inflorescence was, 
of course, followed with interest. In December a scape with 
a panicle in all about 3 m high shot up. When trying to determine 
the flowering specimen I quickly found that it did not belong 
to the genus Agave but to Furcraea Vent., and moreover I discov- 
ered that it could not rightly be referred to any described species 
af Furcraea judging from the material, to which I have had 
access, viz. literature, living specimens in our hot-houses, dried 
specimens and specimens in alcohol in the museum. 

Unfortunately most descriptions of the species of Furcraea 
are more or less incomplete, and some species are not figured. 
It must be regretted that Drummond, who in 1907 wrote on 
the genus, has not given new descriptions of all species adopted 
by him, as he certainly has had an unusually good material at his 
disposal, although, as he says himself, it was deficient as to 
certain points. 


Botanisk Tidsskrift. 37. Bind. 20 


Arbejder fra den botaniske Have i København. Nr. 98; 


— 306 — 


Notwithstanding these insufficient descriptions I feel convin- 
ced that our plant represents an undescribed species. It is one 
of the smaller species of the genus, charcterised by its very 
compressed bulbils, small flowers and geminate spines in the 
margins of the leaves as in F. geminispina. I therefore propose 
to describe it as a new species and 
name it: 


Furcraea stratiotes Boye P. n. sp. 


F. caule brevissømo, foliis circiter 
50 dense rosulatis, 35—53 cm longis, 
3,5—5 cm latis, utrinque glabris lævibus- 
que, margine spinis rigidis geminis in- 
structis; scapo cum panicula 2,8m alto, 
floribus pendulis albidis pedicellatis 
parvis, 22 mm longis, bulbillis numerosis 
valde compressis; fructu ignoto. 

Specimen in Nicaragua lectum 1848 
(A. S. Ørsted) et in hortum botanicum 
Hauniense introductum et cultum. Fl. 
1921. 

Stem lacking or very short; leaves 
(about 50) in a rosette, with a base 
5—6 cm broad, 3 cm thick; above the 
base contracted to 2,5—3 cm, linear- 
lanceolate, 35—53 cm long, 3,5—5 cm 
broad at the middle, long-acuminate, 
the apex with a small whitish mucro, 
not a pungent spine, both surfaces 
Fe Probus smooth and glabrous, margin with 
tion of a leaf showing mar- double brown horny spines, not more 
ginal characters. Nat. size. than 2—3 mm long, with a distance 

of 1—3 cm between two pairs (Fig. 1); 
simple spines are to be found near the tip of the leaf only. 

Scape 2 m high, at the base 2,5 cm, at the middle 2 cm, and 
just below the panicle 1,3 cm broad, finely furrowed. 

The lower bracts of the scape 2 cm broad, 6 cm long, obtuse, 
their margins minutely denticulate (Pl. IV, fig. 2), the upper ones 
smaller, acuminate, entire. 

Panicle (Pl. IV, figs. 1, 3) about 80 cm long, thrice branched; 














sm sm 





— 307 — 








Fig. 2. Branch of the panicle with numerous bulbils and a flower (X 2/3). 


the single branches with small brown membranous bracts. Flowers 
solitary, with 1—2 bulbils (Fig. 2). These are of very varying 
sizes, the largest 3,5 cm long, 1,5 cm broad, very compressed, with 
3—d visible leaves, not densely united 
like a bulb (Fig. 6, 7). Bulbils also are 
developed in the axils of the upper bracts 
of the scape below the inflorescence. The 12 É 
flowers on pedicels 1/,—1 cm long, cer- W 
nuous, with 6 cream-coldured perianth- | 
segments and of a faint scent. The outer 
segments 1,4 cm long, 0,6 cm broad, the 
inner a little broader (Fig. 3, 4). Stamens 
much shorter than the perianth-seg- 
ments. Filaments 2,3 mm long, at the 
base 1mm thick, upwards subulate. 
Anthers 4 mm long, 1,5 mm thick, versa- 
tile (Fig. 5). Ovary 8 mm long, .2 mm 
thick, its transverse section obtuse tri- 
angular. Style 5 mm long, thickened at SE ig. ka KASER MORE 
the base. The flowers all barren; the SF R five (X 1'/,). 
fruit therefore unknown. Fig. 5. Stamens; back-view, 
Fee LEE front-view, and side-view 
One of the most striking peculiarities (x 3). K. Wiinstedt del. 
of the plant in question, I should think, is 
the very compressed bulbils, similar to the flower-buds with the two 
green bracts of Stratiotes aloides (hence the name of the species). I 
tried therefore, to compare these bulbils with those of other spe- 
cies of Furcraea allied to our species, but I found that in most 
descriptions given the bulbils were not mentioned at all. Only 


20= 





— 308 — 


in the descriptions and illustrations of the following species I 
find some information as to the shape of the bulbils. 





Fig. 6. Bulbil and a transverse Fig. 7. Front and side-view of 
section of it. (X 2). bulbil. (X 1'/,). K. Wiinstedt del. 


Furcraea cubensis Jacquin 1763 p. 100, tab. 175, fig. 28 shows a 


terete, bulb-like bulbil, and in the text it is also 
described in this manner. 

Cabuja Trelease 1910 Taf. 37. Terete, bulb-like bulbils. 

elegans Todaro. In the description in Hortus Panormi- 
tanus nothing is said about the shape of the bulbils. 

gigantea Vent. Some material in the Botanical Museum 
of Copenhagen, determined by W. Trelease, shows 
terete, bulb-like bulbils. 

macrophylla Hook. Trelease 1910 Taf. 37. The bulbils 
terete, bulb-like. 

Selloa K. Koch. Bot. Mag. tab. 6148 distinct terete 
bulbils. 

tuberosa Ait. Some material in the Botanical Museum 
of Copenhagen (det. W. Trelease) shows terete, 
bulb-like bulbils. Concerning F. lipsiensis (fide 
Drummond — F. tuberosa Ait.) Jacobi 1869 p. 169 
says, that the bulbils resembles ”"Gartenzwiebeln”. 

undulata Jacobi. To judge from Bot. Mag. 6160 and 
7250, these both represent the same species (fide 
Drummond), the bulbils are terete also in this 
species. 


RE 


Only in F. stricta Jacobi 1869 p. 171—74 the hbulbils are mentioned 
in the following words: ... "sehr zahlreiche ogi- 
vale, theils seitlich etwas plattgedruckte, theils un- 
scheinbar stumpflich dreikantige Bulben”. 

I think it probable that the bulbils of all species of Furcraea 
hitherto known, F. stricta only excepted, are terete, bulb-like, as 
possible deviations from that shape would certainly have been 
noticed by the authors. 

F. stricta resembles in many points F. stratiotes, though 
distinct differences are present. F. stricta is mentioned by Drum- 
mond (1907 p. 57) as being closely related to or even identical 
with F. elegans Todaro or F. macrophylla Hook. I am of opinion 
that Drummond is wrong here. I contend that F. stricta ought 
doubtlessly to be referred to Drummond's group A. Minores 
together with F. undulata. Unfortunately it will certainly be 
impossible ever completely to elucidate F. stricta (cfr. Drum- 
mondeo 0 psIn): 

I think that the species allied to F. stratiotes may be grouped 
thus when employing Drummond's system as shown in the follow- 
ing key: Sect. II. Spinosae. 

A. Minores. 

a. Bulbils terete, 13—30 leaves in the rosette 
F. undulata Jacobi 
(F. albispina Baker?) 
b. Bulbils more or less compressed, 30—50 leaves 

in the rosette. 

1. Bulbils slightly compressed, leaves rough below. 
Flowers large, about 30 leaves in the rosette (fide 
Bakers ss ps) FF SIE F.: stricta. Jacobi 
. Bulbils very compressed, leaves smooth on both 
surfaces, flowers minute, about 50 leaves in the 
FOSEBbEe TESS SE SE En F. stratiotes mn. sp. 

The terete, bulb-like bulbils is certainly a characteristic com- 
mon to all the species of the group B. Giganteae. Of the species 
of this group F. cubensis probably approaches nearest to F. stra- 
tiotes.  F. cubensis however distinctly differs mainly by 1) the 
terete, bulb-like bulbils, 2) the flowers being twice as large, 3) the 
broader leaves and 4) the simple spines in the margin of the 
leaves. 


Do 


The Botanical Museum of the University. Copenhagen. Febr. 1921. 


— 310 — 


Bibliography. 


Baker, J. G., 1888. Handbook of the Amaryllideæ. London. 

Drummond, J. R., 1907. The literature of Furcraea with a synopsis o 
the known species. Missouri Bot. Gard. Eigh- 
teenth Report. St. Louis. 

v. Jacobi, 1869. Nachtråge zu dem Versuch einer systematischen Ord- 
nung der Agaveen. Abhandl. der Schles. Ges. f. vater- 
lånd. Cultur. Abth. f. Naturwiss. u. Medicin. 

Jacquin, N. J., 1763. Selectarum stirpium americanarum Historia. Vin- 

dobonæ. 

Kunth, C. S., 1850. Enumeratio plantarum. Tomus 5. Stuttgart et Tu- 

bingen. 

Todaro, A., 1876—78.… Hortus botanicus Panormitanus. Tom. I. 
Trelease, W., 1910. Observations on Furcraea. Ann. du Jard. bot. de 
Buitenzorg. 3. Suppl. 2e Partie. 

Ventenat, C., 1796. Furcraea. Novum Plantæ Genus descriptum. An- 

nalen der Botanik herausgegeben "von Dr. Paulus 
Usteri. 19, p. 54. 


Explanation of the plate IV. 


Fig. 1. Furcraea stratiotes n. sp. showing the habit of the entire plant. 

— 2. — — — , the rosette with double-spined leaves, and 
the base of the scape with the lowest 
denticulate bract. 

— 3. — — — , the panicle with numerous compressed 
bulbils and some flowers. 





En ny Art af Furcraea fra Nicaragua. 


Af 


Johs. Boye Petersen. 


I Aaret 1848 rejste daværende Mag. A. S. Ørsted i Mellemamerika 
for at gøre botaniske Indsamlinger. Han hjemsendte store Mængder af 
tørrede Planter, der nu er noget af det værdifuldeste, vort botaniske Mu- 
seum rummer. Tillige samlede han Frø og levende Planter til den bota- 
niske Have, der den Gang laa ved Charlottenborg. De fleste af disse Planter 
er selvfølgelig forlængst forsvundne igen; men enkelte af dem er bevaret 
op til Nutiden og flyttedes til den nuværende botaniske Have i 1874. For 
nogle Aar siden blomstrede her for første Gang ÅAgave guatemalensis, og 
i Slutningen af 1921 kom endnu en Plante i Blomst, som stod under Navnet 
Agave sp. og stammede fra Nicaragua. Det viste sig, at denne snart 74 
Aar gamle Plante i Virkeligheden ikke hørte til Slægten Agave, men til 


— 311 — 


Naboslægten Furcraea; men den kunde ikke henføres til nogen af de kendte 
Arter af denne Slægt og beskrives derfor som en ny Årt under Navnet 
Furcraea stratiotes n. sp. Den har en Roset af stive Blade, der paa Randen, 
er besatte med ejendommelige dobbelte Torne, en 2 m høj Blomsterstængel 
der bærer en stærkt grenet Blomsterstand. Blomsterne, der er duftende 
og med nærmest flødefarvet Bloster, visnede alle, saa at den ikke satte 
Frugt; men i Blomsterstanden og langt ned ad Stængelen fremkom i Blad- 
hjørnerne en Mængde stærkt sammentrykte Yngleknopper. Disse minder 
noget om Blomsterknoppen hos Krebseklo (Stratiotes aloides), og herfra 
stammer Plantens Navn. Ved Hjælp af disse Yngleknopper kan det 
nu lykkes at bevare Arten og faa den udbredt i de botaniske Haver. Selve 
den gamle Plante derimod gaar ud efter Blomstringen. Nær beslægtet 
med F. stratiotes er F. cubensis (Jacq.), der hører hjemme paa Cuba, men 
dyrkes flere Steder i Vestindien, idet man bruger dens sejge Bladtaver 
som Tekstilmateriale. 








Typ. Bianco Luno. Kbhvn 





RK 


FAM 


Boranisk TIDSSKRIFT. 37. BD. 


PLATE IV. 


By 


Fig. 














høg 
å = E=3 
2 i ; 
så, TE gReG 
Cd ÅR 
= | 
ss JRÆR i | 
E i i | 
så z cs - 





Særtryk af Botanisk Tidsskrift. 37. Binds 5, Hefte. 1922. 


LIBRARY 
NEW YGRK 
BOTANKAL 

«3 A 


Studies on the Collective Species Viola 
brreolor ET 


By 
J. Clausen. 


SE | 
W 


I. The varying Characters of V. tricolor and V. arvensis. 


In the first paper I published on Viola tricolor (CLAusen 
1921), I mentioned 6 varying characters and the variations of 
those examined in nature. I stated that nearly all possible com- 
binations of 4 characters were found realized among 1000 individ- 
uals. I will here give a brief account of the variations I hitherto 
have examined in Viola arvensis-tricolor. As in the first paper, 
the abbreviations employed for the characters are shown in 
brackets. 

fÆærhefsrresor thepetals: 

Small petals (parv) = arvensis. 

Large petals (grand) — tricolor. 

Should one refer any individual belonging to the collective 
species V. tricolor, either to arvensis or tricolor (in a more 
restricted sense) only this character ought to be applied to 
the distinction. 

2. The stigma: 

Without labellum (n.lab): typical arvensis. 

With labellum (/ab): typical tricolor. 
SEAN ark spor mr ontlor thetstyle: 

Without spot (n.mac): typical arvensis. 

With spot (mac): typical tricolor. 

4. The pollen-magazine: 

Open (ap): typical arvensis. 

Closed (el): typical tricolor. 

Half-closed (sem.cl): hybrids? 


Arbejder fra den botaniske Have i København. Nr. 99. 


D. 


rave, 


The honey-streak: 

Without streak (erad = eradiata): occasionally in plants of 
hybridal origin. 

With non furcate streak (n.fure). 

With furcate streak (fure). 

Mhetdolourstorthepetals: 

Yellowish white (albid): typical arvensis. 

And the colour-variations of tricolor: Blue (vrol), Bright 
yellow (lut), Rose-red (or red-lilac) (rosea), a Pale blue (pall — 
pallida), Pure white (alba). 

The colours must be determined upon elder flowers. 


Most of these variations were mentioned in the paper from 1921. 


ve 


(ge) 


Nhetlens hore thes pr 

It is not expedient to state the length of the spur in absolute 
measures, because it is dependent on external conditions. 
In determining the length of the spur I have applied the 
ratio between the length of the appendices of the sepals and 
the length of the spur, because this ratio is very easy to 
determine. Both the size of the appendices and the length of 
the spur are varying dimensions, and they are in some way 
independent of each other, so that the ratio to some extent is 
the result of two varying characters. 

V. arvensis has a spur as long as the appendices or a little 
shorter. V. tricolor's spur is a little longer. But in the dunes 
of West-Jutland and Læsø, types of tricolor are found, whose 
spur is more than twice as long as the appendices (fig. 1). This 
ratio is due to both the long spur and the short appendices. 
As I have used this character to determine the variation of 
plants from dune populations, I have only distinguished the 
following two types: 


Short spur (brev.calec = brevicalcarata): the length of the 
spur twice the length of the appendices or less. 
Long spur (long.calc = longicalcarata): the length of the 


spur more than twice the length of the appendices (fig. 1 c). 
The for mor hes pre 


Spur straight (rect.calc == calcar rectum): the most com- 
mon type. 
Spur bent upwards (incurv.ealec = calcar incurvatum): 


most commonly found among the tricolor-types with long 
spur from the dunes. 





d e 


Fig. 1. Leaves and flowers from different types. (f/6). 
: V. arvensis typica from moor near Hillerød, Sealand: petals parv, leaf ovat-cren, stipules pinn. 
: V.tricolor typica from grass-field at Emmerlev, Southwest-Jutland: petals grand, leaf lanc-serr, 
stipules palm. 
ec: V.tricolor maritima from Læsø: petals grand, spur long.calc, parvifoliata. 
d: Leaf and stipules of V. arvensis from Tysinge Moor near Tølløse, Sealand: leaf ovat-cren, stipules 
pinn, foliac. 
e: V. arvensis from Tunø, population T, table I. Leaf and stipules very large, nearly as long as 
the peduncle; stipules pinn, foliac. 


søs 





Asta 
” 
E 


EDETNYA 





& 


9. The form ofthespur-bearing petal: 


KOS 


TREE 


12. 


— 367 — 


Commonly the spur-bearing petal both 
in arvensis and tricolor is truncate or 
faintly emarginate; but in the dunes of 
the northern part of West-Jutland and 
upon Læsø types of tricolor are found, 
where this petal is pointed (CLAUSEN 
1921, p. 209, fig. 1). I therefore disting- 
uish the following two types: 





Pointed (acum —= acuminata): fig. 2. 
Not pointed (n.acum = non acum- 

inata). Fig. 2 (1/)). 
The lateral sepals: V. tricolor. maritima 


I have found a special character of from Læsø: lateral 
these in some tricolors from dunes upon  Petals emarg, spur- 
Læsø. The lateral sepals are somewhat Bear IE PE FER SERSS 
incised at the tips: (fig. 2). This character was not previously 
known, and perhaps Læsø is the only place in the world, 
where it can be found; and probably it came into existence 
there endemically, by a mutation: 

Lateral sepals entire (integ — integerrima). 
Lateral sepals emarginate (emarg = emarginata). 
The epidermis-cells of the petals: 

The surface of the petals of the majority of pansies from 
the gardens is velvet-like. This character is due to the epi- 
dermis-cells, which are extended to many densely placed small 
microscopic papils (fig. 3a). The common wild types have 
not so large and so densely placed epidermis-papils, that the 
surface becomes velvet-like; but at Hadsund (Jutland) I 
have found a type of tricolor growing wild with velvet-like 
petals. This character I call velutina (velut). 

Common, not velvet-like petals: non-velutina (n.velut). 
The pollen-grains: 

In a dry state the pollen-grains are shrunken, but they 
are able to absorb water very quickly. Then they swell and 
assume a characteristic form: seen from the edge they are 
ellipsoidal with a protuberant equatorial-belt (fig. 3 6), 
but seen from above they appear either 4- or 5-edged (fig. 
3d;te);rless. frequently 3-= or 6-edgeds.(fig. 3'67 f). Tis the 
protuberant equatorial-belt giving them this edged appearance. 


15 


— 368 — 


The pollen-grains of V. tricolor are 4-edged, but further- 
more the same plant can have a few 3- and 5-edged. The 
grains of V. arvensis are d-edged, but the same plant can 
besides have some 4- or 6-edged or both. Finally, types are 
found that have just as many 4-edged as 5-edged 
pollen-grains. I suppose these to be of hybrid origin. 
The leaf-size: 

This character varies in a high degree, but as it is highly 
dependent on external conditions, the variations are not 





Fig. 3: 
a: Epidermis-cells from petals ("59/,). 
b: Pollen-grain seen from the edge. 
ec, d, e, f: Pollen-grains seen from above. 
ke Be 


easy to determine. The sizes must at least be stated in pro- 
portion to some other organ on the plant, for instance the 
length of the leaves in proportion to the length of the pe- 
duncles. Commonly the length of the full grown peduncles 
is about twice that of the leaves. 

However I have found another type of leaves in the dune 
populations. They are very short and narrow (fig. 1 e). 
These leaves have always been somewhat fleshy, 
too. The surface of a leaf of this type is only about ”/,—/, 
of that of a corresponding leaf of a common type. Fig. 46 
and c are leaves of parvifoliate resp. grandifoliate types at 
the same stage and grown under the same conditions. 

I found leaves of an extremely large size in an arvensis- 
population upon Tunø (an island in Kattegat) fig. 16. The 
leaves were just as long as the peduncles. Hybrids often 
have very large leaves. 


— 369 — 


The leaf with the largest surface is most favourable to 
assimilation, but it gives (all other conditions alike) the 
largest transpiration too. I have compared the epidermis of 
the leaves of the parvifoliate types from the dunes with 


bp) 








NZ 


Fig. 4 ("/,). 
V. arvensis from Koldby Kaas, Samsø (population U, table 1): petals 
parv, leaf lanc-serr, stipules palm. 
b: Leaf of V. tricolor maritima from Læsø: parvifoliata. 
e: Leaf of V. tricolor from fields: grandifoliata. 
The leaves b and ce were both from the rosette at the basis of the plant. 


=> 





a: 


those of the common types and have not been able to detect 
any difference between them in the number of stomata per 
unity of area. Therefore, the small leaves on the plants from 
the dunes must undoubtedly be favourable in reducing the 
strong transpiration in these places. Just in the dunes we 


find almost exclusively parvifoliate types. 


Botanisk Tidsskrift. 37. Bind. 24 


14. 


16. 


HEE 


— 370: — 


The form of the leaves: 
Two extreme types can be distinguished: 
Ovate, crenate leaves (ovat-cren): fig. 1a, common in 

V. arvensis. 

Lanceolate, serrate leaves (l/anc-serr): fig. 15, common 
in V. tricolor. 

But as these two characters are transgrediating, and the 
rosulate leaves are quite different from the upper ones, they 
are difficult to distinguish in practice. Sometimes plants 
can be found that have ovate, crenate leaves hut in all other 
characters are typical tricolor, and on the contrary plants 
that have lanceolate, serrate leaves can in all other char- 
acters be arvensis-like. 

The stipules: 

The most typical V. tricolors have palmate stipules (fig. 
16), but V. arvensis has pinnate stipules (fig. 1 a). As these 
two characters are transgrediating, I have provisionally 
distinguished the two types in the following way: 
Pinnate stipules (pinn): at least one stipule pinnate. 
Palmate stipules (palm): all the stipules palmate. 

Fig. 4a shows an arvensis with palmate stipules. 

The end-lobe of the stipules: 

In some types the end-lobe of the stipules is foliaceous 
and petiolate (fig. 1 d; the end-lobe in fig. 18 is foliaceous 
but not petiolate). When the end-lobe is crenate, it has the 
character of a leaf. 

Foliaceous end-lobe of the stipules: (foliac). The end- 
lobe not foliaceous (not crenate): (n.follac). 

In such stipules as those in fig. 1 d and 1ec the assimila- 
ting area is highly increased. The character foliacea can be 
found combined with either pinnate or palmate stipules. 
The growth-form: 

Most pansies have ascending stems, they are negative 
geotropic. But many of the types from the dunes of West 
Jutland have transversely geotropic stems. Already when 
the young plants leave the rosulate stage, the stems press 
themselves against the surface of the earth. Sometimes the 
pressure can be so strong that, when the plants are drawn 
up, the turgor bends the stems downwards with vigour (see 
fig. 5 b. In fig. dc a branch of the same plant is shown. On 





Fig. 5 (2/3). 

a: V.tricolor from field, S. Lyngvig, Holmsland Klit: erecta, n.atropurp, grandifol. (Population Y, 
table I). 

b: V.tricolor maritima from dune, S. Lyngvig, Holmsland Klit: prostrata, atropurp, parvifol, cae- 
spitosa. The light parts were covered with sand. The turgor has bent the stems downwards with 
vigour. 

c: A branch of b with flower. The light parts of it was covered with sand. The subterraneous parts 
are negative geotropic, the supraterraneous are transversely geotropic. 


24+ 





9 


— 373 — 


the surface of the earth the stem is bent horizontally at 
right angles). The subterraneous parts of stems of these 
plants are negative geotropic as fig. 53 b and c show. I di- 
stinguish two types of growth-form: 

Erect stems (negative geotropic): (erect). 

Prostrate stems (transversely geotropic): (prostr). 

At the beginning of autumn some of the most erect types 
bend their stems downwards horizontally, but the apices of 
the stems are always erect, and thus these phenotypically 
prostrate plants are easy to distinguish from the other gen o- 
typically prostrate. 
The colour of the stem: 

I distinguish 2 types: 
Plants containing a large quantity of anthocyanin 


(atrop = atropurpurea): dark violet stems. 
Plants with fresh green stems (n.atrop = non atro- 
purpurea). 


In the dunes of West-Jutland and Læsø nearly all the 
pansies are atropurpurea. The plants from the fields are 
usually non atropurpurea. The plants with albinotic flowers 
have pale green leaves and stems (chlorina), but as this 
character is coupled with alba, I have not taken it into 
consideration here. 

Duration of Life: 

Viola tricolor and arvensis are as a rule annual. They are 
Therophytes which regard the dry summer-time as the 
most unfavourable season. In many fields arvensis only lives 
from autumn to early summer (is winter-annual). V. tricolor 
can better survive the dry season. On the other hand, in 
dunes and in pine forests most individuals of Viola tri- 
color are perennial. An essential condition for perennity in 
our latitudes is that the plants either are able to produce 
subterraneous side-branches or are able to ramify strongly 
in the surface of the earth. Commonly Viola tricolor and 
arvensis lack this capacity in contradistinction to V. cornuta- 
calcarata. In dunes the sand-heaping and in pine woods 
the falling of acicular leaves replace this capacity of the 
plant, as the lower parts of the stems with their buds 
are covered by these agencies. From the covered parts the 
plants shoot adventitious roots and subterraneous branches 


(fig. 56, c). The subterraneous leaves are white and scale- 
like, without any resemblance to the supraterraneous. 
Consequently many of the perennial pansies from the dunes 
and pine forests are merely phenotypically perennial. 

But the perennial types from the dunes of West-Jutland 

and upon Læsø possess some characters which better enables 

them to survive the unfavourable seasons and conditions, 
and therefore these types are genotypically perennial. 

These characters are: 

a. A very high degree of ramification (the character 
caespitosa) in the earth-crust, exceeding many times that 
of the common Viola tricolor and sometimes that of cal- 
carata-cornuta too. Fig. 5 b is a very moderately ramified 
individual. It is not uncommon to find individuals with 
more than 20 and even up to 50 side-branches. 

b. Transversal geotropism (prostrata), common among 
the individuals in these dunes, is a character that facili- 
tates the covering with sand. 

c. Transpiration is reduced, partly by the character 
parvifolia and partly by prostrata. This reduction is of 
great importance for the survival during the unfavour- 
able season. 

d. The plentiful Anthocyanin in the leaves and stems of 
these types (the character atropurpurea) is supposed to 
be a protection against too much light. 

Therefore the perennity is most frequently the result of 
a co-operation of the named genotypically conditioned cha- 
racters, but as the first one is the most essential, and besides 
the only one not mentioned before I here distinguish only 
two types and determine them according to the ramification: 
Perennial (caesp = caespitosa): high degree of ramification 

in the earth's crust (more than 6 side-branckes). 

Annual (n.caesp): When not covered, only side-branches 
above the surface of earth, the side-branches not 

rosulate and sparse. 

The two types are transgrediating. 


— 375 — 


As will be seen from the foregoing, the characters in several 
cases are rather difficult to distinguish from each other, and the 
list here given is not exhaustive. As for stipules for instance 
there are more types than the four mentioned, but on the other 
hand each V. tricolor or arvensis can as regards stipules be placed 
under one of the 4 types. The limitation of all the quantative, 
transgrediating characters against each other is also difficult, but 
I suppose that a limit, even an arbitrary one, is always to prefer 
to a description, in common terms, of the habitual appearance, 
that never can be so concise and so easy to compare as the state- 
ment of a numeric relation. 


II. Are the Variations induced by Genotypical Differences? 


This problem I have tried to solve in different ways. 

HEE Bsysko bier va bro nen re SnSmnany habitat ses found 
the different characters realized in plants growing together 
under the same conditions. Plants with violet petals and 
yellow petals, with long spur and short spur, with straight 
spur and incurved spur, with erect stems and prostrate stems 
grow intermixed in the same habitat, and when they are 
different, it must be due to the genotypical differences. The 
tables I and II (p. 378 and 381) state the extent in which the 
different characters found are realized in the same habitat. 

2. By cultivation of the types under the same con- 
ditions. In several cases I found marked differences between 
the composition of the populations from habitats the con- 
ditions of which differed considerably. On the narrow isth- 
mus Holmsland Klit between Ringkøbing Fjord and the 
North Sea I found two such populations. A road runs length- 
ways through the isthmus and separates the dunes on the 
western seaside from the clayey and more fertile fields on 
the eastern fjord-side. There is but a few steps between 
these two localities. Both in the dunes and in the fields 
there were populations of Viola tricolor but of two very 
different types what can be seen in fig. 5. In the dunes all 
the plants were parvifoliata (leaves of the thick, somewhat 
fleshy type common in the dunes of West-Jutland), the stems 
and the leaves were atropurpurea, most of them were prostrate 
and caespitose perennials (fig. 5 b, c). Otherwise in the fields 


Do — 


on the other side of the road: all the plants were as fig. da: 
grandifoliata, non atropurpurea, erecta and non caespitosa (Table 
I, population Y, page 379). Apparently the characters of the 
type from the dunes were favourable to the plants under 
the conditions of life there. One might presume that the 
diverging appearance of this afore-mentioned type was a 
modification induced by the external conditions. The most 
exact method to prove whether the extreme characters were 
phenotypically or genotypically induced would have been to 
take the plants from these populations and propagate them 
in clons under identical conditions in order to see if they 
became identical. I did not do so, but instead of that I collected 
seeds from the two types and these were sown under identical 
conditions in the Botanical Garden, Copenhagen. They bred 
true to type: all plants of seeds from the dune became parvi- 
foliata, atropurpurea and all plants of seeds from the field 
became grandifoliata, non atropurpurea, the offspring being 
just as different as the mothertypes in fig. 5a and b. By 
this it was proved that the extreme characters were hereditary 
and not induced by the conditions. 

All the above mentioned characters I have tried under 
cultivation, and they proved to be genotypically conditioned. 
Sometimes a segregation was noticed: some of the prostrata 
plants taken in nature are homozygotic and some are hetero- 
zygotic, and therefore seeds of them show segregation in 
erecta and prostrata. — Seeds of an albinotic tricolor gave 
many albinotic and a few violet plants; (alba is recessive, and 
the violet plants were from seeds pollinated with pollen from 
violet flowering plants). — The first plant cultivated with 
the character emarginata (lateral sepals) was homozygotic 
and gave by self-fertilization only emarginata individuals. 

On the whole, most of the types collected in nature show 
no segregation except in such characters as the flower-colours 
lutea and violacea (tricolor) and in the characters maculata 
and non maculata (the style); likely they are homozygous 
in most characters. The probable cause to this phenomenon 
will be discussed later. 

By researches in Genetics. The different characters 
have been crossed in order to determine the factors and the 
limit of the characters against each other. It will still take 


— 377 — 


some time to complete these investigations, but they show 
interesting facts. They confirm the results attained by the 
two other methods, that the variations of the characters 
are induced by genotypical differences. 


III. Are all Combinations of the Characters possible? 


In the paper from 1921 I reported about the combinations 
of the 6 first mentioned characters. In the meantime, I have 
examined a great number of individuals from different habitats 
in order to decide if all combinations of the remaining characters 
are likely to occur, and to get information as to which combi- 
nations are the most frequent. The characters were examined in 
sets of 4, d or 6 at a time. Within the sets practically all combi- 
nations can be found. Characters from different sets have been 
examined together too and, as they also combine in different ways, 
I suppose it justifiable to infer that, on the whole, all combina- 
tions can be realized. Of course I have not seen all possible com- 
binations of all these characters. It would take more than a lifetime 
to examine only one individual of each combination, and I do 
not believe that they are realized all simultaneously. 

In table I next page a summary is given of the combinations 
of four characters that are easy to determine even on exsiccata. 
The results from 23 habitats are given in the table. In each place 
50 or 100 individuals were taken and determined. The numbers 
im the table are egalized so, that they in all cases indicate the 
number of individuals of the combination (isoreagent) in question 
per 100 individuals from that habitat. The four characters are: 
size of petals, colour of petals, stipules and end-lobe of stipules. 
The order of the characters is not indifferent; they ought to be 
taken in the afore-mentioned order, as the bulk of the individuals 
by this method is placed in two groups, one at the top and one 
at the bottom, corresponding to the two species in question: 
V. tricolor (at the bottom) and V. arvensis (at the top). In the 
middle the fairly rare irrelevant types are to be found. As to the 
flower-colours I have only taken two alternatives here, as there 
is a great difference between the arvensis-colour albida on the one 
hand, and all the tricolor-colours violacea, lutea, rosea and alba 
on the other. 

Each of the 23 habitats has its column in the table indicated 


— 378 — 




















































































































Table 
| | 
Reaction of soil basic | neutral 
Size of | Colour IE ; End-lobe | el, 
petals | of petals | Stipules | of stipules| 4 B CEED | E F | G | H | I J K I 
I 
|! | 
oe foliac | » | 66 SNGERS RES 6) HEY RRS ST BD » 2 »|  » 
væ | m.foliac | X | 34| 92! 76) 36! »|| 15) x | 81 14) 34) ») 
albid | | | ; | 
| tigt jnA foliac | » »|  » » SKEDE > y » ”» » » 
| z n.foliac VERS SØN Sy 6 Ste D NDE 8 2 » » || 
parv | . | | 
|| | | 
præ fohace | » KAN » » Snes | RSS as » » » » | 
7 7 I | | | 
viol, | me folidel Eye ” SERENE AES DISSE EEG: 4 st kere | 
| | | | 
HD | palm foltac soen » » SEN SDS ha) »] 0» » » 
| n.foltac ”» | ”» » Hedes en] ED ES« (85456 SHEET dør | 
I - I - . : i | 
pinn folraec » » » » » » || BIDER, » | » ” | » || 
| |m.foliac|| »|  » » » SER KERES ERE] rr » » || 
albid | | ; | || 
palm | foliac » »l 5» RL sa Er »)| 5 » » » » 
å | m.folrac|| »| »| » SSD 950) BREDT SEERE SER 559 » > || 
gran == == = ; : - - - | 
| pinn folnact || 9 » NET 559 HE len » » 6 » » 
bol; sl] Ensfoliae (REE BE HSG ES SD eN RA Ro el 
lut se] gs | | | 
555 foliac | » ”» SAGE HM er DSE] RE NREE fanen » » 
| | m.foliae |] »| »| »| »| »|x || »| »| 48) 28) 51) 44! 
| ø | i Tad ll | || 
| x 1100/100/100/100| x 1100! x |100 100 | 100 100! 
| | | 
Populations: 
A: Grass field at Onsild, Mid-Jutland, basic, marle strate. 28th May 1921. 
B: Grass field at Andrup, Mols, East-Jutland, basic. 2nd June 1921. 
C: Grass field near Skørping, East-Himmerland, basic, marle strates 28th 


May 1921. 

D: Grass field at Bælum, East-Himmerland, lime countries, basic. 31st May 
1492se 

E: Rye field on lime-hill south of Smidie, East-Himmerland, basic. 31st 
May 1921. 

F: Grass field on Holmsland, West-Jutland (marle), basic. 25th May 1921. 

i: Grass field between Grønfeld and Egens, Mols, East-Jutland, neutral. 
2nd June 1921. 


H: Sandy grass field near Vorgaard, East-Himmerland, neutral. 31st May 
92de 

I: Sandy grass field at Vejrumhede, Mid-Jutland, neutral. 18th May 1921. 

J: Sandy grass field near Terndrup, East-Himmerland, neutral. 31st May 
99 SE 

K: Grass field south of Hadsund, East-Jutland, neutral. ist June 1921. 








— 379 — 


































































































cz 


OLAFS 


: Grass field east of Vejrumhede, Mid-Jutland, faintly acid. 
: Sandy cleared space in plantation of spruce fir, north of Hadsund, East- 




















KE 
faintly acid acid | not known 
: | Summa 
M N | o P | R S ne IS RER | v 5< Y 
el RS » ”» » » 64 » || » ) » | 281 Ea 
SE EDR REE STE RES SRRR SET DE SSSREDN FS: BESES BÆRE FEE TAN RE 
| | S— == || | 2 
x » » || » » » » Wæl| » » » » 
|| 0 
» 6 » ; » » 6 X (13) | 6 » » 60 | Øg 
» » » » » » » » | » » | » » c 
» al, » » » » » » | » DE » 25 PR | 
| |; 246 
”» ” ” » » » » » | » » » » 91 
2 10 » » » » » BE » » » 21 a 
» » » | » » » » | » » » » » 
”» » 6 | » , » » ) » » » 6 É 
| = 12 
» » 2 RED » » » | » » | » » 2 | 6 
D 9 » | » » » » » || » | » » 4 
» » | » | » | » ”» » » |) » | 54 » 6 449 
BE 40 34 NED MED ster 5 8 Se far or 16 107 84 1542 1544 
| | | | FEE ET U= FEE BEES 
» 2 » | dl eg: ” » » || » | » | » 2 681 
bGE HEER HE50N 668 HE7 0 78 55) FAN KEE UR GS PAR RES SR SE 2 
OG ET OOS ER O0 | 100 |10011001100| x || 100 | 100 100 " 1900 1900 | 1900 








: Grass field at Assentoft, east of Randers, East-Jutland, neutral. ist 


June 1921. 
: Grass field east of Viborg, Mid-Jutland, faintly acid. 28th May 1921. 
28th May 1921. 


Jutland, acid. 3ist May 1921. 


: Sandy grass field at Oxbøl, west of Varde, West-Jutland, acid. 24th 


May 1921. 


: Uncultivated sandy Anthoxanthum-Scleranthus formation in Mols Hills, 


East-Jutland, strongly acid. 2nd June 1921. 
Sandy grass field near Issehoved, Samsø, strongly acid. 3rd June 1921. 


: Stony rye field at Tunø harbour, strongly acid. 3rd June 1921. 
: Sandy grass field at Koldby Kaas, Samsø, strongly acid. 4th June 1921. 
: Sandy grass field near Røde-Kro, South-Jutland. 21st May 1921. 


: Dune field at Selvig, Samsø. å3rd June 1921. 
: Grass field between N. and S. Lyngvig, Holmsland Klit, West-Jutland, 
2bth May 1921. 


— 380 — 


Characters: 
Size of petals: Stipules: 
parv = petals < the sepals. pinn = pinnate. 
grand = petals > the sepals. palm —= palmate. 
Colour of petals: End-lobe of stipules: 
albid = yellowish white. foliac =— foliaceous. 
viol, (lut) =— blue or one of the n.folitae = not foliaceous. 


other tricolor-colours. 


by capital letters, which refer to the explanation below. Each 
of the 16 combinations or isoreagents has a rubric in the table, 
and the number per cent is stated off each habitat. In some 
cases the number of individuals from a special habitat has been 
too inconsiderable to be stated per cent, and sometimes no stati- 
stics have been made. In these cases the Isoreagents noticed 
in the habitat are indicated by a cross. 

In some of the habitats only arvensis-like combinations are 
found, in others only tricolor-like combinations, in others again 
nearly all combinations are represented. Parv, albid and grand, 
viol are the two most common combinations, respectively 712 
and 1130 individuals from 1900. The combinations parv, viol and 
grand, albid are rare (only 46 and 12 out of 1900). 

Table II next page shows the combinations of some of the 
characters marking the tricolors from the dunes of Læsø and 
West-Jutland. Mr. C. A. JØRGENSEN has been so kind to make 
up the lists Z, Æ, Ø from Læsø in 1921, as it was of interest for 
me to know in which combinations the character emarginata was 
able to occur. (I observed this character in summer 1921 in 
plants from seeds I had collected on Læsø 1919). For the popu- 
lations AA and BB only form of stipules, length of calcar and 
colour of stems are indicated. 

All the Viola in the dunes are grand, lab, non albida (viol) 
and certainly parvifoliata. The last mentioned character is difficult 
to compare from different localities, except by cultivation under 
the same conditions. Most of them are atropurpurea too. But 
im other characters all combinations are realized. 

All combinations of characters, I have tried, have proved 
the same, and there is a multiplicity of possibilities for variations 
within the collective species. 



























































— 381 — 
Tablet: 
'Stipu=| | Lateral | 
Petals sene Calcar fed Z Æ Ø ||Summal Stems | AA | BB 
I 
| rect unteg g i 8 AL n. atrop 4 
i: : | emarg || 2 81 høre UR Baek: ; 
rev. ca Sj] eds AR 
. | | 
|incurv| eg | dit é atrop 301: 36 
ends | ENGANG AED RAS KE RS 6 | 
| pin ESSSSSSETNEND RE 18153555 == === == == 
| rect integ 2 | » » 
|; i emarg we URETS » 
tlongscalej—= - FS ren Fr 
| | £ | 2 ) 8 20 
grand, | | INCUTV | unteg É | B y 
K | | emarg NE 12 » 
BE ER er ER ae ES FE PO er NDA SEA 
(lut) rect | ”Pteg 116574 be SA fra 7 
b i emarg | 16 16 » 
rev. cale SEER SEN | NESA (eg fald SR NOGEN, 
re hemnieg 16 VEL AEG 
| ' | emarg | 10| 12! » 
| palm | = mme RR 
EN integ 16 » » 
| ka Aa) emarg 8 4 » 
; 9 É; 
|incurv| "eg = val 36 Så atrop | 10 ”» 
| emarg 41 20 4 28 
ES ESESLTSRLLrL Se NE SE eee 
100 100 " 100 300 100 | 100 
lo acum... 74 | 84 | 100 


Populations: 


Zi, Æ, Ø: Dunes on Læsø, 12th July 1921 (C. A. JørGENsEn), Z nearest to 
Højsandene and Ø nearest to Vesterøhavn. 
AA....: White dune at Nymindegab, 24th May 1921, many rosea. 


BB....: Grey dunes from Ringkøbing Fjord to Nissum Fjord. 26th May 
1921. 
Characters: 
Stipules: Lateral petals: 
pinn —= pinnate. integ = entire. 
palm — palmate. emarg = emarginate. 
Galcar: 


; Stems: 
brev.calc: calcar twice as long 


2 atrop = atropurpurea. 
as the appendices or less. P are p 
i n.atrop = not atropurpurea. 
long.calc: calcar more than twice 
as long as the appendices.  Spur-bearing petal: 
rect = calcar straight. acum — pointed, (the remaining 


incurv = calcar bent unwards. were not pointed.) 


— 382 — 


IV. The cause of the variation. 


The multitudinous different isoreagents (geno- 
typically conditioned variations, microspecies) within 
the collective species Viola tricolor L. have arisen from 
crosses between V. tricolor and V. arvensis and between 
the segregation products from these crosses. 

In the paper from 1921 I proposed this theory, and I am 
now able to confirm it. The evidences are threefold: 


1. Analytical evidence. 


Seeds of plants from populations with both arvensis, tricolor 
and the intermediate types give by sowing not only the mother- 
type but also the other combinations. This is characteristic for 
hybrids. An instance will show this clearly: 

In 1921 I collected seeds from the population F mentioned 
in the paper from 1921 (p. 210—211, table I and II), a stone- 





















































IReUSNLOSIEENE 
— - 
F VEPAGE RVE2 ÆT 
| 
; y2 D ) ; 
| TEE n.mac il SNE » arvensis 
Mac Pol! 4 4 
| . ; 9 PER > 
HG ODDE HEELS 2 
parv | mac 3 TÆEE 0 the mother type 
| SNS. 
mac ) AN ) 
lut md 2 
| mac » SUN » 
| lab, viol, mac 2 » » 
| z n.ma ifl | 
EET ER n.mMae i » 
| mac 2 il JÆS 
' | | 
|jeEr Mac 15) | 
n. lab | viol nd | 1 GEN 
| mac &7| ts Bd) 
| | | | re | 
grand Fess | n.mac » SÆN » 
| mac il » » 
| albid, mac 6 | » ”» 
Laos | Tet « | 
: n.maec rn TE 1 6 
viol | Rg | (KARA 
| | mac 18 SSÅNET 1 | tricolor 
| | 
| 100 AL ARM HEbe 7] 
| 


— 383 — 


quarry at Frederikshavn (see table III F in this paper). The 
seeds were derived from uncontrolled fertilization in nature. 
Seeds of two of these plants, both parv, n.lab, viol, mac (the size 
of flower was that of arvensis, the colour was tricolor's) were sown 
as V. 246 and V. 247. The result can be seen from table III. 

When these four characters are taken into consideration, the 
two plants have given, besides the mother type, 9 different com- 
binations and among these one that I did not find in the original 
population (parv, lut). Some of these must be due to segregation 
and some of them to cross-fertilization, but in both cases it sup- 
ports my theory. Moreover, from populations with arvensis or 
tricolor alone I have not obtained such segregations. 

Some of the segregated types were selfed in 1921 and they 
gave in 1922 the following results: 


V. 414: Mother plant: grand, lab, lilacina, eradiata, n.mac. 
petalis latis (in all a very peculiar type, that I have not seen 
in nature) gave only 9 plants, but they were all just as the 
mother plant. 

V. 415: Mother plant: parv, n.lab, lut, n.mac, petalis angustis gave 
8 plants all of the same type as the mother plant, but the 
size of the petals varied somewhat, as some of the plants had 
petals of the size of the sepals; other plants had smaller petals. 

V. 416: Mother plant: parv, n.lab, albida (with a hue of violacea), 
mac segregated the following types (table IV): 




















kabler HV 
| albid 24 
parv n.lab | lilac 2 All 36 individuals were mac. 
| lut 1 Some of the plants had broad 
TA DIGG] 9 petals and some of them nar- 
- row. 
STED albid | 1 lilaa = lilacina (a hue of 
rand bla HE 3 violacea). 
lab albid | » 
lilac 3; 





V. 417: Mother plant: grand, lab, viol, mac, petalis latis gave 11 
plants all as the mother plant (tricolor-type). 

V. 418: Mother plant: parv, n.lab, albid, mac segregated the 
following types (table V): 


— 384 — 








breve 
albid | n.mac | 7 
fr sggge sl uds 
parv, n.lab | | 
| 
Fler n.mac | 0 
Mae | 4 





24 individuals (7 n.mac: 17 mac). 


Some of these individuals have apparently become rather 
constant already and others segregated further. 

All this is an evidence of the hybrid origin of the two mother 
plants to sowings V. 246 and V. 247. 


2. Synthetic evidence. 


As mentioned in my first paper, I crossed different types of 
V. tricolor with different types of arvensis. F; was intermediate 
in its appearance (in some characters there was full dominance). 
F,, obtained in 1922, presented just the same multitudinous types 
as the populations from nature which I had suspected to be 
segregations from crosses between arvensis and tricolor. Such a 
segregation-field resembles just one of the populations I, J, K, 
M, N or O in table I, page 378. (The detailed results from these 
experiments will be published later on, when the results of the 
inquiry into the behaviour of the chromosomes after the hybridi- 
zation can be stated). 


a 


3. Cytological evidence. 


In the preliminary note I stated that tricolor had 13 chromoso- 
somes. Ås for arvensis, 17 chromosomes were found in several 
cases, but also types with 15 chromosomes existed. The further 
investigations have revealed that, as a rule, all relevant and stabil- 
ized types of tricolor and arvensis have respectively 13 and 17 
chromosomes. 

Fig. 6 a shows the homotypic metaphasis of an arvensis (parv, 
albid, n.mac, pinn, folitac) from Tunø (population T, table I, 
pag. 379). The type of the plant was like fig. 16, pag. 365. The 
divisions here goes on very regularly. 17 chromosomes. 

Fig. 6 bshows the diakinesis of a tricolor (grand, pall, mac, pinn, 
caespitosa) from Selvig, Samsø (population X, table I). Also 


< så A j F: Beer SS ; BD. 
ET ESS FONG AA 
LA ak BEN KERES 
SEE sed gs rr re 


Hiss GR) 
a: V. arvensis (population T, table I) from Tunø: homotypic meta- 
phasis in a pollen-mother cell. 17 chromosomes. 
b: V. tricolor (population X, table I) from Selvig, Samsø: diakinesis in a 
pollen-mother cell: 13 chromosomes. 





ØE ÆRE 
FE, UN As 
; 6 eg ) É | 
f ud SIG | É e 2 eet i 
| j 
| 3" e i 9, & 
| [Na de FS ) 
X 9 Å rn 7 
EET 1 < ss c 
Pa BY 
£ ” = 
g ” By i 
Fz x Å S; j 
f rs » lad eg å 
j É g: e 2, 
7 | Pe & ; ae Se x 
Na 3 
ik sr 6 bs ST i 
xx 
FE] 
sæ Fon, sæ KÆ 
Pr i ; dg 
[ d KS 
xx 1: 88 eg ] D Pr 


.£ 
& 
ø. 
& 
e. 
e 
X 


RE eee > 1 


Fis) 
: Reduction division in a spontaneous hybrid parv, lut. (V. 246—14). 
: homotypic metaphasis: 13 chromos. 
: heterotypic metaphasis: 14 chromos. 
: homotypic metaphasis: 15 chromos. 
: early heterotypic anaphases seen from the pole; varying number 
of chromosomes. 
early heterotypic anaphasis in artifical hybrid parv, lut (F:). 


Botanisk Tidsskrift. 37. Bind. 95 


Ya a SR 0 


z 





— 387 — 


the division of this is very regular. The conjugation between the 
components is end to end and is complete. 13 gemini. 

The intermediate types from the mixed populations are not 
so regular in their divisions, as shown in figs. 7 a—g and 8 a—e. 
It is the reduction of a plant from sowing V. 246, table III, p. 382 
i. e€.: a plant segregated from the very mixed population F in table 
III). Its formula was: parv, n.lab, lut, n.mac, pinn, foltac (arven- 
Sis” size of petals, but the colour of tricolor). It was the plant sown 
as V. 415 (p. 383). By its irregular divisions it discloses its 
hybrid origin. 

In the early heterotypic anaphases seen from the pole most 
frequently 14 bivalent and 1 or 2 single chromosomes can be 
seen, as the following schedule over the pictured nuclei shows: 









































; univalent | | 
bivalent | (marked X) | upper plate | lower plate 
Eir27%d 14 1 (large) |15 chromos. |! 14 chromos. 
geder 13 HE ERNE oe En SENER 
2 little 
Ft 14 IE TATE ESS Ejes ER GE br gere 
1 little 
—  g| 14 1 (little) 14 — 15 — 
| (the two marked X X 7 
only faintly joined) | | 
— === TE == — Fr = === - SITE =— — — =-— = — == = == 
es sk is DSE, VE vA I N SER HE SEN 
| | lybrid 











The chromosomes distribute irregularly to the poles, so that 
the anaphase-plates contain different numbers of chromosomes. 
One chromosome more or less seems not to afflict the viability 
of the plants. Just as irregular are the artificial hybrids. In the 
same anther nuclei with 13, 14, 15, 16, 17 and even with 11 and 
12 chromosomes can be seen. Fig 7 h is of a beginning hetero- 
typic anaphasis from F, of a cross tricolor X arvensis (13 chromos. 
X 17 chromos.). In the upper plate are 14 chromosomes and in 
the lower one are also 14, but one from each (marked X) is single 
and not conjugated, just as in the spontaneous hybrid. 

Fig. 8 shows the homotypic anaphases of the same spontaneous 
hybrid. The following schedule shows the irregularities in this 
division: 

25% 


— 388 — 








Chromosomes lost 





Fiz. 3 Number of | 
g. 








| chromosomes " in heterotyp. in homotyp. | 
| division — | division 
Tetrad a | 14— 14 Bl | Cas 
Søg b sketetre || ASUSers 0 | 2 
— .c ble Il 
Ade eet gt | VEG 
— €e | 4 + 4 
er, DELE 1 VERS 8) Artificial hybrid 


There is a large accordance and similarity in the nuclear 
divisions between the artificial hybrids and the suspected, spon- 
taneous hybrids. 

When 13 tricolor-chromosomes and 17 arvensis-chromosomes 
are brought together, they behave in different ways in the hete- 
rotypic metaphasis. They can form either 15 bivalents, 14 bivalents 
and 2 univalents or 13 bivalents and 4 univalents. The most 
frequent is 14 bivalents. We must suppose that 13 zricolor-chro- 
mosomes join with 13 arvensis-chromosomes and, in addition, 2 
arvensis-chromosomes join with each other forming the 14th pair. 
The univalent chromosomes in some cases go to different poles, 
in other cases to the same pole both, and in other cases again 
one or more of them are lost in the first or in the second division. 
Most frequently they are lost in the homotypic division. We 
must suppose that they can be lost during the vegetative mitoses 
too (Fig. 7d, g and hk). As I never yet have found tricolors with 
less than 13 chromosomes, I must suppose that pollen or eggs 
with less than this number, or the zygotes with less than 26 chro- 
mosomes degenerate. 

This type of distribution of chromosomes after crosses was 
not previously known. The only one it resembles is the distribu- 
tion in F, of a cross between Solanum nigrum diploid x tetraploid (72 
chromosomes x 36 chromosomes) according to a discourse deliv- 
ered by H. WINKLER in the Deutsche Gesellschaft fir Vererbungs- 
wissenschaft (H. NAcAHTSHEIM 1921). F; of this cross in its vege- 
tative cells has 108 chromosomes, but in the reduction division it 
exhibits 54 bivalents, so that all the 36 chromosomes of the diploid 
parent must unite with 36 from the tetraploid and the remaining 
36 from the tetraploid must unite mutually to form 18 pairs. 





Fig. 8 (2500/,), 
a-e: Homotypic ana-telophases in spontaneous hybrid parv, lut. 
(V. 246—14). 
f: Homotypic telophasis in artificial hybrid parv, pall (F,). 


A 
RR 
| 
e BD 
j a?e 2 2 | ' e ap & 
98 | (> 
| e…2 na 
| de 2 8? | Å 
i s… e AR e | 
2 / Se | 
i; & / 8 7 
/ & Eg 4 Se e? 
a NR ERE bb Cc 





Fig. 9 (2500/ ), 
a-b: V. tricolor nana. 
a: Interkinesis; 2 chromosomes are lost. 
b: Homotypic metaphasis, regular. 13 + 13 chromosomes. 
V. tricolor lutea; homotypic metaphasis. 12 + 12 chromosomes in 
the nuclear plates + 2 lost in heterotypic division. 





— 391 — 


F, of this cross agrees with F, of tricolor X arvensis, as the eli- 
mination of chromosomes in Solanum does not take place before 
in F,. In one F,-individual RosE STOoPrPrEL found 45, 41, 40, 38, 
37, 36 and in another 54 and 55 chromosomes and so on. Both 
in Viola and Solanum some of the bivalents must be formed by 
ehromosomes from the same parent. In Solanum F, is regular, 
but in Viola the irregularities begin already in F2. 

As usually only the two chromosome numbers 13 and 17 
occur in nature, the chromosome-sets of the hybrids must for 
some reason or another be unstable, or the plants with these 
combinations of chromosomes can not be fit to the strong com- 
petition in nature. It is likely, that the offspring of a hybrid 
sooner or later ends in either a tricolor or an arvensis, i. e. in a 
13- or in a 17-chromosome plant. But the chromosome-set of a 
tricolor that has come into existence in this way need not be 
identical with the parental tricolor. Likely a chromosome-ex- 
change has taken place between the two chromosome-sets, so 
that the new tricolor has chromosomes both from the parental 
tricolor and the parental arvensis and in the same manner the 
new arvensis has chromosomes from the parental tricolor too. 
Consequently the factors carried by these chromosomes must be 
transferred together with these, so that we must expect to find 
tricolors that show some of the characters of arvensis and vice 
versa. In many places it is very difficult indeed to find a Viola 
typical in all its characters. 

Even in the stabilized types, the reduction division can be 
irregular in some cases: In my cultures I have a curious little 
dwarftype. In 1919 I found in Sophienholm Hills, Sjælland, 
(CLausEen 1921, table I- and II, population G, p. 220—221) a 
plant which I determined as parv, pall. It had very narrow petals 
and, according to its type, I expected it to approach arvensis. 
But, under cultivation, I observed that the petals or the sepals 
might vary slightly, so that the petals usually were a little longer 
than the sepals. It had a small labellum. I expected a segregation 
similar to sowing V. 246 and V. 247 p. 382, but in two generations 
it has bred true to type. All the offspring were nana-individuals 
with narrow petals slightly longer than the sepals, but the absolute 
size of them was less than that of many arvensis. The time of devel- 
opment to the flowering stage is the same as arvensis”, only about 
1//, month; considerably shorter than tricolor's (about 2"/, months). 


— 392 — 


All the plants have been mac and faintly labellata. Regarding 
the colour of petals, segregation has taken place, but only in 
tricolor-colours: pall and lut. Apparently some of its characters 
caused it to be referred to tricolor and some to arvensis. Å cyto- 
logical investigation showed that it usually had 13 chromosomes 
(hapl.) as tricolor (fig. 96). But in some of the anthers the divi- 
sions were very irregular, and most of the pollen from these 
anthers degenerated. A rather regular interkinesis of this sort is 
shown in fig. 9 a. 2 chromosomes here are lost in the first division. 
Possibly all the irregular pollen degenerates and therefore the 
type can be constant. According to its chromosome number I 
regard it as a tricolor, but it was certainly established through 
a segregation after hybridization and it has inherited some of 
tricolor's and some of arvensis? characters. 

One of the bright yellow tricolors (grand, lut, n.mac), a well 
established type that only segregate the two types pall, mac and lut, 
n.mac shows similar irregularities in the reduction division. The 
chromosome number is 13 haploid, but in a few cases two chro- 
mosomes are expelled in the heterotypic division. Fig. 9 c shows 
the homotypic metaphasis in such a dyad. Each of the nuclear 
plates has 12 chromosomes, and between them are seen the two 
expelled chromosomes. Apparently there is some instability in 
the chromosome stock. The cause of this instability is probably 
a preceding cross. I assume that all the bright yellow tricolors 
descend from crosses between tricolor and arvensis. 


V. Are all the genotypically conditioned Combinations 
(Isoreagents) to be regarded as ''Species?”? 


In Viola there is a multitude of possible types. When crosses 
are possible and give fertile offspring, and when the variation is 
caused in this way, we may expect from time to time to find 
all the combinations of the characters realized. With the clas- 
sification here adopted thereis 2.2.2.3.3.6.2.2: 2.2. 237. 
ORE NO ORD NOD BO SAGE possible com binationsske Butkas 
there are more characters varying than the former (for instance 
broad petals and narrow petals, but difficult to distinguish from 
each other), there are still more combinations. The number of 
possibilities will increase very rapidly, when new varying char- 
acters are found, and in the same proportion the likelihood of 


— 393 — 


finding them all realized during ones life-time will decrease. It 
is practically impossible to make a classification of so 
many systematic units. 

But many of the individuals are heterozygotes, their offspring 
is inconstant and therefore they have no systematic value, some 
will say. It is a mere assertion and only supported by custom, 
that constancy in offspring is the real standard for systematic 
value. The ecological value of characters as prostrata, atropurpurea 
and caespitosa is the same, whether they are heterozygotically 
or homozygotically realized. When the purpose is to classify the 
individuals, it is more to the point to take a standard that has 
some relation to the individual itself and not to its offspring. 
The identical reaction against the conditions is such a standard. 
All individuals reacting identically against the conditions are of 
the same ecological value. 

On the other hand, even if we maintain constancy in off- 
spring as the standard for systematic classification, almost the 
same number of systematic units are possible as when the reaction 
is taken as standard; however the units are possibly still more 
seldom realized. Furthermore my investigations show, that in 
Viola the homozygotic types are far more frequent in nature 
than might be expected, so that the difference between the two 
apprehensions is more of theoretical than of practical significance. 
In all cases we get a very large number of systematic units within 
V. arvensis and V. tricolor. 

It is interesting to see how the systematists hitherto have 
dealt with a polymorphous species like Viola. Wirrrock (1896) 
has given about 40 sub-species, varieties, sub-varieties, forms 
and sub-forms. The diagnoses comprise nearly all the varying 
characters, and in most cases the varieties are based upon one 
or a few individuals. WiITrTROcK's varieties and forms, in most 
cases correspond with. one of my 5,308,416 combinations, and 
it would be very fortunate if the same combination of all the 
characters was found once more, that is to say: there is the greatest 
possibility of finding one of the other 5,308,376 combinations that 
have no name. Such a division, onthe one hand, is quite insufficient, 
but on the other we cannot reasonably divide all the earth's 
140,000 fold” species into so many new species, each with a new 
name. 

Moreover I have been able to recognize several of WIrrrRocK's 


— 394 — 


types as segregation products from tricolor X arvensis. Such 
segregation products are the following types (Wirrrock 1897): 


V. arvensis Murr. ”communis Wittr. var. gotlandica Wittr. (table 

X, fig. 144—152). 

=curtisepala Wittr. (table XII, fig. 196—203; 
table XIII, fig. 220—224). 

=sublilacina Wittr. (table XII, fig. 182—195); 

”"sublilacina var. atropurpurascens Wittr. (table 
XIII, fig. 216—218); 

”striolata Wittr. (table XII, fig. 204—205). 


When WiIrrTrROckK supposed that he was able to recognize the 
parental varieties to some other spontaneous hybrids, he mentions, 
it is quite illusory. 

This investigation shows how little reason there is in the 
modern splitting up of Linnean species. Had one of these system- 
atists found my individual from Læsø with the curious emargin- 
ate, nearly laciniate lateral petals, its erect stature and narrow, 
deeply serrate (fig. 4 6, pag. 369), glaucous and fleshy leaves, 
I am sure he would have made a new species of it, especially as 
all the descendents were of the same type. This individual was 
made the original specimen, and all its characters were inserted 
in the diagnosis, both the palmate stipules, the long and 
incurved spur, the acuminate basal petal and the dark spot 
in front of the style. Now I caused the statistical investigation 
on the locality to be carried out (table II Z, Æ, Ø, pag. 381) and 
it showed that the character emarginata might be combined with 
different characters. Of the 16 combinations possible in table II, 
15 were realized among 100 individuals, this being the total 
number investigated from these 3 populations. If a new species 
should be based upon each diverging character such a small locality 
as Læsø would increase the number of species very much, because 
the individuals cross, and the new character soon will be combined 
with a lot of all the: earlier known characters. 

Every one, who is engaged in plant-determination, knows 
the difficulties caused by the fact that most specimens only agree 
with the diagnosis in some of the characters. In critical plant 
groups as Viola, where hybrids and stabilized segregation products 
from crosses are frequent, not only within the species but between 
the different collective species too, it is in many cases quite im- 


— 395 — 


possible to place the individual, which is examined, under any 
of the described systematic units. The result of these difficulties 
is a multitude of new "species.” The genus Hieracium is another 
instance hereof. The increase of "species" in this genus has been 
especially great in the last 15 years. The number of Hieracium- 
names embodidied in the main volume of Index Kewensis is 
about 1400. The increase has come as follows: 


In supplementum I, anno 1896: c. 900 species 


zz HR RO ERR 
ze EN ESSE fo HK SAG 40 ER 
e: ET BARE VI] IE, 
AN SENSE NE rer eee OU 


The number of ”species” up to the year of 1915 was about 
5000, but it increases rapidly from day to day. Most of the new 
species” are Scandinavian. The diagnoses are very long and 
comprise a 40—50 characters each. And I am sure there are 
millions of ”"species” not yet described. In Hieracium each little 
character is fixed by apomixis, but there is no doubt (according 
to ROSENBERG'Ss and OSTENFELD's investigations) that the different 
types and the large variety is due to preceding crosses between 
older species. Apparently circumstances here are to some extent 
parallel to Viola. 

It is quite clear that this course leads to chaos. 
On the other hand we cannot be content with the old 
division in species. In many accurate scientific investigations, 
for instance in the relation between the plants and the habitat, 
it is necessary to work with far smaller and more accurate limited 
systematic units than those of the old ”Linnean” species. But 
for these investigations, where sometimes one and sometimes 
another combination of characters is needed, it is necessary to 
work with the single, genotypically conditioned characters 
and their combinations (i. e. the isoreagents), which can be placed 
in a schedule as the tables I and II (pag. 378 and 381). Millions 
of new names of species are not needed for that reason. 

Even if we cannot state the reason for it, we feel sure that 
there is a reality behind the notion of the Linnean species, in the 
same manner as the genera are a reality. The Linnean species 
become a superior classification above the new "species” (varieties, 
microspecies and isoreagents) just as Genus is a superior class- 


— 396 — 


ification above Linnean Species. Therefore it is a confusion 
of ideas to co-ordinate the Species and the Microspecies by using 
the binar nomenclature for both. For many purposes we have 
no use for a finer classification than the Linnean species. 


In order to prevent this confusion I advance the following 


Propositrorns: 


The notion Species shall be maintained in its Linnean 
significance but according to our present knowledge as a 
superior classification. 

The Species in this significance, especially the critical ones, 
shall be critically revised with regard to the variation of the 
characters. New diagnoses shall be given; in these are embo- 
died only the characters that distinguish the species from 
all other species of that genus. No varying characters are 
embodied in the diagnoses. In a list after the diagnoses, all 
the varying characters and the genotypically conditioned 
variations of these (similar to my list of Viola) are given. 
The minor units shall, in order to distinguish them from 
the Linnean species, be named Microspecies (OSTENFELD 
1921, p.117) in the sense heritably fixed units, and Isore- 
agents (= combinations of genotypically conditioned cha- 
racters), when only the reaction of the individuals 
themselves against the conditions is taken into con- 
sideration (RAUNKIÆR 1918). 

The term Forma shall be applied as a designation of the 

variations caused by the external conditions solely 
(as for instance Polygonum amphibium L. f. terrestre and f. 
natans). 
Ternar nomenclature shall be applied for the Microspecies 
and Isoreagents. Just as the name of the species involves 
the name of the genus too, the name of the microspecies 
shall involve both the name of the genus and that of the 
species. (Not Hieractum marginelliceps Dahlst. but Hieracium 
silvaticum marginelliceps Dahblst. and not Viola maritima 
Schweigg. but Viola tricolor maritima Schweigg.). 


When the varying characters and the genotypically conditioned 


variations of these are given after the diagnosis, every one can 


— 397 — 


construct a system of Isoreagents from this list according to the 
special investigations for which he will use them. 

The splitting up of Linnean species in new »species« has fre- 
quently been based upon accidental observations of deviating in- 
dividuals which are described, instead of trying to make a per- 
spicuous classification of the types within the species based upon 
studies in the extent and the cause of the variation and other 
questions in connection herewith. In many cases this result is due 
to the fact that these systematists either do not accept the conse- 
quences of, or are not fully familiar with all the results of modern 
biology. The systematist's investigations are so difficult, that it 
is needful that he uses all the resources at his disposal. 
At the investigation of the small systematic units within the single 
Linnean species he cannot be content with employing the old me- 
thods alone, which suited the classification of the Linnean species 
themselves so long ago. 


VI. Must Viola tricolor and arvensis be regarded as two 
distinct Species? 


During the last 200 years these two systematic units alter- 
nately have been regarded as two distinct species and as two minor 
units belonging to the same species. 

As early as in 1745 A1B. HALLER distinguished two species 

(HALLER 1745). In Historia Plantarum Helvetiae (1769) 
"he distinguishes the two species in the following way: 

n. 568 (V. tricolor Riv.): "flore calyce duplo longiore.” 

n. 569 (V. bicolor Riv.): "flore calyce paulo majori.” 

LINNÉ regarded them as one species (Species Plantarum 
1753). 

MurrAaY (Prodr. Stirp. Gottingensium 1770) separates 
them under the names V. tricolor L. and V. arvensis Murr. (syn: 
V. bicolor arvensis C. BauH.) and for description he refers to 
HALLER'S forenamed description of them both. — Most frequently 
they have been looked upon as two species. 

I regard them as forming two species. If we assume the 
criterion of hybridization as the only criterion in all cases, they 
would not be so. They cross and produce fertile offspring. And 
moreover, there are smooth transitions between them. But then 
all the Melanium section would constitute one species. In Viola 


— 398 — 


the facts are so extraordinary, that I do not believe that the com- 
mon rules are sufficient. 
For following reasons I maintain them to be two species: 
1. The chromosome numbers are very distinct: 13 and 17. 
Two well defined types: grand, lab, viol and parv, n.lab, 
albid exist and are far more frequent than the transition 
types. 
3. The two species, when crossed, cannot give segregation in 
Mendelian proportions owing to the difference in chromosome 
number and the irregular distribution of chromosomes in F,. 


DO 


VII. What is typical V. tricolor and typical V. arvensis? 


I know no better distinction between the two species than 
HALLER's above quoted diagnoses: grandiflorda — tricolor and 
parviflora = arvensis. But the dividing line between grandiflora 
and parviflora is not quite the same as HALLER's. According to 
general use I determinate parviflora (arvensis in widest sense) 
as all plants with the upper petals as long as or shorter 
than the upper sepal (including both the homozygotic parvi- 
flora and the intermediate heterozygotic parviflora) and grandi- 
flora (tricolor in widest sense) as all plants with the upper 
petals longer than the upper sepal. 

But when the question is what is typical arvensis and tricolor 
i. e. the probable combination of the original species, 
that crossed have produced the great multitude of types, we 
have no other guide than the frequency of the combinations in 
nature. 

Looking at table I, pag. 378 we will find, when only the three 
first characters are considered, that one combination of parviflora 
is far more frequent than the three other combinations. This 
combination is parv, albid, pinn and must be considered as the 
typical arvensis. The other combinations are of unfrequent occur- 
rence: parv, albid, palm and especially the two combinations parv, 
viol (n.albid) (only 46 individuals among 758). 

Among the grandiflora individuals two of the four combin- 
ations are far more frequent than the remaining two. The com- 
bination grand, albid is very rare (only 12 individuals among 
1142). The remaining 1130 individuals are all grand, viol (n. albid) 
but 449 have pinnate and 681 have palmate stipules. It is 


— 399 — 


a very wide spread opinion, that tricolor has pinnate stipules 
(BEckErR 1905, pag. 48, about tricolor: "”Stipulae pinnatifidae”), 
but I presume it is wrong. Palmate stipules are more frequent 
than pinnate. Many of the tricolor (grandiflora) individuals descend 
from crosses between tricolor and arvensis and I suppose that all 
the tricolor individuals with pinnate stipules are stabilized segre- 
gation products from these crosses. In other words: that the 
original typical tricolors had palmate and the original typical 
arvensis had pinnate stipules. If we had a territory with tricolor 
alone, the statistic proportion from this territory might decide 
the question. Such a territory we do not have, as arvensis occupies 
a territory being co-extensive with and extending beyond trico- 
lor's. When the facts are so, we must try to compare a territory 
where tricolor is far more common than arvensis with one where 
arvensis is just as frequent as tricolor. In Jutland we have such 
two territories. V. tricolor (grand, viol) is one of the most predom- 
inant plants in West-Jutland. In early summer the fields are 
blue with these flowers. In many of these fields I never saw an 
arvensis. In East-Jutland fields with arvensis alone alternate 
with fields with tricolor alone and with fields with both arvensis 
and tricolor. The following table gives the distribution of palmate 
and pinnate individuals among the tzricolors in 7 habitats from 
West-Jutland and 7 habitats from East-Jutland: 








— FE 3 
Number of | 0 io-tic 
individuals |" relation 




















West | pinn 178 20910 
Jutland | palm | 432 71 9/0 
grand viol == RER EØF IE" 
East pmn | PA IP 47 9/9 
Jutland palm 333 53 9/0 


The table shows that in West-Jutland a far greater proportion 
of tricolors have palmate stipules than in East-Jutland and there- 
fore I suppose palmate stipules to be typical of the pure tricolor. 

As to the dark spot in front of the style it is rare in parvi- 
flora individuals and very frequent in grandiflora individuals and 
I suppose it to be a tricolor-character. 


— 400 — 


The characters that distinguish the typical arvensis from 
the typical tricolor are the following (the first named characters 
are the most significant): 


Viola arvensis typica: parviflora stigma non labellatum, petala 
albida, stipulae pinnatifidae, stylus non maculatus. 

Viola tricolor typica: grandiflora, stigma non labellatum, petala 
violacea (lutea, rosea, alba), stipulae palmatisectae, stylus 
maculatus. 

Viola tricolor maritima (typica): Folia parva, carnosula, 
caules atropurpurei, caespitosi, prostrati. 


VIII. Notes on the Geographical Distribution of V. arvensis 
and V. tricolor in Denmark. 


On a journey through Jutland in early summer 1921, under- 
taken with the support from the Botanisk Rejsefond in order 
to examine the variation of the Danish Viola-species belonging 
to the Melanium section, I had occasion to make some observ- 
ations concerning the distribution of the types. 

One of my routes stretched from Aabenraa in Eastern South- 
Jutland to Flensborg Fjord. From here it ran across South-Jut- 
land to Tønder and Højer on the west coast. Further northwards 
along the coast to Hjerpsted and from here eastwards to Visby. 

Another route stretched from Varde westwards through the 
heathery countries to the North Sea and northwards through 
the dunes along the coast and through the heathery countries 
to Nymindegab at the southern end of Ringkøbing Fjord. From 
Ringkøbing westwards through the fertile Holmsland between 
Ringkøbing Fjord and Stadil Fjord and southwards through the 
sandy isthmus Holmsland Klit between Ringkøbing Fjord and 
the North Sea. Further northwards along the dunes almost to 
Nissum Fjord and eastwards to Ulfborg. 

AÅA third route stretched from Viborg through the reclaimed 
heathery territories in Mid-Jutland north eastwards to Hobro at 
the east coast and northwards to East-Himmerland, which is 
very abundant in lime. The lime in most places is very near 
to the surface of the ground, and at several places it juts out 
in lime-hills. Southwards again through East-Jutland via Had- 
sund to Randers and south-east to Kalø and Mols with the sandy 


— 401 — 


Mols Hills at Kalø Vig. Finally I visited the two islands Tunø 
and Samsø in Kattegat. 

Along the first route in South-Jutland V. tricolor was the 
predominant of the two species. Only in the eastern part some 
arvensis were found. These tricolors were the most palmate ones 
I ever have found (population V, table I, pag. 379). Some of these 
(from Røde-Kro) had palmatilobate, broad-based, triangular 
stipules resembling cornuta's and similar to the type BECKER 
has described from the Færøes as V. tricolor subsp. færøensis 
W. Becker in Bot. Færøes III 1907, pag. 856. No maritima 
types were found at the West coast by Emmerlev or Hjerpsted. 

Along the second route through the heaths and dun: s of 
West Jutland (populations P, Y, F table I, pag. 378, AA and BB 
table II pag. 381), V. tricolor was the all-predominant species. In 
my statistical schedules I have no V. arvensis. My intention was 
to study the maritima types here. South of Nyminde I found 
only one considerable population of V. tricolor in the dunes. It 
was at Børsmose mid-way between Blaavands Huk and Nyminde. 
They were tricolor individuals of the common type, only a little 
phenotypically altered in appearance by the extreme conditions. 
The leaves were not fleshy and the stems were normally green, 
non atropurpurea. No real maritima (parvifoltata, carnosula, atro- 
purpurea) were found till Nyminde, where the population AA, 
table II, pag. 381, is from. All the individuals were parvifol, carno- 
sula. Only 10 per cent were nm.atropurp, the remaining were 
atropurp. From here to Øhuse south of Nissum Fjord (population 
BB, table II), where I left the dunes, maritima was very frequent 
and the only one living in the dunes, even if the typical tricolor 
grew in the fields on the other side of the road that separated the 
dunes from the fields. Most of these maritima were a little acu- 
minata but this character was not so strongly marked as in those 
I found in 1919 in the dunes of Skagen (CLausenN 1921, pag. 
210, population K). The point on the spur-bearing petal was 
very short. 

The typical maritima lives also further south than Nyminde. 
From Fanø I have received seeds of a true parvifol, carnosula, 
atropurp, prostrata type (leg. Professor WINGE). 

In spite of the varying characters the maritima types from 
Fanø, Nyminde-Nissum, Skagen and Læsø have something in 
common: the small, narrow, glabrous, fleshy leaves, the narrow 

Botanisk Tidsskrift. 37. Bind. 26 


— 402 — 


sects of the stipules, the atropurpureous stems (often the nerves 
of the leaves are atropurpureous too), the narrow petals, the short 
appendices of sepals and the long spur. Usually they are cespitosely 
branched in the surface of the ground. As far as I have been 
able to examine the maritima types in exsiccates from other 
coasts both in Denmark, Sweden and Germany (The Baltic 
Sea) and from the Dutch, Belgian, French and British shores of 
the North Sea and the Channel they are rather different from 
the West-Jutland type which is the most extreme of them all and 
likely the fittest type to the extreme conditions. It is, in fact, 
the West-Jutland type that LANGE describes and pictures in 
Flora Danica fasc. 45, pag. 4, plate 2647 under the name of 
V. tricolor (L.) var. arenaria (Sond.), but it is surely not this 
variety, which Wirrrock (1897, pag. 69) correctly remarks. The 
specimen pictured in Flora Danica is acuminata. — Each ter- 
ritory has its special composition, and apparently the Skagen 
type with its always prostrate stems with very long internodes 
and its very narrow, long and tapering petals, that gives it an 
appearance very strange in a Viola, is the most extreme of them all. 

The route from Viborg to Hobro was characterized by a 
multitude of types and combinations (populations I, M, N, A, 
table I, pag. 378). It was one large genetic experiment arranged 
by nature. It was characteristic that in the first field east of 
Viborg, where I found Viola tricolor, I immediately saw that the 
type was somewhat different from the type in the West-Jutland 
fields. Soon I discovered a plant, which I recognized as a hybrid 
between tricolor and arvensis. Necessarily I then might expect 
to find arvensis, and really, I soon discovered several typical 
arvensis. In the same manner facts were from field to field. In 
East-Jutland, especially in East-Himmerland between Mariager 
Fjord and Limfjord, arvensis is the most common of the two 
Danish Melantum species. In many parts and many fields not 
one tricolor can be discovered, while arvensis is very common 
(populations C, D, E, table I). But in many places here, areas with 
arvensis alone (populations B, G, table I) alternate with areas with 
tricolor alone (populations L, R, table I) and with areas with them 
both and the intermediate type (populations H, J, K, O, table I). 
Apparently the two species meet in the middle of Jut- 
land and cross together. 

Upon Samsø at Selvig I found a dune population (X, table 


ROSE 


I); but it was not the typical West-Jutland maritima. The indivi- 
duals were all grandifol, n.atropurp. The only maritima character, 
they were in possession of, was caespitosa. Finally the whole 
population had pallida petals. Both tricolor and arvensis lives 
on Samsø (populations 5, U, table I). 


IX. The Distribution after the Conditions of the Habitat. 


One of the most interesting facts concerning the distribution 
of the isoreagents is that the isoreagents belonging to V. tricolor 
maritima (parvifol., carnosula) have never been found in other 
places than the dunes. 

Another interesting fact is the distribution of the isoreagents 
according to the degree of acidity in the soil. Previously FERDI- 
NANDSEN (1918) has stated, that V. tricolor is acidophilous and he 
suggests its possible occurrence on ground that lacks lime to 
86 per cent. The corresponding index for V. arvensis he states to be 
40 per cent.,so that this species should be faintly acidophobous. 
FERDINANDSEN has recorded tricolor or arvensis from 31 of his 
statistic tables (31 habitats). He classifies the degree of acidity 
in the soil in the following classes: basic, faintly basic, neutral, 
neutral-faintly acid, and acid. In the basic classes only arvensis 
is found, in the classes neutral and faintly acid both arvensis 
and tricolor and in the acid class only tricolor. The following table 
shows the distribution of the habitats among the classes of acidity: 








| Reaction of soil 





Classes of acidity 


faintly neutral- 











basic basic neutral faintly acid aCid 
| | | 
Number of | arvensis | 7 3 8 5 | 
habitats with : | | == Ve SPORER 
TS th | tricolor | | 1 4 4 
ll | | 


| 


Primarily it was not my intention to examine this question. 
Therefore I was not prepared for a more detailed investigation. 
But, during my journey, I observed that the distribution was 
so characteristic that even a rather rough determination of the 
degree of acidity might suffice to give a result. I applied litmus 
paper, hence only a rough classification in basic, neutral and 
acid was obtained. But this classification agrees with the general 


26% 


— 404 — 


impression of the habitat, and it also coincides with the results 
that might be expected from the geological structure of the ground 
on the habitats. 

In table I, pag. 378, the habitats are arranged according to 
the degree of acidity in the soil: basic, neutral, acid; 3 habitats 
that only gave a very faintly acid reaction are inserted in a special 
class. 

When the habitats F, T and U are excepted, the isoreagents 
are distributed as follows: 


On basic ground: parv, albid (pure arvensis), populations A—E. 

On acid ground: grand, viol (pure tricolor), populations P—S. 

On neutral-faintly acid ground: all the combinations: parv, 
albid; parv, viol; grand, albid and grand, viol (pure arvensis 
and pure tricolor and the intermediate combinations), popu- 
lations G—O. 


This result is in accordance with FERDINANDSEN'S above 
mentioned. 

The cause of the many types present on neutral—faintly acid 
soil is certainly that both the extreme combinations (arvensis 
and tricolor) meet on this soil, and therefore there is a possibility 
of crosses. 

Even if the reaction of the soil is not the only distributing 
factor, it is surely one of the most important. The geographical 
distribution of arvensis and tricolor in Jutland is certainly rather 
due to the reaction of the soil in East- and West-Jutland than 
to any special geographical causes. This is evident from the fact 
that the composition of the populations can change within very re- 
stricted areas parallel to the changes in the reaction of the soil. 
A diagram of the route from Viborg to Hobro will show it: 


faintly a faintly et faintly 
| ner | [neutral] | SÆr | [basic] | BR 
M ——> I ————7 N ——————— 3 Å 3 Å; (not in table I). 


parv, albid — grand, viol parv, albid all combi- 
and intermediates alone nations 


At Onsild (A) the populations suddenly changed. All the 
large blue flowers disappeared coinciding with the fact that seed 
and grass became markedly more vigorous. The explanation of 


— 405 — 


this sudden change was a marle stratum. The diameter of this 
basic oasis was not more than about 3 km, and coinciding with 
the fact that the soil had become acid again the large blue flowers 
reappeared (A+). 

Another striking change was observed in Himmerland, as 
the following diagram will show: 




















[basic] [neutral] [faintly acid] 
ar am, 
C—— > E ——> D ——> H ——> J ——>0 
[parv, HØ alone] [all combinations] 


The distance between D (Bælum) and H (near Vorgaard) 
was not more than about 1 km, but the populations C—E—D were 
from the high lime districts and the populations H—J—O from 
the sandy valley of one of the rivers from the glacial period. I 
was not aware of this boundary line before Viola itself reminded 
me of it when I saw the large, blue flowers again. 

I do not believe that the cause of this distribution is, that 
tricolor is incapable of growing in basic soil and arvensis incap- 
able of growing in acid soil. But plants that react upon the 
degree of acidity have an optimum at which they become most 
vigorous (CARSTEN OLSEN 1921), at other concentrations of pp. 
they become more weak. Apparently arvensis has its optimum 
in the basic or neutral soil and tricolor in the acid soil. The strong 
competition in nature then causes that they usually only can 
pull through when the reaction of the soil approaches the opti- 
mum for the species in question. 

There are 3 exceptions in table I (pag. 378). The population 
F with grand, viol on basic soil and T and U with parv, albid 
on acid soil. When the number of exceptions is so relatively large, 
it has its natural explanation in the fact that I have selected 
those populations which I suspected would deviate from the 
rule. The population F is from Holmsland (not Holmsland 
Klit) between Ringkøbing Fjord and Stadil Fjord. It is a fertile 
basic oasis with loamy soil in the West-Jutland acid and sandy 
heath land. There is marle on Holmsland. Seed and grass was 
very vigorous, likewise Viola. I did not succeed in finding one 
arvensis here. 

The population T is from a stony rye field near the harbour 








— 406 — 


of Tunø. Only parv, albid was found here. These plants were 
particularly vigorous; but the rye was weak and the soil distinctly 
acid. I have never seen Viola with a larger assimilating surface 
than these individuals. This fact was due to both the large leaves 
and the extraordinarily large stipules. Leaf and stipules of one 
of these plants is pictured in fig. 1, pag. 365. I presume that 
these arvensis descended from stabilized segregation products of 
crosses between tricolor and arvensis. V. tricolor lives on the island 
too. The chromosome number was the usual in arvensis, namely 
17, and the divisions were regular (fig. 6 a, pag. 385). 

Population U is from a sandy grass field at Koldby Kaas 
on the island Samsø. The soil was distinctly acid and the grass 
sparse. Only parv, albid was found and but few. In contradis- 
tinction from the specimens from population T they belonged to 
a small dwarf type. A characteristic feature was, that 13 of the 
14 plants had palmatisect stipules as tricolor. Fig. 4 a, pag. 369 
shows leaf, stipules and flower of offspring of one of these individu- 
als, cultivated in the Botanical Garden this year. All the offspring 
was dwarf and palmatisect. The chromosome number is not 
determined yet. I am quite sure that this arvensis type is a segre- 
gation product from a cross between arvensis and tricolor. 

The "sentiment”, if we may say so, of an individual against 
the degree of acidity in the soil is a physiological character pos- 
sibly contingent on one or more factors. Usually these factors 
are coupled with the factors that stipulate the size and the colour 
of the petals. The two species have a different optimum as to 
the degree of acidity, arvensis at a high pr -value and tricolor 
at a lower one. It is not likely that these factors for a certain 
optimum should be coupled absolutely with the factors for the 
morphological characters. When the two species cross, we there- 
fore should expect to get arvensis types (parv, albid) with optimum 
at a low px. (acidophil) and tricolor types (grand, viol) with opti- 
mum at a high p4. (acidophob.) In some cases the preceding re- 
combination of factors is manifested by the fact, that a morpho- 
logical character has been separated from the complex of cha- 
racters it usually is connected with. Such is apparently the case 
with the population U, where the character palmata together 
with the character acidophilous have possibly become separated from 
the character-complex grand, viol and instead have been connected 
with the character-complex parv, albid. 


— 407 — 


In other cases, as in the population F from Holmsland, no ex- 
ternal signs of a preceding cross and recombination have been found, 
but there can be no objection to that supposition that a factor 
for optimum at a high py.-value might cross over to the factor- 
complex for grand, viol without being accompanied by factors for 
morphological characters, so that the explanation, also in this 
case, can be a preceding cross between tricolor and arvensis 
with the following recombination of factors. 


X. Adaptation as a Result of Mutation, Crosses and Selection. 


In a short paper Turesson (1922) has shown that some 
so-called "adaptative” types inhabit certain localities, so that 
the re-appearance of a distinct locality coincides with the re- 
appearance of the variety typical to that locality. This fact is 
most strikingly shown in Hreracium umbellatum. In the south- 
eastern corner of Skåne (Sweden) woods, dunes and sand-fields 
alternate within short distances. The woods are inhabited by 
an erect broad-leaved type, the dunes by a lesser erect, more 
narrow-leaved and strongly shoot-regenerating type; in the sand- 
fields grows a prostrate, hirsute type. Where woodland is replaced 
by dunes, the woodland varieties are replaced by the dune vari- 
eties and these in their turn are replaced by the prostrate types 
when the sand-fields begin. At the boundary line between two of these 
zones a narrow zone inhabited by all kinds of intermediate types 
can be found. — The same applies to three Atriplex-types. The east 
coast of South-Sweden is inhabited by one type, the west coast 
along the sound by another and north of the sound on the ex- 
posed west coast lives a more deviating, small-leaved, prostrate 
type. On the whole TureEesson found a parallelism between the 
extremeness of type and the extremeness of locality. He sup- 
poses the varieties differentiated from previously existing types 
and their combinations when crossed. 

The parallelism between these Hieracium- and Atriplex-types 
and the extreme West-Jutland types of Vzola is striking. Certain 
populations are composed by isoreagents which are ecologically 
adapted” to the conditions of the habitat. In the vagabondizing 
Viola populations on cultivated ground such an adaptation is 
of course not so pronounced. These populations do not live so 
long at the same place that they can become balanced. But 


— 408 — 


in the dune populations an accordance to the extreme conditions 
can be recognized: in the small transpiration surfaces of the leaves, 
m the somewhat fleshy leaves, in the large and deep going roots, 
in the plentiful anthocyanin in stems and leaves, in the extensive 
ramification in the earth's crust, by which they become perennial, 
and in the transversely geotropic stems, they exhibit genotypically 
conditioned characters that are of great importance to the plants 
at these localities. 

The older Lamarckian interpretation regarded these "adapt- 
ations” as a direct effect of the extreme conditions. The Selec- 
tionism saw an effect of the continued selection between the 
fluctuating variations. That there is a fitness to function can- 
not be denied. The question is merely in which way it has come 
into existence. 

The selection cannot create new types (JOHANNSEN) but it 
can select the fittest among already existing types or among 
types that arise after crosses, in other words: it can select 
the fittest combinations of the existing genes, but it 
cannot create new genes. 

As to the supposed direct effect of the conditions, we neither 
have been able to trace such an effect nor disprove it. It agrees 
badly with our present apprehensions concerning genotypic struc- 
tures and genotypic constancy. The only instance we know of 
genotypic inconstancy is the mutations. In which way they 
take place and by which influences they are induced we do not 
know. And as to the nature of mutations we know nothing. The 
only way in which we might suppose that the conditions could 
act upon the genotypic structure is through the mutations, either 
by increasing the number of mutations or by affecting the direction 
of the mutations. 

But we need not a supposition of a direct effect of the external 
conditions to explain the parallelism between the extremeness 
of type and the extremeness of locality. This might be explained 
as an effect of mutation, crosses and selection solely. The muta- 
tions, crosses and segregations mainly provide the material for 
the selection, and the selection itself brings about the "adapta- 
tion”. Qccasionally, crosses between different species occur 
and supply one species with a character from another that enables 
it to live in places, where it previously could not live. In Viola : 
tricolor and arvensis this is unusually frequent (tricolor on basic 
soil and arvensis on acid soil). 


— 409 — 


When the existing and not yet existing but possible hereditary 
different types are as many as in V. arvensis—tricolor, the selection 
has a large material to act upon, and by and hy, eliminates the 
least fit types. 

The West-Jutland dune types are in the character parvi- 
foliata recessive to the common grandifoliata types. Likely the 
parvifoliata originally came into existence by a mutation in a 
place far from the dunes. This character is not profitable in places, 
where the transpiration is normal. But as the factor for it is 
recessive, it can ”"emigrate concealed” (RAauNnKkIæÆrR 1920, pag. 4) 
heterozygotic, even if all the homozygotic parvifoliate individuals 
die before they can disperse seeds. When it reaches to a locality, 
where the character induced by this factor is profitable, it becomes 
homozygotic by elimination of both the heterozygotic and homo- 
zygotic grandifoliate individuals. If A is the factor for grandifol 
and a the factor for parvifol the circumstances might be illustrated 
by the following diagram: 




















grandifol grandifol parvifol 

4 EH | £E 

FODEE (aa dies) (Aa and AA dies) 
grass field dune 


aa cannot live in the grass field- owing to the strong com- 
petition there. Aa and AA cannot live in the dunes because of 
the strong transpiration there. Therefore these two isoreagents 
do not intrude on each others territories. Even if they cross at 
the boundaries, the type not fit to the conditions is eliminated. 
Only in this way the aforementioned peculiar circumstances at 
Holmsland Klit (pag. 375, fig. da and b) can be explained. 

Stabilization of conditions in a place will certainly, by and 
by, cause the isoreagents fit to the conditions at the spot in ques- 
tion, to become more and more homozygotic; those with the 
recessive characters first, but also those with the favourable 
dominant characters, as all the recessive here have gradually 
been eliminated when they are segregated. The heterozygotes 
themselves decrease in proportion to the dominant homozygotes 
unless the first should be better fit to live than the latter. Accord- 


— 410 — 


ing to this it will be understood why so many types from nature 
are pure in the essential characters and show no segregation. 
Probably the homozygotes are far more common in nature than 
originally supposed. 





The principal cultivation experiments were carried out in 
the Botanical Garden of the University, Copenhagen. 
The cytological investigations were carried out in the Laboratory 
of Genetics of the Royal Veterinary and Agricultural 
College, Copenhagen, and some of the cultivation experiments 
from 1922 here mentioned took place at its experimental field 
at Lyngby. I tender my best thanks to Professor RAUNKIÆR, 
Director of the Botanical Garden, and to Professor WINGE Phil. 
Dr., Director of the Laboratory of Genetics, for the kindness 
with which they have placed these institutions at my disposal. 

I thank the "Botanisk Rejsefond” for the kindly aid 
given me for my statistical and ecological investigations in nature, 
especially in Jutland, and I thank Mr. C. A. JørGEnNSEN for his 
kindness in making the three lists from Læsø. 


Copenhagen, the Botanical Museum of the University. "/s. 1922. 


Literature. 


Becker, W., (1904): Systematische Behandlung der Viola arvensis auf 
Grundlage unserer phylogenetischen Kenntnisse. (Mitt. des thuring. bot. 
Vereins. Neue Folge XIX. Heft 19). — Weimar. 

Clausen, J., (1921): Studies on the Collective Species Vzola tricolor L. 
(Preliminary Notes). (Bot. Tidsskr. Bd. 37). — København. 

Ferdinandsen, C., (1918): Undersøgelser over danske Ukrudtsforma- 
tioner paa Mineraljorder. — København. 

Flora Danica, Vol. XV. Fase. -45. 

Haller, Alb., (1745): Flora Jenensis. — Jena. 

— (1769): Nomenclator ex Historia Plantarum indigenarum Helvetiae. 
— Bern. 
Linné, C., (1758): Species Plantarum. — Stockholm. 
Murray, Å., (1770): Prodr. designationis Stirpium Gottingensium. — 


Goettingen. 
Nachtsheim, H., (1921): Die Grindung der Deutschen Gesellschaft fur 
Vererbungswissenschaft. (Die Naturwissenschaften. Heft 22). — Berlin. 


Olsen, C., (1921): Studier over Jordbundens Brintionkoncentration og 
dens Betydning for Vegetationen, særlig for Plantefordelingen. (Compt. 
rend. trav. Laborat. Carlsberg. Vol. 15). — København. 


— 411 — 


Ostenfeld, C. H., (1921): Some experiments on the origin of new forms 


im tbe genus Hrieracium. (Journal of Genetics. Vol. XV). — Cam- 
bridge. 
Raunkiær, C., (1889): Vesterhavets Øst- og Sydkysts Vegetation. — 


København. 

— (1918): Ueber den Begriff der Elementarart im Lichte der modernen 
Erblichkeitsforschung. (Zeitschr. f. ind. Abst. und Vererbungslehre. 
Bd. 19). — Berlin. 

— (1920): Erindringsord til Forelæsninger over Plantegeografi. — Køben- 
havn. 

Turesson, G., (1922): The Species and tne Variety as ecological units. 
(Hereditas III). — Lund. 

Wittrock, B. V., (1897):. Vzola-Studier I. (Acta Horti Bergiani, Bd. 2, 
Nr. 1). — Stockholm. 





Studier over Samlearten Viola tricolor L. II. 


Resumé 
af 


J. Clausen. 


I. I Fortsættelse af den foreløbige Meddelelse i nærværende Binds 
3die Hefte om de varierende Karakterer hos Viola tricolor — arvensis skal 
her gives en Meddelelse om en Del yderligere varierende Karakterer: 


6. Kronbladfarve: 
Hos typisk arvensis: gulhvid (albid). 
Hos typisk tricolor følgende Farver: 
blaa (vol), højgul (lut), rosa ell. rødlila (rosea), blegblaa 
(pall) og ren hvid (alba). 
7. Sporelængde: 
langsporet (long.calc): Sporelængden mere end 2 Gange Bæger- 
bladenes Vedhæng; hos Klit-Typer (Fig. 1e, S. 365). 
kortsporet (brev.calc): Sporelængden ikke mere end 2 Gange 
Bægerbladenes Vedhæng. 
8. Sporeform: 
krumsporet (7ncurv.calc): hyppigst hos langsporede. 
retsporet (rect.calc). 
9. Formen af det sporebærende Kronblad: 
tilspidset (acum): Fig. 2 S. 367; hos vestjydske og læsøske Klit- 


typer. 
ikke tilspidset (n.acum): Fig. 16, S. 365; den almindeligste 
Form. 
10. Sidekronblade: 
udrandede — svagt fligede (emarg): Fig. 2; findes kun paa 


Læsø; muligvis ingen andre Steder i Verden; 
hele (integ): den almindelige Type. 


Lg 


13. 


14. 


16. 


ze 


Ile) 


— 412 — 


Kronbladenes Overflade: 
fløjlsagtig (velut): er foraarsaget ved, at de Vorter, hvortil Kron- 
bladenes Overhudsceller er udtrukne, er særlig tætte og kraftige 
(Fig. 3 a, S. 368); denne Karakter kan ogsaa findes hos vildtvok- 
sende V. tricolor. . 
ikke fløjlsagtig (n.velut): det almindeligste. 


. BStøvkornene, der i fugtig Tilstand set ovenfra er enten: 


hovedsagelig firkantede (Fig. 3 d): tricolor, 

hovedsagelig femkantede (Fig. 3€): arvensis eller 

omtrent lige mange fir- og femkantede. 

Mellem de firkantede kan findes enkelte trekantede (Fig. 3 c) og mel- 
lem de femkantede enkelte sekskantede (Fig. 3f). Set fra Kanten 
er alle Støvkornene som Fig. 3 b. 
Bladstørrelse: . 
smaabladede (parvifolata): Fig. 1c, S. 365 og 46, S. 369. En 
Type fra Klitter; de smaabladede er vistnok altid tillige noget 
kødfulde; 

Blade af normal Type (grandifoliata): Fig. 1a og b og Fig. 406. 
Bladfladen er 3—4 Gange større end hos de smaabladede Klit- 
typer. Bladene er ikke kødfulde. 

Bladform: 

ægformede, rundtakkede (ovat-cren): (Fig. 14): hos arvensis; 

lancetformede, savtakkede (lanc-serr), (Fig. 16): hos tricolor. 


. Akselblade: 


fjerdelte (pinn): hos arvensis (Fig. 1 a); 
haanddelte (palm): hos tricolor (Fig. 1 6). 
Akselblades Midtflig: 
løvbladagtig udvidet (foliac): Fig. 1 d og e. (Midtfligen takket). 
ikke løvbladagtig udvidet (n.foltac): Fig. 4a, b og ec, S. 369. 
Vækstform: 
nedliggende Stængler (prostr): alle Stængler er transversalt 
geotropiske (Fig. 5 b og c, 8. 371). Findes mest ide vestjydske 
Klitter. 
oprette Stængler (erect): Hovedstænglen er negativt geotro- 
pisk (Fig. 5 a). Det almindeligste Tilfælde. 
Stængelfarve: 
anthocyanrøde Stængler (atrop): findes mest i Klitter; 
frisk grønne Stængler (n.atrop). 


levetid 


fleraarige, d. v. s. tueformet forgrenede i Jordskorpen (caesp): en 
Karakter, der findes hos de fleste vestjydske og læsøske Klit- 
typer (Fig. 56). Ofte 20—50 Sidegrene i Jordskorpen. 
enaarige, ikke tueformet forgrenede (n.caesp): kun Sidegrene 
ovenfor Jordskorpen. 
Flere af disse Karakterer griber ind over hinanden og er derfor under- 
tiden vanskelige at skelne fra hinanden. 
II. Disse forskellige her nævnte Variationer er arveligt forskellige. 


Et af de mest slaaende Eksempler herpaa afgiver Planterne paa Fig. 5, 8. 371. 


— 413 — 


De to Planter er begge fra S. Lyngvig paa Holmsland Klit. 5a er 
fra en Græsmark paa Tangens mere leragtige Inderside og 5 6 er fra Klit- 
ten paa den anden Side af Vejen, der skiller Mark og Klit fra hinanden. 
5 a var opret, storbladet, friskgrøn, enaarig, medens 5 b var nedliggende, 
smaabladet, anthocyanrød, flersårie re altsammen Egenskaber, der 
kan have Betydning for re der lever i Klitter. Alle Eelam paa 
Marken var af Type som ! ao (Bestånd Y, Tabel I, S. 379) og alle fra Klit- 
ten som 5 b (Bestand Fm Tabel II, S. 381). Og dog var det ikke Modi- 
fikationer fremkaldt af de meget forskellige ydre Kaar, men virkelig 
arvelige Forskelligheder betingede af indre Anlæg. Afkom af disse 2 
Planter blev nemlig dyrket i Bot. Have under samme Kaar, og des- 
uagtet bibeholdt det Forskellighederne, idet alt Afkommet fra Marken 
blev af Type som 5 a og alt fra Klitten af Type som 5 b. 

III. Saa langt som jeg har kunnet undersøge, har det vist sig, at 
alle Kombinationer af de omtalte Variationer er mulige, men 
de er langtfra lige hyppige allesammen. 

Tabel I, S. 378 giver Resultatet fra 23 Voksesteder A—Y hovedsage- 
lig fra Jylland. Der er taget Hensyn til 4 Karakterer, nemlig: 

Kronbladstørrelse: smaakronet (parv) og storkronet (grand). 

Kronbladfarve: gulhvid (albid) og blaa (vrol) (inclusive de andre 

tricolor-Farver). 

Akselblade: fjerdelte (pinn) og haanddelte (palm). 

Akselblades Midtflig: løvbladagtig (foldac) og ikke løvbladagtig 

(n.folzac). 

De paa et Voksested fundne Kombinationer af disse 4 Karakterer er 
beregnet pr. 100 Individer. 

Tabel II, $. 381 viser paa lignende Maade Kombinationerne mellem 
nogle Karakterer, der er typiske for Klitplanterne: Voksestederne Z—Ø 
er fra Læsø, AA og BB fra Klitter i Vestjylland. Alle Planterne er stor- 
kronede, men de varierer iøvrigt i: 

Akselblade: fjerdelte (pinn) og haanddelte (palm). 

Sporelængde: kortsporet (brev. .ealc) og langsporet (long.calc). 

Sporeform: retsporet (rect) og krumsporet (incurv). 

Sidekronblade: helrandede (integ) og udrandede (emarg). 

Forneden er der angivet den procentvise Mængde af Planter med 
tilspidset nedre RE abe (acum). For Bestandene AA og BB er 
kun angivet Akselblade, Sporelængde og Stængelfarve: frisk grøn (n.atrop) 
og anthocyanrød (atrop). 

Begge Tabellerne viser, at saa godt som alle Kombinationer er fundet 
realiseret. 

IV. De mange i Naturen forekommende Typer hørende til 
V. tricolor eller arvensis, stammer fra Krydsninger mellem tricolor 
og arvensis og mellem disse Krydsningers Udspaltningspro- 
dukter. 

De stærkt blandede Bestande i Naturen som i Tabel I: H, I, J, K, M, 
N og O er saadanne Udspaltninger, og Planter fra saadanne Bestande 
spalter yderligere. Tabel III, $. 382 viser de Typer, der fremkom af to 


— 414 — 


smaablomstrede, blaablomstrede Planter (parv, n.lab, viol, mac) V. 246 
og V. 247 fra Bestand F, Tabel III, Frederikshavn 1919. 

Ved kunstig Krydsning af V. arvensis med V. tricolor faas der i 2den 
Generation en Ukemnns. af Typer, der ganske ligner Typerne fra de 
omtalte, stærkt blandede Bestande fra Naturen. 

Endelig viser Cellekærnens Delingsforhold hos de formodede Bastarder 
i Naturen fuldstændig Overensstemmelse med Delingerne hos de kunstigt 
dannede Bastarder. V. tricolor (grand, viol) har 13 Kromosomer (Fig. 6 b, 
S. 385) og arvensis (parv, albid) har 17 (Fig. 6 a). Bastarderne derimod viser 
alle mulige Tal fra 13 til 17 i samme Støvsæk. Fig. 7 a—g og 8 a—e, S. 385, 
389 er Afbildninger af Delingerne i en frivillig Bastard fra Naturen (parv, lu 
fra V. 246, Tabel III, S. 382) og Fig. 7 h og 8 f er de tilsvarende Figurer 
af kunstige Bastarder. Som det ses, er der meget stor Overensstemmelse. 

V. De Smaa-Arter, som Wirrrock (1897) har beskrevet, er opstillet 
paa de foran omtalte varierende Karakterer. Da der med den Inddeling, 
jeg har anvendt, er 5,308,416 Kombinationer mulige, skal der et rent Held 
til at faa fat i en af de 40, som WITTROCK har beskrevet. Der er langt større 
Chance for at faa fat 1 en af de andre 5,308,376 Kombinationer. WITTROCK's 
Smaa-Årter kan genkendes som Bastardudspaltninger. 

Det store Antal Muligheder viser, at man ligesaa godt kan holde op 
med at opstille Smaa-Arter indenfor de ere Årter paa mm hidtil anvendte 
Maade, hvor alle Karakterer tages med i Diagnosen. Dersom man skal 
naa til Bunden paa den Maade, kommer man snart op paa et Antal af 
Smaa-Årter, der gør det umuligt for et Menneske indenfor sin egen Leve- 
tid at — ib et Individ af hver af de mulige Kombinationer inden- 
for en enkelt DER 

Til mere detaillerede økologiske Undersøgelser kan man ikke nøjes 
med den gamle linnéske Artsinddeling, men maa skelne mellem langt 
mindre systematiske Enheder. Til dette Brug kan man mest rationelt 
arbejde med selve Karaktererne (de anlægsbetingede) og deres Kom- 
binationer (Isoreagenterne) stillet op i et Kombinationsskema som Tabel 
I og II. Derved kan man nøjes med et langt mere begrænset Antal Navne. 
(De 5,308,416 omtalte Viola-Kombinationer kan udtrykkes ved at anvende 
45 forskellige Variationer af Karakterer. Som Regel har man Brug for 
et mindre Antal Karakterer). 

VI. Skønt V. tricolor og arvensis krydses og frembringer frugtbart 
Afkom, betragter jeg dem som to veldefinerede Arter. Der eksisterer 
nemlig 2 Typer: parv, n.lab, albid (arvensis) og grand, lab, viol (tricolor), 
der er langt hyppigere end Mellemtyperne (Tabel I); svarende dertil er 
2 Kromosomtal, 17 og 13, langt hyppigere end de andre mellemliggende 
Tal, der ogsaa synes at være forbundet med Uregelmæssigheder ved 
Delingerne. 

VII. Dersom V. tricolor — arvensis skal deles i 2 Grupper paa en 
saadan Maade, at ethvert Individ, der træffes, kan komme ind under enten 
den ene eller den anden Gruppe, bør man anvende Kronbladstørrel- 
sen som eneste Skelnemærke, saaledes at alle smaablomstrede (parv) hen- 
føres til arvensis og alle storblomstrede (grand) henføres til tricolor. Dette 
stemmer med MurRaAY's (1770) og HALLER's (1769) Diagnoser. 


— 415 — 


Dersom der derimod er Tale om de typiske Arter, kan man nok tage 
flere Karakterer med, og da følgende: 


Typisk V. arvensis: smaablomstret, Støvfang uden Læbe, Kron- 
blade gulhvide, Akselblade fjerfligede, Griffel uden Plet paa For- 
siden. 

Typisk V. tricolor: storblomstret, Støvfang med Læbe, Kronblade 
blaa (højgule, rosa ell. rent hvide), Akselblade haandsnitdelte, 
Griffel med mørk Plet paa Forsiden. 

Typisk V. tricolor maritima: som V. tricolor, men desuden: Blade 
smaa, noget kødfulde, Stænglerne anthocyanrøde, tueformet for- 
grenede, nedliggende. 


VIII. I Vestjylland (Bestandene F, P, Y, Tabel I) er V. tricolor langt 
den almindeligste af de 2 Arter, og i Klitbæltet er den saa godt som ude- 
lukkende af maritima-Typen (AA og BB, Tabel II). I Østjylland, spe- 
cielt i det kalkrige Øst-Himmerland, er arvensis den almindeligste Art 
(C, D, E, Tabel I), selv om ogsaa Marker med arvensis alene (B, G, Tabel I) 
veksler med Marker med tricolor alene (L, R, Tabel I) og med Marker 
med begge Arterne + Mellemtyperne (H, J, K, O, Tabel I). I Midtjyl- 
land mødes de to Arter og krydses med hinanden som Bestandene I, M 
og N, Tabel I, viser. 

IX. Fordelingen efter Voksestedets Kaar. Det er karakteri- 
stisk, at V. tricolor marittma (d. v. s. den fleraarige Type med smaa, kød- 
fulde Blade) udelukkende bebor Vestkystens og Læsøs udsatte Klit- 
bælter. Den Type, der vokser i Vestjylland, er forskellig fra de Typer, 
der vokser ved Østersøens Kyster og ved de engelske, franske, belgiske 
og nordtyske Kyster af Vesterhavet og Kanalen. Den yestjydske Type 
er den mest udprægede af dem alle ser dens smaa, smalle, glatte og noget 
kødfulde Blade, Akselbladenes smalle Afsnit, de anthocyanrøde Stængler, 
de smalle Kronblade, Bægerbladenes korte Vedhæng og den lange Spore. 
Naar dertil kommer den overordentlig stærke Forgrening i Jordskorpen, 
de ofte transversalt geotropiske Stængler med meget lange Stængelled og 
det i en lang Spids nedløbende sporebærende Kronblad hos nogle af Typerne, 
vil man forstaa, at de er meget afvigende fra andre Viola tricolor. At de 
fleste af disse Krake er gunstige paa disse udsatte Steder med den 
stærke Blæst og den stærke Fordampning er sikkert nok, og maritima- 
Typerne frembyder saaledes et meget smukt Eksempel paa den Parallelisme, 
der ofte findes mellem Særprægethed i Kaar og Særprægethed i Vegeta- 
tionstype. Disse ejendommelige Typer findes fra Nyminde til Skagen, 
paa Fanø og paa Nordsiden af Læsø. Den mest særprægede af dem alle 
er sikkert Skagen-Typen. 

Forøvrigt har Undersøgelsen vist, at Fordelingen af tricolor og arven- 
sis i Jylland snarere er helnee: af Jordens Surhedsgrad end af specielle 
geografiske Aarsager. De forskellige Typer fordeler sig paa følgende Maade: 

Paa alkalisk Bund: marv, albid (= ren arvensis): Bestandene 

A—E, Tabel I. 
Paa sur Bund: grand, viol (= ren tricolor); Bestandene P—S, 


Tabel I. 


— 4l6 — 


Paa neutral, svagt sur Bund: alle Kombinationerne: parv, albid; 
parv, viol; grand, albid og grand, viol (ten arvensis, ren tricolor og 
alle Mellemtyperne); Bestandene G—O, Tabel I. 

De tre Undtagelser, Bestandene F, T og U, har antagelig deres Aar- 
sag 1 en foregaaende Krydsning, hvorved Faktorerne for Evnen til at 
trives godt paa Surbund eller alkalisk Bund er blevet kombineret med 
Faktorerne for andre morfologiske Karakterer end de, som de plejer at. 
være kombinerede med. 

X. Den Overensstemmelse der findes mellem Særprægethed i Kaar 
og Særprægethed i Vegetationstype behøver ikke at være foraarsaget ved, 
at Kaarene kan ændre Typerne, ja det er sandsynligt, at Kaarene ingen 
Indflydelse har i saa Henseende. For at forklare Overensstemmelsen 
behøver man heller ikke at ty til denne Mulighed, idet den kan forklares 
ved Antagelsen om Mutationer, Krydsninger med paafølgende Udspalt- 
ninger og det .derpaa følgende naturlige Udvalg mellem de herved 
fremkomne Typer. Disse Virksomheder vil bevirke, at paa et Voksested 
med stabiliserede Kaar vil Typerne efterhaanden komme i Ligevægt, 
idet de mindst egnede udskydes, og de vil tillige bevirke, at Typerne paa 
saadanne Voksesteder vil blive arveligt konstante med Hensyn til de 
Karakterer, der har Betydning for dem i Konkurrencen paa disse Steder. 





Contents. ER 
1. The varying characters of Viola tricolor. and arvensis .........…. 363 
2. Åre the variations induced by genotypical differences? .......... 375 
3mrArerall"combinationskorttnercharactersspossible eee nen 377 
4. The cause of the Variation: 
aa) sAnalyticaleyrdenceesrr er Bror are Eee ERE 382 
Bb) Synthetical 'evidences ere erne Herren Ea ner aL EEN 384 
Gu Gytolorieal evidens Er En 384 
5. Are all the genotypically conditioned combinations to be regarded as 
spe dess nsNE Tnls myrer ERE re ES eo ER ERE ENE ERE 392 
6. Must V. tricolor and V. arvensis be regarded as two distinct species? 397 
(«What isstypical tricolor tand typicalsPSarvensis ERE ERE RE ER DSS 
8. Notes, on the geographical distribution of V. tricolor and Y. arvensis 
mi Denmarks se an EEN ER 400 
9. The distribution according to the conditions of the habitat...... 403 
10. Adaptation as a result of Mutation, Crosses and Selection ...... 407 
Dansk Resumé (Studier over Samlearten Viola tricolor L. II). ... - 411 








Bianco Lunos Bogtr. Kbhvn. 


Særtryk af Botanisk Tidsskrift. 37. Binds 5. Hefte. 1922, 


LIBRARY 
NEW YORK 
BOTANICAL 

GARGEN 


Heléococcum aurantacum n'sen' et nsper 
By 
C. A. Jørgensen. 


In the autumn of the year 1921 Prof. L. KornEerur ROoseEn- 
VINGE observed on a sample, which he had collected in the moor 
im the botanical gardens of the University of Copenhagen, some 
small globular perithecia of a fungus he did not know. 

As we did not succeed in finding it mentioned in the myco- 
logical text-books (Schroeter, Rabenhorst, Clements), the fungus 
was handed to me for close examination in order to determine 
it by means of the existing literature on this subject. 

In spite. of a careful examination of the species described 
within the section of the mycology (Plectascineæ-Perisporiales), 
where the present fungus according to its structure must be refer- 
red to, I have not succeeded in finding any suitable diagnosis of 
a genus with which it agrees. 

It seems therefore, that the fungus is not hitherto described, 
and even if the possibility is not excluded that a suitable diagnosis 
is to be found somewhere in the literature, I feel justified in 
describing the fungus as new to science, on the basis of the afore 
mentioned resultless investigations. 

Heleococcum aurantiacum nov.gen.etn.spec. Perithecia super- 
ficialia, globosa, 7/,—/, mm diam., primo albida, dein aurantiaca, 
peridio subtile, membranaceo, pseudoparenchymatico, ex 4—6 
stratis cellularibus formato, glabro, areolato, ostiolo nullo, sporis 
maceratione peridii liberatis; ascis globosis, sessilibus, numerosis, 
sine ordine collocatis, paraphysibus nullis; sporis hyalinis, uni- 
septatis, ellipsoideis, membrana crassiori, glabra, 253—30 4 long., 
10—15 w lat. (Fig. 1a, b, c & d). Habitat in solo humido horti 
botanici Hafniensis. 

Botanisk Tidsskrift. 37. Bind. 27 


— 418 — 





Eisæls 
a) Å ripe perithecium (X 110). 6) median section through the same (X 110). 
c) An ascus (X 650). d) Ascospores (X 650). 


Among the already described species T'estudina terrestris Biz- 
zozero approaches nearest to Heleococcum. They are alike in 
having globular perithecia with an areolated surface; the asci, 
which contain 8 two-locular spores, are in both forms globular 
and lie promiscuously within the perithecium and conidia lack 
in both species. 

Testudina differs from Heleococcum in its dark perithecia 
and spores but its special characteristic is, that the spores are 
liberated by the disintegration of the peridium into small polygon- 
ous fragments. It also differs as to the size of the spores and the 
spores of Testudina have a rugged surface. The most character- 
istic feature of the genus Heleococcum is the large, smooth, 
orange-coloured two-celled spores and the light colour of the peri- 
thecium; the spores are liberated by the disintegration of the 
peridium. The colour of the perithecium is localized in the wall of 
the spores, the peridium being quite hyaline. 


— 419 — 


It is a difficult task to decide in which family Heleococcum 
(and Testudina) are to be included. When following the elassifica- 
tion of Ed. Fischer im Engler & Prantl "Natiurliche Pflanzenfa- 
milien” I, 1, 1896, both forms have to be classed among the 
Plectascineæ, to which Ed. Fischer refers the families Gymno- 
ascaceæ, Aspergillaceæ, Onygenaceæ, Elaphomycetaceæ and Terfe- 
ziaceæ, the common feature of which is that the asci lie pro- 
miscuously in the ripe perithecium. 

Heleococcum cannot however be included in any of these 
families. It certainly approaches to the Aspergillaceæ in the 
structure of its perithecium, but differs in its two-celled spores 
and in the fact that conidia are entirely wanting. 

I prefer therefore, as Clements in his "Genera of Fungi” 1909, 
to consider the Perisporiaceæ more comprehensively and to in- 
clude the Aspergillaceæ in this family, being recognizable by its 
globular, membranaceous or coriaceous perithecia without any 
apical pore. Heleococcum is then naturally placed in this family, 
where it together with Testudina forms a connective link between 
the Aspergillus (Eurotium) on the one hand and the typical Peri- 
sporieae on the other. 

The ripe ascospores already germinate in 24 hours after being 
sown in a hanging drop of sweet wort (Fig. 2 a). When the 
germination takes place at a rather low temperature in which the 
development of bacteria in the culture is repressed, while the 
fungus is not affected, large myceliums can grow out. They consist 





Fig. 2. 
a) Germinating ascospores (X 650). 5) Threads of mycelium with series 
oil drops (X 650). c) brownish, swelled cells of older mycelium. 
97% 


of hyaline, cellular, branched, 5—8 uw broad hyphæ, containing 
a series of bright drops of oil (Fig. 2 b). Older hyphae of the 
my celium become brownish and the cells swell slightly (Fig. 2 c). 
In spite of repeated experiments in transferring mycelia to wort- 
gelatine and sterile earth, I have not succeded in obtaining peri- 
thecia in my cultures. 


Bianco Lunos Bogtr. Kbhvn. 


























£ 


3 5185 00258 9768 








——— — — Ex BE Fry - ——=—=— —. mm em —« —e ” 2 p i > 





é SR DSR RE TESS RER ES PISSE NERE SS SEN ISS SS SES RS TORS SAS SR Re ner er SES ER SAR SES SEE, 

"7 L 278. 23-55 has ge Sne are & 237 å 23842. - 
Se SE SS Ses VST Sme Rs Sen R SR Ur Er LE RE KOR ERE BEDES SEE STETS ESKE TES: ” re fer mn Bet Sø HELE ta | 

rnrdesegnsmertedej mn tedetekrsntrtningr prins kk 38383 ELF TEE SST ET Eee OR EN mile] tel fe re LERNER 


4 = 


SVT ST RT SSR ES 
PRESS KS RES 


renee SEEST EESE 


== 

a— teltet glen 
poserer br 

sas CSE TRT STS 


z Rs 
723 RT STEL DTI BTS 


peer 
mt 


SET BEDT 3 
LOC SLUT ST IS SST YLE 


"35922292 82 


5 RESIDENT TESTE 
SEES SEE = 
Dalskels SS SES DE DES SE RE SOE 


SE 
SDR VER aE 


rr 


St itetat = s 2 å arer 


OD 4 
SES SSP EE