Polytrichadelphus magellanicus sensu lato (Musci) and its ascomycetes different fungi on different hosts

11 Pages·2001·4.1 MB·English

by  Peter Döbbeler

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Polytrichadelphus magellanicus sensu lato (Musci) and its ascomycetes
different fungi on different hosts
P. DÖBBELER
Abstract.
DÖBBELER, P.: Polytrichadelphusmagellanicussens\x\2^.o{M\xsc\)znd'\is ascomycetes
- different fungi on different hosts. - Sendtnera 7: 35^5. 2001. ISSN 0944-0178.
Forty-six herbariumspecimensofthe mossPolytrichadelphusmagellanicuss.l.(Poly-
trichaceae) from all parts of its (sub)antarctic distribution area were analysed for
associated fungi. On thirty-eight specimens fruit-bodies representing ten species of
ascomycetes were found. P. magellanicus s.str. from Patagonia and Tierra del Fuego
was found to be an excellent host, infected by nine fungal species belonging to seven
genera. Each ofthe fifteen infected collections was associated, on average, with four
different fungal species.However,on the twenty-three infectedspecimensof/', inno-
vans fromNew Zealand,the AucklandIslands,AustraliaandTasmania, only Epibryon
pogonati-urnigeri was present with regularity. Except for the unspecific Bryochiton
perpusillusboth hostmosseshave theirownparasites. This underlinesthe distinctness
ofP. magellanicuss.str. from southern South AmericaandP. innovans from Austral-
asia. These taxahave probably developed independently from each other over a long
time period. Relationsbetween P. innovans andPogonatum urnigerumarediscussed.
Zusammenfassung:
Sechsundvierzig Herbarbelege des Laubmooses Polytrichadelphus magellanicus s.l.
(Polytrichaceae) aus allen Teilen des (sub)antarktischen Verbreitungsgebietes wurden
aufPilzbefall hin analysiert. AufachtunddreißigProben konnten Fruchtkörper nach-
gewiesen werden, die zehn Arten von Ascomyceten repräsentieren. P. magellanicus
s.str. aus Patagonien und Feuerlandist mit neun vorkommenden Pilzarten, die zu sie-
ben Gattungen gehören, ein ausgezeichneter Wirt. Jede der fünfzehn befallenen Auf-
sammlungen weist im Durchschnitt vier verschiedene Pilzarten auf Aufden dreiund-
zwanzig Probenvon P. innovans2msNeuseeland,den AucklandInseln, Australienund
Tasmanien ist hingegen fast nur Epibryon pogonati-urnigeri vorhanden, allerdings
mit erstaunlicher Regelmäßigkeit. Abgesehen von dem unspezifischen Bryochiton
perpusillus haben beide Wirtsmoose ihre eigenen Parasiten. Das unterstreicht die
Eigenständigkeit von P. magellanicus s.str. aus dem südlichen Südamerika und P. in-
novans aus Australasien. Vermutlich haben sich beide Sippen über lange Zeiträume
getrennt voneinanderentwickelt.DiskutiertwerdenBeziehungenzwischenP. innovans
und Pogonatum urnigerum.
Introduction
Tworandomly collected specimens ofPolytrichadelphus magellanicus (Hedw.) Mitt. (Polytri-
chales, Musci) from Tierra del Fuego proved to be extremely good hosts for parasitic asco-
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mycetes. Sixandsevenspecies,respectively,wererecorded,representingno lessthan fivegenera
and fourorders.Apparently, hundredsofascomatamay beformedonasinglehostplantwithout
causing any symptoms visible tothe naked eye (Döbbf-ler 1999b).The present study analyses
the fungal diversity within differentpopulations ofP. magellanicus in southern South America
andP. innovans (Müll.Hal.)A.Jaeger in Australasia. The latter taxon often was not separated
fromP. magellanicus(e.g. BEEVERetal. 1992, Fife 1995,Ramsay 1997,Scoit&Stone1976, van
Zanten 1978:460).
Material and methods
Specimens ofPolytrichadelphus magellanicus s.l. from the bryological herbaria B, F, G, GZU,
M W
JE, and werescreenedforthepresenceoffungi.Beforestudyingtheplants it was necessary
to wet them thoroughly. This clearly demonstrates their endohydric organization as well as
highlightingthe waterrepellentlamellaemargins.Smallor sparselydistributedascomatawereonly
detectableby microscopicscreeningofindividualhost leaves.Lugol'ssolution(Merck9261)was
used to test iodine reactions. Infected plants, or parts ofthem, were separated and deposited
underthe fungal name inthecorrespondinghostherbarium.
Specimens examined
Polytrichadelphus magellanicus s.str. as host (specimens 12 and 13, reported by Döbbeler
1996,are included)
1 Chile. Prov. de Cautin, Volcano Villarrica, ca. 1000 m, Hollermayer 56 (W). Recorded species:
Crocicreas magellanicum, Dawsophilapolycarpa, Potriphilaneurogena, Rogellianectrioidea.
2 Chile. Prov. de Cautin,Andes ofVillarrica,(probably 1897),Neger(JE). Recordedspecies: Cro-
cicreas magellanicum, Dawsophilapolycarpa, Potriphila navicularis, P. neurogena.
3 Chile.Prov. deValdivia,Corral,nearQuitaluto, ca.400 m, 2.2.1935,Hosseus698 (JE).Recorded
species: Bryochitonperpusillus, Crocicreas magellanicum.
4 Chile. Prov. de Valdivia, Cordillera Pelada, road from La Union to El Mirador, 800 m, 12.12.
1985, Hellwig 8560 (M). Recorded species: Dawsophila polycarpa, Rogellia nectrioidea.
W
5 Chile. Prov. de Valdivia, West slope ofCordillera Pelada, 8.8 km by road of El Mirador on
road between La Union and Punta Hueicolla, 580 m, 40°07'S, 73°16'W, 18.2.1976, Crosby
J2758 (B, W). Recorded species: Crocicreas magellanicum, Dawsophilapolycarpa.
6 Chile.Prov. deChiloe,Trumao, path to "Madrina", 200 m, 28.5.1932,jM«ge 424{¥). Recorded
species: Bryochtion perpusillus, Crocicreas magellanicum, Dawsophilapolycarpa.
7 Chile. "Waldhagen", [Nothofagus] pumilio-ixQQ line, 1200 m, 1.1.1938, Schwabe 34 (JE).
Recordedspecies:Dawsophilapolycarpa, Epibryon eremita,Potridiscuspolymorphus, Potriphila
navicularis, P. neurogena, Rogellia nectrioidea.
8 Chile. Prov. de Coihaique,Checkpoint Cisnes Medioofthe Carretera Austral, IV.1985, Hellwig
811 (M). Recordedspecies.Bryochitonperpusillus,Crocicreasmagellanicum, Dawsophilapoly-
carpa.
9 Chile.Prov. de Magallanes,Punta Arenas, 1917, Micken(B).Recordedspecies:Croc/'crea^wage/-
lanicum, Dawsophilapolycarpa, Potriphila navicularis, P. neurogena.
10 Chile. Prov. de Antärtica Chilena, Canal Beagle, 11.1922, Gusinde 339 (W). Recorded species:
Crocicreas magellanicum, Dawsophilapolycarpa, Potriphila navicularis, P. neurogena, Ro-
gellia nectrioidea.
1 1 Argentina. Tierradel Fuego,Dep. Ushuaia,RioOlivia,27.9.1979, MatteriNC2153(B). Recorded
species: Bryochiton perpusillus, Crocicreas magellanicum, Dawsophilapolycarpa.
12 Argentina. Tierra delFuego,Dep. Ushuaia,about30 km E ofUshuaia,roadfrom Ushuaiato Paso
Garibaldi, ca. 190 m, 22.3.1997, Mues (M). Recordedspecies: Crocicreas magellanicum, Daw-
sophilapolycarpa, Epibryon eremita, E. interlamellare, Potriphila navicularis, P. neurogena
(Type).
463
. .
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1 Argentina. Tierra delFuego,Dep. Ushuaia,nearPasoGaribaldi,ca.430 m, 20.3.1997, Mues(M).
Recorded species: Crocicreas magellanicum (Type), Dawsophila polycarpa (Type), Epibryon
eremita (Type), E. interlamellare, Potriphila navicularis, P. neurogena, Rogellia nectrioidea
(Type).
1 Argentina. Tierra delFuego australis,Dep. Ushuaia,Montes Martiales, 320 m, 54°46'S,68°18'W,
30.12.1969, Roivainen (Plantae Argentinae 1480, s\xhPolytrichadelphus magellanicus), (G).
Recorded species: Dawsophilapolycarpa.
15 Argentina. Tierra del Fuego, Dep. Ushuaia, Cerro Martial, 350 m, 5.1.1989, Schäfer-Verwimp
& VerM'imp 10849{G).RecoTdcdspecics:Crocicreasmagellanicum, Dawsophilapolycarpa, Epi-
bryon eremita, E. interlamellare, Potriphila navicularis, P. neurogena.
Polytrichadelphus innovans as host
1 NewZealand,(without further notes), com. Dr. Schwarz (W).Recordedspecies: Epibryonpogo-
nati-urnigeri.
17 New Zealand. NorthIsland, NorthAuckland, Waipouaforest, 50 km NW ofDargaville, ca. 300
m, 9.1.1974, van Zanten 7401214 (B). Recorded species: Epibryon pogonati-urnigeri.
18 New Zealand. North Island, Paradise Valley, Fish hatchery, near Rotorua, 7.1.1969, O. & 1.
Degener 31830 (GZU). Recorded species: Epibryon pogonati-urnigeri.
19 New Zealand. North Island, Waiotapu, IX.1929, Meebold (M). Recorded species: Epibryon
pogonati-urnigeri.
20 New Zealand. North Island,Bay ofPlenty, Kaingaroaplantation forest, SE ofRotorua, ca. 600
m, 25.10.1959, van Zanten 1251 (B). Recorded species: Epibryon pogonati-urnigeri.
21 New Zealand. North Island, Tauhara,Taupohill, about 2500', IX.1929, Meebold (M). Recorded
species: Epibryon pogonati-urnigeri.
22 New Zealand. NorthIsland, Waitakerei Range, 1206', IX.1929, Meebold(M). Recordedspecies:
Bryochiton perpusillus.
23 New Zealand. North Island, Gisbom district. Lake Waikaremoana, along road from
Waiotukupuna-bridgeto CascadeFalls,ca. 600m, 29.8.1968, vanZanten 681886(iE). Recorded
species: Epibryon pogonati-urnigeri.
24 New Zealand. North Island, Mt. Egmont, 3000', VIII.1929, Meebold (M). Recorded species:
Epibryon pogonati-urnigeri.
25 New Zealand.North Island,Mt. Egmont, DawsonFalls, 6.3.1959,H. & E. Walter(M).Recorded
species: Epibryon pogonati-urnigeri.
26 New Zealand. North Island, Hawke's Bay, near Whakarara, Gardiner's Mill, 31.12.1950,
Sainsbury (W). Recorded species: Epibryon pogonati-urnigeri
27 New Zealand. North Island, Wellington, York Bay, V.1929, Meebold (M). Recorded species:
Epibryon pogonati-urnigeri.
28 New Zealand. SouthIsland, Westland,Teramakau Valley, 600 ft, 6.2.1903, NaylorBeckett(JE).
Recorded species: Epibryon pogonati-urnigeri.
29 New Zealand.South Island,West CoastArea, Nof Whataroa, 7.2.1970, Vitt 4505 (B).Recorded
species: Bryochiton perpusillus. Epibryon pogonati-urnigeri.
30 NewZealand.South Island,SouthOtago,The Chaslands,alonghighway93, 14.1.1973, Vischiß).
Recorded species: Epibryon pogonati-urnigeri.
31 New Zealand. Stewart Island, 1.1889, Bell (M). Recorded species: Epibryonpogonati-urnigeri.
32 Auckland Islands. Auckland Island, South end of North Harbour, 12.1.1973, Vitt 10223 (M).
Recorded species: Epibryon pogonati-urnigeri
33 Australia. Victoria, Baw Baw National Park, 6 km SE ofMt. Erica, road cutting on Mt. Erica,
700-800 m, 37°35'S, 146°23'E, 10.1.1986, Rambold 3276 (M). Recorded species: Epibryon
pogonati-urnigeri.
34 Australia. Victoria, Baw Baw National Park, 6 km SE ofMt. Erica, road cutting on Mt. Erica,
900-1000 m, 37°35'S, 146°23'E, 10.1.1986, Rambold 3280 (M). Recorded species: Epibryon
pogonati-urnigeri.
35 Australia.Victoria,Mt. Donna Bvang,5km N ofWarburton, 1200m, 12.4.1974, Streimann 991
(B). Recorded species: Epibryon pogonati-urnigeri.
36 Tasmania, (without further notes), Lucas (M). Recorded species: Epibryon pogonati-urnigeri.
37 Tasmania. NWofMt. Field,Florentine Valley,21.1.1959, H. &E. Walter(M). Recordedspecies:
Epibryon pogonati-urnigeri.
38 Tasmania. Hobart, Mt. Wellington, 14.4.1903, FleischerB2074 {B).Recordedspecies:Epibryon
pogonati-urnigeri.
The ascomycetes recorded
BiyochitonperpusillusDöhhdQr(DÖBBELER, Mitt.Bot. Staatssamml.München 14: 226, 1978)
Hosts: Polytrichadelphus magellanicus, P. innovans (and otherbryophytes, see belowj
Distribution on Polytrichadelphus: Chile, Argentina,NewZealand
Specimensexamined: 3, 6, 8, 11, 22,29
Bryochitonperpusillus is a very widespread and variable species recorded from distantly
related bryophytes, often with a withered appearance, indicating that taxonomic uniformity
shouldbeproved. Partially damaged hepatics, Wke.Ptilidium occurwithin the spectrum ofhosts
besides the Polytrichaceae, mcXxxdmg Dawsonia (Döbbeler 1978, 1981). B. perpusillus most
regularly colonises Polytrichum piliferum Hedw. where it develops subcuticularly from a
conspicuous network ofthick brown hyphae, especially in the apical, abaxial region ofnormal
lookingorhardlyaltered leaves. On Polytrichadelphus, Bryochitonperpusillus occurs in lower
concentrations. Ascomata are sparse on the lower, withering leaves ofa plant, suggesting the
saprophytic tendency ofthe fiingus. Generally, apical regions ofindividual leaves arepreferred
and oftenboth sides ofaleafhavefruit-bodiesas well asthe interlamellarspacesbeing infected.
Herethe ascomata may lookvery similartothoseofEpibryonpogonati-urnigeri. However, the
latter speciesprefersupperhealthyleaveswherethe fungusis completelysurroundedbythe leaf
lamellae. A positive iodinereaction withintheostiolarregion immediately reveals E. pogonati-
urnigeri.
Crocicreasmagellanicum Döbbeler(Döbbeler, HaussknechtiaBeih. 9: 81, 1999b)
Host: Polytrichadelphus magellanicus
Distribution: Chile, Argentina
Specimens examined: 1-3, 5,6, 8-13, 15
Due to a lack of contrast with its substrate, the white to nearly hyaline apothecia of
Crocicreas magellanicum areoftendifficultto detect, especiallywhenhiddenbetween the leaf
lamellaeandsparse innumbers.Microscopicallyscreenedleavesrepeatedlyprovedtobeinfected,
though apothecia could notbe seen by lowstereo-microscopicmagnification.
DawsophilapolycarpaDöbbeler(Döbbeler, HaussknechtiaBeih. 9: 83, 1999b)
Host: Polytrichadelphus magellanicus
Distribution: Chile, Argentina
Specimensexamined: 1,2,4—15
The two species Dawsophila callichroma Döbbeler and D. pygmaea Döbbeler frequently
colonisedifferentmembersofthegenusDawsoniaandarepresent within its wholeAustralasian
distribution range (Döbbeler 1981). Though special attentionwaspaidto seeking Dawsophila
on Polytrichadelphus innovans, which shares its distribution forthe most part withDawsonia,
ascomatacouldnotbedetected.
Epibryoneremita Döbbeler(Döbbeler, HaussknechtiaBeih. 9: 85, 1999b)
Host: Polytrichadelphus magellanicus
Distribution: Chile, Argentina
Specimensexamined: 7, 12, 13, 15
Several ascomata ofcollection 15 had approximatelythirty-two sporesper ascus unlike the
eight spores perascus found in theotherthree specimens, including the type specimen. Spore
sizeamongspecimenswas similar.Thelargerasci (upto 60 fimlong) foundin the32-sporedasci
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simplyaccommodate a largersporenumberthanthesmallerasci (35-40 ^m long). As thereare
no other differences to the type material, spore number and ascus size do not seem to be of
taxonomic significance. RelationstoEpibryonelegantissimum Döbbeler on Dawsonia superba
Grev. inNewZealand(Döbbeler 1981) mayexist,judgingby fruit-bodymorphology,hymenial
featuresandthe peculiarmicrohabitatpreference.
Epibryoninterlamellare(Racov.) Döbbeler(Döbbeler, Mitt. Bot. Staatssamml. München 14:
279, 1978)
Hosts: Polytrichadelphus magellanicus (andotherPolytrichaceae)
Distribution onPolytrichadelphus: Argentina
Specimensexamined: 12, 13, 15
This frequentandwidespreadspeciesinfectsdifferentPolytrichaceae (seeDöbbeler 1999b).
Epibryonpogonati-urnigeriDöhheler(Döbbeler, Mitt. Bot. Staatssamml. München 14: 296,
1978)
Hosts: Polytrichadelphus innovans (and otherPolytrichaceae, seebelow)
Distribution onPolytrichadelphus: NewZealand,Auckland Islands, Australia, Tasmania
Specimensexamined: 16-21,23-38
The young, upperleavesofan infectedplantnormallycontainonly afewascomatawhichare
restrictedtothe apical leafregion. Due to a lack ofspace, they are very laterally compressed.
Oftenthe ascomatal walls adjacentto the lamellae are reduced or completely missing. In older
leavesascomataare moreevenlydistributedand spherical.Heavily infected,6mm longleavesof
differentcollectionscommonlyyielded abouthundredascomata Inonecasean individualleafof
mm
8 length was colonised by approximately 180 ascomata. The species is quite uniform on
Polytrichadelphus innovans, although slightdifferences inspore size and otherfeatures between
collectionson differentPolytrichaceaeorfrom differentgeographicalregionsapparently exist.An
essentialfeature oiEpibryonpogonati-urnigeriisits iodinereaction,which isnot knownin other
bryophilous ascomycetes. The ostiolar region turns reddish in Lugol's solution and KOH
pretreatmentgives amore distinctbluish colour(hemiamyloid reaction, Baral 1987).
Four polytrichaceous hosts are known for Epibryon pogonati-urnigeri in addition to
Polytrichadelphus innovans, namely Pogonatum urnigerum (Hedw.) P.Beauv., P. dentatum
(Brid.) Brid. (Döbbeler 1978, 1985), Dawsonia longiseta Hampe and D. polytrichoides R.Br.
(Döbbeler 1981). Withthe exception ofP. dentatum, for which only a few data are available,
these mosses are very regularly or (in certain regions) even indiscriminately infected by E.
pogonati-urnigeri. Both species ofPogonatum represent the subgenus Dendroidea (Hyvönen
1989). It is rather well documented that E. pogonati-urnigeri occupies wide areas of the
geographical range ofP. urnigerum (Döbbeler 1978, 1985) which is distributed in northern
Eurasia, North Americaand in scattered high mountain localities in Africa, southeast Asia and
PapuaNew Guinea (Hyvönen 1989). Afterstudyinghundreds ofEuropean Polytrichaceae it is
obvious that species like Pogonatum aloides (Hedw.) P.Beauv. {P. subgen. Pogonatum) or
Polytrichastrumalpinum(Hedw.)G.L.Sm.(syn. Pogonatumalpinum(Hedw.)Röhl.)do nothost
E. pogonati-urnigeri. Dawsonia longiseta and D. polytrichoides make up the more primitive
section ofthegenusDawsonia(D. sect.Dawsonia)andarerestrictedto eastern Australia (van
Zanten 1973). On studyingrepresentative materialDöbbeler (1981)observedthatascomataof
Epibryonpogonati-urnigeri were always present on both D. longiseta and D. polytrichoides
whereas forD. superba (D. sect. Superba) the situation is quite different. Though occurring
within the range of the parasite in New Zealand, Australia and Tasmania, parasitism by
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E.pogonati-umigerioccurredonly exceptionally. The mosses in question are not ecologically
separate as they (as most Polytrichaceae) preferopen, exposed soil as on river banks and road
cuttings.
How should one interpret the range of hosts for Epibryon pogonati-umigeril It exhibits
specificity between sections within Dawsonia and Pogonatum and between Pogonatum
urnigerum andPolytrichastrumalpinum, but itdoes notdiscriminate amonggeneraas different
asPogonatum, PolytrichadelphusandDawsonia.The differences betweenthese generamaynot
be as great as one might initially suppose because they are distinguished by sporophytic
characters, i.e. capsule morphology. The gametophytic plants are well-developed and share
several common features. The leavesare characterised bya distinct sheath and swelling tissue.
Theybeardensely formedadaxial lamellaewith specialisedmarginal cells which are covered in
Polytrichadelphus andDawsonia with angular flakes ofepicuticular wax. In this respect they
differfrom other Polytrichaceae (Proctor 1992). Though species oiPogonatum vary greatly
regardingwaxy deposits,P. urnigerum appears "to behighlydistinctive,with epicuticularwax
onthepapillose lamellamargins in the form offlakes orplateletsnotunlikethose oiDawsonia
and Polytrichadelphus'' (Clayton-Greene et al. 1985: 552; see also Proctor 1979). On a
macroscopic level, the upperleafsurfaceofthesemosses displays aglaucous bloom caused by
the waxy coverings. Another leaf character common to Pogonatum urnigerum and
Po/v/nc/7flJe//7Az/5;>7«ovö«5aresimilarlamellaehairs(Döbbeler 1999a,Figs.4, 5). Nevertheless,
recentmolecularresults on thephylogenyofthe Polytrichales byHyvönen etal. (1998) do not
support avery close relationshipbetween thegenera in question.
PotridiscuspolymorphusDöbbeler&Triebel (Döbbeler&Trjebel, Hoppea 61 72, 2001)
:
Hosts: Polytrichadelphus magellanicus (and other Polytrichaceae, especially Polytrichum
juniperinum Hedw., Döbbeler&Trjebel 2001)
Distribution onPolytrichadelphus: Chile
Specimenexamined: 7
This distinctive species with a palaeoaustral distribution is more frequent than previously
thought. Polytrichadelphusseemsonly to bean occasionalhost. Withinaratherrich collectionof
P. innovans from New Zealand (collection 20) a few intermingled plants of Polytrichum
juniperinum wereheavily infectedbyPotridiscuspolymorphus, whereasno ascomata could be
detectedon thepredominantPolytrichadelphus.
PotriphilanavicularsDöbbeler(Döbbeler,Nova Hedwigia62: 62, 1996)
Host: Polytrichadelphus magellanicus (andotherPolytrichaceae)
Distribution onP. magellanicus: Chile, Argentina
Specimensexamined: 2, 7, 9, 10, 12, 13, 15
Potriphilanavicularishasa bipolardistributiontype and is very frequentonPolytrichastrum
alpinum (Döbbeler 1996). On Polytrichadelphus magellanicus, as in nearly all southern
hemisphererecords,onlythe anamorphicstate is present. Inadditionto normalleaves,bractsand
scale leaves shouldalsobe screened. Usually only a few pycnidia are detectable on dozens of
potential host plants. The host of Potriphila navicularis from the Auckland Islands is not
Polytrichadelphus magellanicus, as recorded by Döbbeler (1996), but Polytrichastrum longi-
setum (Brid.)G.L.Sm.
41
Potriphilaneurogena Döbbeler(Döbbeler, HaussknechtiaBeih. 9: 88, 1999b)
Host: Polytrichadelphus magellanicus
Distribution: Chile, Argentina
Specimensexamined: 1, 2, 7, 9, 10, 12, 13, 15
Ascomata also develop within male bracts surroundingtheantheridia.
Rogellia nectrioidea Döbbeler(Döbbeler, Haussicnechtia Beih. 9: 91, 1999b)
Host: Polytrichadelphus magellanicus
Distribution: Chile, Argentina
Specimens examined: 1,4, 7, 10, 13
In several cases the ascomata ofboth Rogellia nectrioidea and Potriphila neurogena were
observedonthe samehostleaf.
Geographical range ofPolytrichadelphus magellanicus and P. innovans
Austral populations ofPolytrichadelphusdiXQoftenregardedasasingle species,P. magellanicus.
All of the herbarium material cited (specimens 1-38) was determined as P. magellanicus
(specimen 7 asP. horridus Mitt.). However, Smith(1971) considersPolytrichadelphus ofNew
Zealand and Australia as a distinct taxon, P. innovans. "Compared with the American
P. magellanicus, theplants aresmallerandmoreslender,withshorter,subsecundleaves.The leaf
sheath is broader,with more pronouncedshoulders,and the teethonthe marginoftheblade less
well developed" (Smith 1971: 41). He additionally noted differences in capsule morphology.
Moreover, the presence oftwo distinct species is clearly indicated by completely different
lamellae hairtypes as documentedbyDöbbeler(1999a) and repeatedlyobserved in this study.
Ifone acceptsP. innovansas adistincttaxon, thenP. magellanicuss.str. isconfinedtoChileand
Argentinasouth ofabout40 latitude, the Fuegian region, Juan Fernandez Islands, the Falkland
Islands, Tristan daCunhaandGough Island (Greene 1986, He 1998, Proctor 1992, Robinson
1975, Schjavone 1993, Schiavone & HyvOnen 1992, Smith 1971). The assumed disjunct
occurrencein thenorthernAndesofPeru(Schultze-Motel&Menzel 1987,Gradstein&Frahm
1987, Menzel 1992) is based on a BRYOTROP-specimen [Peru. Dep. Amazonas, Prov.
Chachapoyas, road Chachapoyas-Cajamarcakm 413, mountain rain forest, 3000 m, 5.9.1982,
Frahm, Geissler, Gradstein, Philippi& Schultze-Motel916(Herbarium Frahm!). Femaleplants
with only one capsule and several successive, apparently lateral setae typical for the genus
Polytrichadelphus.]. This species clearly differs by habit and leaflamellae characters from the
southern South American Polytrichadelphus magellanicus. The Australasian P. innovans is
known fromNewZealand,southeasternAustralia,Tasmania,theAucklandIslandsandCampbell
Island (Smith 1971, Vitt 1974, 1979).
Comparison ofthe ascomycetes on Polytrichadelphus magellanicus and P. innovans
Careful analysis of forty-six collections of both species of Polytrichadelphus revealed the
presence often different ascomycetes, eight only present on the American, one only on the
Australasiatic host and one common to both. Ofthe seventeen collections ofPolytrichadelphus
magellanicus from Chile and Argentina, fifteen are infected by one to seven species of
ascomycetes. Altogetherfifty-eight specimensofascomycetesarefound,onaveragenearlyfour
42
on each host collection. Thus P. magellanicus represents one ofthe best hosts, not only within
Polytrichaceae but also within bryophytes in general, with regard to the number ofassociated
fungi andthefrequencyofinfection.Dawsophilapolycarpa, themostcommonspecies,is present
on fourteenhostspecimens;Crocicreasmagellanicum,ontwelve;Potriphilaneurogena, oneight;
P. navicularis, on seven; Rogellia nectrioidea, on five; Bryochiton perpusillus and Epibryon
eremita, on fourandPotridiscuspolymorphus, on one host collection. Twenty-eight specimens
ofPolytrichadelphus innovans from New Zealand, Australia and Tasmania and one collection
from the Auckland Islands were examined. Twenty-three proved to be colonised. Epibryon
pogonati-urnigehwas present on twenty-two specimens, whereas Bryochiton perpusillus was
encountered only twice. This means that on average only one associated fungus per host
collection was found.
The present inventory should not imply that the fiingal diversity ofthe two austral species
of Polytrichadelphus is completely covered. Only the more frequent, previously described
parasitesare discussed. On several occasionsascomataofan undescribed speciesweredetected
but are not discussed in this paper. In most cases the potential hosts consisted ofonly a few
plants gathered for bryological purposes. More extensive collections are required to obtain a
sufficient number of ascomata for a proper investigation of the other sparsely distributed
parasites. Fieldstudiesarealsorequiredto seekoutthe relativelyconspicuous,necrotrophicflmgi
that areunlikelytobe representedinbryophyteherbariummaterial.AnexampleisLizonia,which
occupies the antheridial cups ofvarious polytrichaceous mosses. One should also search for
orange coloured apothecia formedontheground between individual plants, asthey may belong
to rhizoidparasitic speciesofOctospora orLeucoscypha. WhetherPolytrichadelphusbelongsto
thehostrangeofthe lattergeneraandalsoLizonia isnotcurrentlyclear.
Interpretation ofbryomycological evidence
The most surprising result of this bryomycological analysis is the diversity of the fungal
associates ofthe two austral species ofPolytrichadelphus studied. Only one species out often
was encounteredseveraltimeson bothhosts, namtXyBryochitonperpusillus, avery commonand
critical taxon which is also recorded from hepatics. Five ascomycete species are known
exclusivelyonP. magellanicuss.str.,amongthemPotriphilaneurogenaandRogellianectrioidea.
The lattertwo species develop within the leafnerves, a distinct niche rarely occupied by the
multitude ofpolytrichophilous ascomycete species.
The marked differences in suitabilityofthe two moss species as hosts was also unexpected.
Polytrichadelphus magellanicus isanexcellent substrate, whereasP. innovans is lessattractive.
The absence ofcommon fungal parasites supports the distinctness ofboth moss taxa as has
already been shown by morphological features. Another question concerns the phylogenetic
relationship ofthe Polytrichadelphus species. Geographical isolation even for long geological
periods does not automatically lead to genetic diversification as demonstrated by Frey et al.
(1999)usingmolecularmethods.Inthe ancientLopidium concinnum(Musci) which hasexactly
thesamedistribution rangeasP. magellanicus s.l. littlegeneticdivergence has occurred within
the isolated populations since theirseparationbygeotectonic events about 80-60 million years
ago.
The vicarianttaxaF. magellanicusandP. innovans{Smith 1972:46)are "undoubtedlyclosely
related"(Proctor 1992:317).Thisinferredrelationshipisnotmirroredby anoverlapofthemore
significant flingal parasites. Furthermore, Dawsophila frequently colonises various species of
Dawsonia (Döbbeler 1981) and P. magellanicus, but avoids P. innovans. Epibryon elegan-
43
tissimumoccurringonDawsoniaandE. eremita infectingP. magellamcus seem to represent a
furtherlinkbetweenAustralasianandSouthAmericanpolytrichaceoushosts, leavingP. innovans
apart. The bryomycological facts documented herein indicate that P. magellamcus and P. in-
novans have hada very long,separate evolution. It wouldbe interestingto know moreabout the
ascomycetes on the relatives ofPolytrichadelphus, both in South America and Australasia.
I am indebted to the curators ofthe herbaria B,F, G, GZU, JE, M and W for loans ofspecimens.
Prof Dr. F. Hellwig(Jena)providedseveralspecimens collectedinChile manyyears ago.Dr. H. No-
wak-Krawietz (Berlin) helpedwith the literature. Prof Dr. W. Frey (Berlin)and Prof Dr. H. Hertel
(München) madecomments on an early draftofthe manuscript.Iam gratefulto Prof Dr. T. H. Nash
III (Tempe) and M. CuUen (Dublin) for linguistic correction ofthis paper.
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Table 1: The hosts, the associated ascomycetes and their frequency (by number of specimens
recorded).
Fifteen specimens ofPolytrichadelphus magellanicus s.str.
(southern South America) are infected by: