Notes on Hymenoscyphus - II: On three non-fructicolous species of the 'fructigenus-group' with croziers

PERSOONIA

Published byRijksherbarium/ Hortus Botanicus,Leiden Volume 16, Part 2,pp. 191-207 (1996)

Notes on Hymenoscyphus — II.

On three non-fructicolous

species

of the

_{‘fructigenus-group’}

with croziers

Jan Hengstmengel

Rijksherbarium/HortusBotanicus,P.O. Box9514,2300 RALeiden,The Netherlands

Descriptions, illustrations,and_keysaregivenof_asmall_partof the _{‘fructigenus-group’}of theascomycetegenus Hymenoscyphus.Thespeciesconcerned arecharacterizedbyasci orig-inatingfrom croziers.HymenoscyphusscutuloidesandH. fucatusvar.badensis are describ-ed asnew,whilethe combination H.fucatusisvalidlypublished.

D Also called _{‘scutula-stirpe’} (Dennis, 1956:66, 82;cf.White, 1944:609, 613)or'caudatus-group'

(Dumont,1981:60),but_preferablynamed_{'fructigenus-group'}because H._fructigenus(Bull.: Fr.) S.F.

Gray,thetypespeciesof thegenus,formspartof it. 2) _{From H.}

scutula,a _{member of the group.}

The taxonomiccoreof the genus

Hymenoscyphus

S.F.

_Gray

is formed

_by

_a

homoge-neous group of

species

which have the same

stipitate-cupulate habit,

the_same

_excipular

structure, and almost thesame

shape

of

asci,

ascospores, and

paraphyses.

This group, here called

_{‘fructigenus-group’}

comprises

some dozens of

_lignicolous

_(inch

fructi-colous),

caulicolous, and foliicolous

_species,

such as H. albidus

(P. Karst.)

W.

_Phill.,

H.

albopunctus

(Peck)

Kuntze, H. caudatus

_{(P. Karst.)}

Dennis,H.

_{fastidiosus (Peck)}

Arendholz,the H.

_{fructigenus-complex,}

H. humuli

_{(Lasch) Dennis,}

H. salicellus

_(Fr.:

Fr.) Dennis,

H. scutula

(Pers.: Fr.)

W.

Phill.,

and H. serotinus

_{(Pers.: Fr.)}

W. Phill. Their

_exciple

is

_composed

oftwo

_layers:

the inner

layer (medulla)

is_athin-walledtextura

porrecta,theouter

_{layer (cortex)}

isa

mostly

thin-walledtextura

prismatica.

The

_excipular

hyphae

are

running parallel

withororientedata low

angle

tothe

_excipular

surface and

they

arenotembedded in_a

_gelatinous

matrix. Characteristic of the

_{‘fructigenus-group’}

are

the

_{ellipsoid-fusiform}

toobovoid-fusiform ascospores. Such spores show the upper half ofan

ellipsoidal

toobovoidal

_body

and the lower half ofafusiformone.Thesporesarein

general approximately bilaterally symmetrical, owing

toan

abaxially angulate

oreven

hooked apex,

occasionally

combined withsome

slight curving

or

single-sided flattening

of the spore. This

particular shape

is

_nowadays

often indicatedas 'scutuloid'2

, a

term intro-duced

_by

Baral

_(in

Baral &

_{Krieglsteiner,}

1985:

_120).

As the

_{torpedo shape}

is

_providing

foran

optimal discharge,

some asymmetry causes the spores torotate round their axis. This

_spinning

can

easily

be seenunder the

_microscope,

at least inmountsof

rehydrated

material in ammonia 10%.Moreover,the spores of several

species,

e.g.H. salicellus and H.

_scutula,

are

provided

attheir ends withone or more

hyaline,

thread-like

appendages,

commonly

indicatedas 'cilia' but- because of their

immovability

and

consistency

-

pref-erably

called 'setulae'

_(bristles).

Whether these

_appendages

haveafunction in

spinning,

in attachmentto substrata, or in any other process, is unknown

(cf. Hawksworth,

1987:

Most

species

of the

_{'fructigenus- group’}

contain asci which

_originate

from

aporhyn-chous

_Dangeardian

elements. Hence

_they

lack an

anastomosing

archattheir base. In this article attention is

_paid

tothree

_species

whose asci arise from

_{pleurorhynchous}

Dangear-dianelements,alias

_croziers,

and

_generally

do showabasal

_anastomosing

_{arch. The}

pres-enceof croziers and arches

_respectively

is, indeed,notthe

_only

character

_by

which these three

_species

are

distinguished

from others. Ifso,I wouldnothave

_regarded

themas sep-arate

species

but as taxa ofan

infraspecific

rank. Such is the caseine.g.Phaeohelotium

3) Also called _'tip'(e.g.Giiumann &_Dodge, 1928:130)or'downward protuberance' (Huhtinen, 1990:

66-67).

4) _{Also called}

'stipe' (Gaumann&Dodge, 1928: 130),'stalk cell' (Huhtinen,1990:67)or_'pedoncule'

(Martens, 1932:259).

3) Haines(1989:315) simultaneouslyintroduced 'crozier'[gen.'crozieris',pi. 'crozieres']asaLatinterm

('cum crozieribus'),but thereisnoneed for suchanobjectionable neologism.

As

_{generally known,}

asci arise atthe

_{(provisional)}

_{end of ascogenous}

_hyphae,

from 2-or3-celledstructurescalled

_'Dangeardian

elements'

_(see

_e.g.

_Chadefaud,

_1943).

With-in the

_inoperculate

_ascomycetes two_{main types of such} structureshave been

_found,

viz. the

_{pleurorhynchous}

_typeand the

_aporhynchous

_type

_(Berthet,

1964: 98 et

seq.).

In the

pleurorhynchous

typethesnoutof the elements is

_laterally

turned offoveran

angle

of about 180°. Hence such elementsare

hook-shaped

and called 'crozier'

_(French

'crochet

(ascogene)',

German 'Haken', Latin

_'uncus').

In the

_aporhynchous

or

(secondarily)

hookless _typethe

forming

of_a lateralsnout is

_thought

tobe

_suppressed,

sothat the asci

seem tobe

_{'simple-septate'}

_(as

called

_by

Huhtinen,1990:

66-67).

When the foremost binucleate cellor'crook' ofacrozier

_develops

intoanascus,the terminal

(uninucleate)

cell3

mayremain visibleasa small

by-pass arching

over the_septumbetween the_ascus and the

_preceding

cell4

.

A_separateterm for this small archseems not tobe incommon

use, atleast in

_English

literature. For thatreasonseveral authors have resortedtoa

cir-cumscription

as 'Asci

_{originating/produced/arising}

from croziers'

_(e.g.

White, 1943; Dumont,

1976;

Korf & Lizon,

1994)

or 'Asci aus Haken entstehend'

_{(e.g. Arendholz,}

1979),

without

making

clear whether the arch remains visible. Some other authors have

misapplied

the term 'crozier' tothe arch

_{solely (e.g.}

Dennis, 1956:76, 79; Haines,

19895

_).

_However,

a similarprocessof

forming

arches is well known frommany

basi-diomycetous

fungi,

viz. in the

_development

of

_hyphae

of the

_secondary

_mycelium

(incl. basidia).

In these

_fungi,

the

_anastomosing

arch

_has,

formorethan acentury,been called

_{'clamp-connection'}

or,

shortly,

'clamp'

(Fr.

'boucle' or 'ansed'anastomose',

Germ.

_'Schnalle',

Lat.

_'fibula').

Since there is

_general

_agreementon the

homology

of

early development

of basidia andasci,andonthe

homology

of

_{clamp forming}

in

basidio-mycetesand arch

_forming

in_ascomycetes

_(see

_e.g.Martens, 1932:261; Moreau, 1950; Moreau, 1954:

_1563;

Berthet, 1964:98,

99,

118; Arx, 1967: 183, 184; Boidin, 1971:

143,

144;

Dorfelt,

1989:

_47;

cf. Gaumann &

_Dodge,

1928:421,

422),

it_seems

_justified

and-forcorrect

interpretation

-advisableto

apply

theterm

'clamp(-connection)'

to

rele-vantascomycetes too.

Yet,

in several dictionaries the

_'clamp'

is considered_{as a}character

unique

to

_{basidiomycetes (e.g.}

Hawksworthet

al., 1983; Dorfelt,

1989).

Iwould, how-ever,advocatealess exclusive

application

of thisterm.

Accordingly

Moreau & Moreau

193 Hengstmengel: Notes onHymenoscyphus-II

(Hymenoscyphus)

imberbe

(Bull.: Fr.) Svrcek,

whereintwootherwiseidentical forms

canbe

distinguished:

onewithcroziers andanother with

aporhynch

elements.

Since therewas nofresh materialavailable, the

presented full-descriptions

have been based_ondried

_specimens

whichwere

rehydrated

in ammonia 10%.

Microscopical

obser-vation and

_measuring

werecarriedouton hand-made sections and

squash preparations

supplied

withasolution of 0.2%cottonblue in lacticacid.

Measurements ofasci, ascospores and

paraphyses (length

of end

_cells)

arebased on

samples

takenatrandom,with the

_exception

ofextremes

(between

round

_brackets)

which

arebasedonselect elements outside the

sample(s)

referredto.The number of

samples (k)

and the number of elements per

sample (n)

areindicated between squarebrackets,

just

as

the calculatedtaxon averages

(length,

width,and

_{length/width-ratio respectively)}

and standard deviations.

KEY TOTHE SPECIES TREATED

la. Asci 72-90

pm

long (average length

< 87.0

pm). Ascospores

14-21 pm

long

(average length

< 19.0

_pm),

with

_{length/width-ratio}

of 3.4-5.9

_(average

ratio<

_5.00),

occasionally

apiculate

butnot

obviously

setulose 3. H. menthae b. Asci 80-136pm

long (average length

>87.0

pm). Ascospores

18-36

pm

long

(av-erage

length

> 19.0

_pm),

with

_{length/width-ratio}

of 4.8-8.8

_(average

ratio>

5.00),

mostly provided

attheendswith

_obvious,

_upto4.0 pm

long

setulae 2 2a.

_Ascospores

18-27x_{3-4 pm}

_{(average length}

<24.0 pm, average width<4.0

_pm),

with

_{length/width-ratio}

of 5.1-7.7

_(average

ratio>

_5.50);

setulae 1.0-4.0 pm

long

2. H. scutuloides

b.

_Ascospores

23-36x4-6 pm

(average length

> 24.0 pm, average width> 4.0

pm);

setulae 1.5-2.5 pm

long

'•

_{H. fucatus}

1.

Hymenoscyphus

fucatus

_{(W. Phill.)}

Baral &

_Hengstm.,

comb. & stat. nov.

Peziza fucata Cooke & W. Phill.inherb, [invalid: ined.];in Cooke,Grevillea 4(1876) 132,pi. 65,

fig. 300 [invalid:nomennudum;seealso Carpenter,Mem. NewYork Bot. Garden 33 (1981) 214].— Hymenoscyphusfucatus (Cooke& W.Phill.)Baral inBaral &Krieglsteiner,Beih. Z._Mykol. 6(1985)

128[as '(Phill.)';invalid: nomennudum],

Pezizascutulavar.fucataW.Phill.,Elv. brit. (3) (1877)n. 120[invalid:nomennudum].— Hymeno-scyphascutulavar.fucataW.Phill.,Man. Brit.Discomyc. (1887)137 (basionym).—Phialea scutula

var.fucata (W. Phill.) Sacc.,Syll.Fung.8 (1889)266. —Helotium scutula var. fucatum(W. Phill.) Rehmin Rabenh. _Krypt.-Fl.ed.2, 1(3) (1893)793 [asvar. ‘fuscata’]. —Helotium scutula forma

fuca-tum(W. Phill.) Massee,Brit.Fung.-fl.4(1895)254[asforma‘fucata’].

Helotium _superbumVelen.in herb. etms. 1923 [invalid: ined.]fide SvrCek(1985: 159, 188).— Helo-tium macrosporum Velen., Monogr.Discomyc. Bohem. 1(1934)194 [illegitimate:laterhomonym];non

Helotium macrosporum Peck,Ann.Rep. State Bot. 26(1874)82.

Hymenoscyphus

fucatus

has

relatively long

and wide _{spores. Two other}

non-ligni-colous

_species

of the

_{‘fructigenus-g}

roup’,

viz. H. dearnessii

(Ellis

&

_Everh.)

Kuntze and

H._suspectus

(Nyl.) Hengstm.,

have ascospores whicharesimilar in

length,

butnarrower

Hengstmcngel: Notes on Hymenoscyphus-II 195

Foranextensive

description,

anenumeration of the examined collections and further

comments,seeunder the

distinguished

varieties.

KEY TO THE VARIETIES

a. Asci 83-101x9-11 pm.

Ascospores

23-35 pm

long (average length

<27.5

_pm)

and

4-5 pm wide b. H.

_fucatus

var. badensis

b. Asci 113-136x 10-14 pm.

Ascospores

24-36 pm

long (average length

>27.4

pm)

and 4-6 pm wide a.H.

fucatus

var.

fucatus

1a.

Hymenoscyphus

fucatus var. fucatus—

Fig.

1

Apothecial morphology

—

Apothecia stipitate-cupulate,

uptoabout 1 mm

high

when

rehydrated, loosely clustered, rarely mutually

grown

together

atthe_{base, erumpent}

_through

(locally blackened) epidermis

or

superficial

ondecorticatedpartsof the

substratum;

some

dozens uptomorethanahundred

apothecia

over a

length

of 10cmof the substratal stem.

Cupule saucer-shaped,

upto0.8mm in diameterwhen

rehydrated. Receptacle

and

_stipe

smoothto

subpruinose. Stipe cylindrical

to

obconical,

uptoabout 0.8mmwhen

rehydrat-ed,

aboutas

long

asthe diameterof the

cupule.

Anatomy

—Asci

[k

= _1,n=

_{10] cylindric-obconical}

to

_{cylindric-clavate,}

113-136_pm

long [average

length

± standard deviation: 123.8 ± 8.2

_pm],

_{10—12(—14)}

_pm wide

[average

width ± standard deviation: 11.1 ±0.7

_pm],

with

_{length/width-ratio}

of 9.7-13.7

[average

ratio ± standard deviation: 11.2 ±

1.2], 8-spored, originating

fromcroziers;apex truncatedconical; annulus

turning

medium blue in Melzer's _reagent.

_{Ascospores [k}

=2,

n=

20]

obovoid-fusiformto

_{ellipsoid-fusiform,}

sometimesalmost

_{cylindrical, straight}

to

slightly

curved,24-36 pm

long

[29.1± 2.7

_pm],

_{4-6 pm wide}[5.0± 0.3

_pm],

with

_length/

width-ratio of 4.8-7.2

_[5.8

±

0.6],

1-celled when mature,afterwards

2-celled, hyaline,

thin-walled, smooth, provided

with 1-2

_large

orabout 4-6 medium-sized and

occasion-ally

somesmall

guttules, obliquely biseriate,

atapexand base

mostly provided

with 1-2

(-3) short,

upto

1.5(—2.0)

pm

long, straight

or

slightly

curvedsetulae; apex roundedto

abaxially angulate,

sometimesbeaked;base

_(almost)

acute, sometimes ratheracute.

Para-physes

[k

= _{1, n} =

_{10] subcylindrical,}

1.5-2.0 pm

_wide,

atthe_topoften

_slightly

wider thanatthebottom,

hyaline,

according

toBaral

_{(in litt.) provided}

with

_highly

refractive

guttules

when

_fresh,

with 3-5_septain the_{uppermost100 pm, forked about}

_half-way

(some-times also in upper

half);

terminal cell 22-40 pm

long

[29.0± 6.0

_pm],

0.9-1.5 timesas

long

asthe subterminal cell

_[1.3

±

_0.2],

with rounded

_tip;

_{subterminal cell 18-30 pm}

_long

[23.4

± 4.6

_{pm], Subhymenium}

_up to about 60 pm thick,

composed

of branched and

strongly winding hyphae, partly provided

with

_anastomosing

arches.

_{Exciple 2-layered.}

Medulla_a thin

_layer

oftextura porrecta with about 2-4 pmwide,thin-walled

_hyphae.

Cortex

_consisting

oftextura

prismatica; hyphae

4-10 pm wide but

covering hyphae only

2-4 pm

wide,

thin-walled,

running parallel

withororientedatalow

angle

to the

excipular

surface,

notembedded ina

gelatinous

matrix; individual cells about 8-23 pm

long.

Occurrence—

Saprotrophic

_onstems of

Polygonum lapathifolium,

P.

_robustius,

and

Collection examined.GERMANY:Baden-Wilrttemberg, Schwabisch-Hall, Teurershof,MTB6824/3,in rush zoneofapond,ondead stemofPolygonum lapathifolium,23 July1986,L.Krieglsteiners.n.(herb.

Baral3057).

The

_{typical variety}

of H.

_fucatus

has been

_fully

redescribed and illustrated

_by

White

(1944: 609-613, figs. 25-30). According

tohim its ascimeasure 118-135x12-15 (tm

and its ascospores 24-34x 5-6.8 pm,ateach end beset withonetoseveral

_small,

incon-spicuous

'cilium-like

_processes'.

It is

_noteworthy

that these setulae havenotbeen mention-edor

depicted by

Dennis

_(1956:

79,

fig. 69G),

who also examinedan

isotype-collection.

Hymenoscyphus fucatus

has

_originally

been found and collected in

Shropshire, England,

ondeadstemsof

Polygonum lying

inwater.

A

_probable

record in 1923 in the Czech

Republik

canbe deduced from SvrcSek

_(1985:

159,

188, pi. IX, fig. 4). For,

in hisrevision of thetaxa described

_{by Velenovsky}

in the

genus

Helotium,

he statesthat the

lectotype

collection of Helotium macrosporumVelen.,

foundonstemsofCicerbita

alpina

in

Bohemia,

is identical with H. scutula

var. fucatus.

White

_{(1944: 610-613)}

collected it in both 1936 and 1938 in thesame

locality

in New York

_{State, USA,}

_onold stemsof

Polygonum

robustius

lying

inaswamp. Pallo collected the

species

in 1975_{on a}herbaceous

_dicotyledon

stemin the Western

_Caucasus,

Russia

(Vaasma

etal., 1986:

_26).

Baral

_(in

Baral &

_{Krieglsteiner,}

1985:

_{128) reported}

its occur-rencein 1975on stemsof

_Solidago

_{sp. in}

_{Baden-Wiirttemberg, Germany (no}

herbarium material

_preserved).

Blank has found H.

_fucatus

sensuBaral in 1987on astemof

Sola-numdulcamare in

Thayngen,

Switzerland

(not preserved;

Baral in

litt.).

Weber

(1992:

28, 122)

has also examinedaSwiss collection of

_it,

found

by

Baral & Blank in 1990on a

dicotyledon

stem in thecantonGraubiinden and determined

by

Baral

_(herb.

Baral

_4193).

Astothe last three records it hastobe

pointed

out,however,that H.

_fucatus

sensu Baral

differs from the_typein

lacking

arches attheascusbase

(Baral

in

_litt.;

Weber, 1992:

_121).

Another

_alleged

find was from

Germany

in 1989 on stems of Aruncus silvester in

Bavaria, but the

description

and

_figures

of this material show neither croziers orarches

norsetulae

(Engel,

1993:5,

8; Engel

&Hanff, 1993:

44).

A

_supposed

record in the Netherlands

_{(prov. Flevoland,}

Abbertbos,10 Oct. 1981;herb.

Swart-Velthuyzen

367)

has turnedout torepresent

typical

H. scutula.

1b.

_{Hymenoscyphus}

fucatus var. badensis

Hengstm.,

var. nov. —

Fig.

2

A varietate typicadiffert ascisminoribus,83-101 x9-10_pm,etascosporis etiampaulominoribus,

(23—)25—31(—35)pmlongis.

Apothecial morphology

—

Apothecia stipitate-cupulate,

0.4-1.5mm

high

when

re-hydrated,

scattered, erumpent

through

substratal

epidermis;

about 60 fruit-bodieson6cm

long fragment

ofaleaf.

Cupule

cup-to

saucer-shaped,

0.2-0.9mm in diameter when

rehydrated,

with

slightly

raised toflat

_{margin. Hymenium slightly}

concave toflat,

light

yellow

when dried.

Receptacle pale yellow

whendried,

subpruinose. Stipe cylindrical,

up to1.3mm

long

when

rehydrated,

aboutas

long

asor

longer

than the diameter of the

_disc,

0.1-0.2_mmacross,

pale yellow

when dried,

subpruinose.

Anatomy

— Asci

[k

= _1, n=

_{10] cylindric-clavate}

to

obconical-clavate,

83-101 pm

Hengstmengel:Notes onHymenoscyphus-II 197

remaining

visibleat mature

asci;

apex moreof less bullate and with thickened wall when immature,but

_slightly

truncated conical and with

_hardly

thickened wall when mature; annulus

_turning

medium blue in Melzer's_reagent.

_Ascospores

_[k

=

_1,

n=

_20]

obovoid-fusiformto

_{ellipsoid-fusiform, rarely subcylindrical, straight}

or

slightly

curved,

bilaterally

symmetrical, (23—)25—31 (—35)

pm

long [26.9

±1.9

_pm],

_{4-5 pm wide}

_[4.6

± 0.2

_pm],

with

_{length/width-ratio}

of

_{(5.1 —)5.3—7.0(—8.8) [5.8}

±

_0.5],

1-celled,

hyaline, obliquely

holotype: a.habit(x 20),b. asci(x 750),c.ascusapices in differentstages (x 500),d.ascospores (x 1500),e.paraphyses(x 750).

Hymenoscyphus scutuloides,holotype:a.habit(x 20),b. asci(x 750),c. ascus apices(stained withiodine)in different_{stages (x 1500),}d. ascospores(x 1500),e.paraphyses(x 750).

Hengstmengel: Notes onHymenoscyphus-II 199

biseriate, provided

with 1-2

_large

and

_possibly

one orafew small

guttules

when mature, afterwards with

_increasing

number

_(up

toabout

_eight)

of

_{shrinking guttules}

and

_finally

slightly granulose

to

optically

empty, thin- andsmooth-walled,atapex and base

frequent-ly (at

least about 50% of the

_{extremities) provided}

with 1-3

_tiny

setulae; apex bluntor

roundedto

_{abaxially angulate}

or sometimes

_laterally

beaked

_{(not hooked);}

base acute; setulaeat most_{2.5 pm}

_{long, extremely}

thin,

mostly

curved.

_Paraphyses

_[k

= _1, n=

_10]

cylindric-obconical,

about _{1.0 pm wide} nearthe base and upto _{1.5-2.0 pm wide}atthe

tip,

ratherscarce,

hyaline, according

toBaral

_{(in litt.)}

with

_highly

refractive

_{guttules (only}

when

_{fresh), provided}

with

_2-3(-4)

_septain the_{uppermost80 pm, sometimes forked}or

anastomosing

in thelower

half;

terminalcell

24-50(-60)

pm

long [39.6

± 9.1

pm],

1.0-2.8 timesas

long

as the subterminal cell

_[1.8

±

_0.5],

with rounded

_tip;

subterminal cell

14-31 pm

long [22.5

± 5.0

_{pm], Subhymenium}

above the

_stipe

uptoabout 25_pm

thick,

consisting

of 2-3 _pm

wide, strongly

branched and

_{winding hyphae, partly provided}

with arches.

_{Exciple 2-layered.}

_{Medulla up} toabout 10 pmthick, atexturaporrectawith 1-2

pmwide,thin-walled

_hyphae.

_{Cortex up}toabout 30 pm

thick,

atextura

prismatica; hyphae

about 5-10 pm wide and nearthe

edge

about 2-3 pm wide but

covering

hyphae only

about 1-2 pm

wide,

thin-walled,almost

_parallel

with the

_excipular

surface, notembedded ina

gelatinous matrix;

individual cells about 11-24pm

long.

Occurrence—

Saprotrophic

on

sedge-like leaf;

October.

Collection examined.GERMANY:Baden-Wiirttemberg, WeingartenerMoor (Oberrheinebene),MTB 6916-17,113 m,reed-land,onleaf of'

Carex’ _(accordingtofinder),1 Oct.1986,W. _Winterhoff86570

(holotype;herb. Baral).

The asci of this

_variety

are

significantly

smaller than those of thevar

fucatus.

More-over, the ascospores tendtobesmaller, and their average

length

is

significantly

smaller than that of the

_{typical variety.}

_{The difference in spore size is}even more

convincing

ifwe

lookat the dimensions of_{turgescentspores}as found

_{by Baral,}

viz.

_23-30(-33)

x

4.5-5.5 pm in the latter collectionversus

(28-)30-38(-40)

x_{5.5-7 pm in the collection of}

var

fucatus (Baral

in

_litt.).

2.

_{Hymenoscyphus}

scutuloides

_Hengstm.,

_spec. nov. —

Figs.

3, 4

?Hymenoscyphusscutula(Pers.: Fr.)W. Phill.sensuBreitenbach &Kranzlin,Pilze Schweiz 1 (1981) 170-171,proparte.

Apothecia stipitato-cupulata, erumpentia, stipite longitudinediametrum _cupulaecirciter_aequantivel paulosuperanti.Asci(80—)85 —102(—105)x8—9(—10) pm, inoperculati, octospori, ex uncisorti;apex plus minusve truncate conicus, annulo iodo medie caerulescente. Ascosporae irregulariter obovoideo-fusiformesut in Hymenoscyphoscutula, (18-)20-27x 3-4 pm, maturitate continuae, demum (uni-)

septatae,hyalinae, guttulatae,in asci parteinferiore uniseriatae,sursum obliquebiseriatae,parietibus tenuibus laevibusque,adapicem basemquevulgo 1—3(—4)setis filiformibus plerumque1-3pm longis

instructae;apex obtusus vel rotundatus usque lateraliterangulatusvelpaulouncatus, interdum satacutus; basisacutavel subacuta.Paraphyses cylindraceaevelcylindraceo-obconicae, longitudineascosaequantes, inferne 1.0-2.0_{pm, superne} 2.0-3.0_{pmlatae, septatae,}in dimidio inferiorealiquandofurcataevel ana-stomosantes._Excipulumbistratum. Medullaetexturaporrectaconstans. Cortexe texturaprismatica

con-stans,hyphis parallelisvel sub_anguloparvoadpaginam excipuli currentibus,ingelatinahaud inclusis,

parietibustenuibus vel_paulo incrassatis.

Apothecial morphology

—

Apothecia stipitate-cupulate,

upto2mm

high

when

rehy-drated,

concolorous

_{light yellow}

when

_dry,

_{scattered, erumpent}

_through

substratal

epider-mis;uptomore thanahundred fruit-bodiesover a

length

of 10cmof the substratalstem.

Cupule

cup- to

saucer-shaped,

upto1.5 mmin diameter when

rehydrated,

with

_slightly

raisedto

entirely plane margin. Hymenium

concave toflat.

Receptacle

smoothto

subpru-inose,

occasionally slightly radially

fibrous.

_{Stipe cylindrical}

toobconical, upto 1.5mm

long,

aboutas

long

as or

slightly longer

than the diameterofthe

_cupule,

upto0.3 mm

across, smooth to

_subpruinose,

atthe base sometimes surrounded

_by

asmall collaret of

epidermal

tissue.

Anatomy

— Asci

[k

= 1, n =

10]

obconicalto

_{cylindric-obconical,}

_(80-)85-102

(-105)

pm

long [96.0

± 4.9

_{pm], 8-9(-10)}

_{pm wide}

_[8.6

± 0.5

_pm],

with

length/width-ratio of 9.6-12.9

_[11.2

±

_{1.0], 8-spored, originating}

from croziers and the

_resulting

ana-stomosing

arches atthe base of the asci

_{generally remaining visible;}

apexmore orless

truncatedconical,thick-walled around the

pore;annulus

turning

medium blue in Melzer's

reagent,

especially

in the central

part.

Ascospores [k

=1,n=

_{20] bilaterally symmetrical}

(asymmetrical

in

_{side-view), obovoid-fusiform,}

flattenedononesideor

slightly curved,

(18-)20-27

pm

long [21.7

± 1.5

_{pm], 3-4pm}

wide

_{[3.6±0.3 pm],}

with

length/width-ratio of 5.1-7.7

_[6.1

±

_0.7],

_1-celled,

_a few

_{(older ones) 2-celled,}

_{hyaline, guttulate,}

in the lower_partof theascus uniseriate and

upwards passing

into

_obliquely

biseriate,

thin-walled,

smooth,

atapex and base

mostly provided

with

_1—3(—4)

setulae;apex bluntor

roundedto

oblique-angulate

or

slightly hooked, occasionally

rather acute; baseacuteor subacute; setulaefiliform,

1.0-3.0(-4.0)

pm

long,

sometimes also

_adhering

atthe flanks of the spore;

germination

observed in 2-celled spores,

laterally

from the

uppercell.

Para-physes [k

= 1, n =

_{10] cylindrical}

or

upwards slightly widening, equalling

the

_asci,

be-low _{1.0-2.0 pm and above 2.0-3.0 pm}

_{wide, provided}

with 2-4 _septain the_uppermost

100

pm,

occasionally

forkedor

anastomosing

in the lower

half,

somewhat

granulose,

partly staining

blue withcotton blue;terminal cell 20-58 pm

long [38.0

± 11.5

_pm],

0.7-4.9 timesas

long

asthe subterminal cell

_[2.3

±

1.3],

with rounded

_tip;

subterminal cell 11-35 pm

long [20.0

± 7.8

_{pm]. Subhymenium}

in the central_{part up}toabout 50 pm thick.

_{Exciple 2-layered.}

Medullanearthe

_stipe

_up to_{about 20 pm}thick,

composed

of

parallel, radially running,

2-4 pm

wide,

thin-walled

_{hyphae (textura porrecta).}

Cortex

a._young asci (x 750),b. _ascospores (x 1500).

Hymenoscyphus scutuloides (from Huijsman 55.H-99),

Hengstmengel:Notes onHymenoscyphus-II 201

about 60 pm

thick,

inclusive ofouter

covering layer,

atextura

prismatica; hyphae parallel

with_ororientedatalow

angle

tothe

_excipular

surface, notembedded ina

gelatinous

ma-trix,

with thinor

slightly

thickenedwalls

_(up

toabout 1.0

_pm),

_{5-9 pm wide but}

_covering

hyphae

thin-walled and

_only

_{about 3 pmwide; separate}cells about 5-30 pm

long.

Occurrence—

Saprotrophic

_onherbaceousstemsand_{on canes}of Rubus

sp.;

August-September.

Collectionsexamined. NETHERLANDS: prov.Gelderland, Winterswijk, Bekendelle,ondead herbaceous stem, 20Sept. 1953,R.A. Maas Geesteranus 9510 (holotype; L).— SWITZERLAND: cantonLuzern, Schiipfheim, ondead caneof Rubus _sp.,21 Aug. 1955,H. S. C.Huijsman55.H-99(L).

An indication of the

_shape

and colour of the

_apothecia

in fresh condition is

given

in

notes

accompanying

the Swiss

_collection,

which state:

[apothecia]

'young

deeply

cup-shaped,

lateron more

flat,

but

_long

time

_{remaining cup-shaped,}

_lastly

flat; colour very

light

cream-lemon;disc

_slightly

darker than rest.'

The_type

_{collection, originally}

identified_as ‘Helotium scutula

_(Pers.

ex

Fr.)

Karst.',

has been examined

by

Dr. K.P. Dumont

(New

York Botanical

_Garden)

in March 1981,

but_wasnotannotated

by

him.

As

_{already expressed by}

itsname,H. scutuloides showsagreatresemblancetoH. scu-tula. The latter

_species,

however, has

_larger

asci

_(120-142

_x 9-11

_pm)

which donot

originate

from

_croziers,

_{while its spores} are

slightly

broader

(4-5 pm)

and

usually

pos-sess

only

onesetulaateach end. In view of the forementioned resemblance it is

quite

possible

that H. scutula in thesenseofsomeauthors includes H. scutuloidesas well.

Thisseems tobe thecase in Breitenbach & Kranzlin

(1981:

170-171,No.

190).

Their

description

and

_figures

of H. scutula are

mainly

basedon acollection, foundon

herba-ceous stemsin the Swisscanton Luzern

_(compare

examined

_collection!)

and

_probably

representing

H. scutuloides. The presence of

anastomosing

arches at the base of the asci is notmentioned,but

suggested by

their

_fig.

190B.

_Perhaps

H. scutuloides also has been found

_by

Berthet

_(1964:

40-41,

101)

ondeadstems of

_Solidago

canadensis in

France,for the relevant collection of 'H. scutula'

is describedtobe of the

pleurorhynchous

type.

A related

_species,

also

_resembling

H. scutula and with asci saidtobe

_produced

from

'tiny'

croziers, has been described

_by

Dumont &

Carpenter (1982: 582-587, figs.

5,

6)

under thenameH. ‘affin. scutulus’. This

_species,

however, foundon various substrates

in the

_neotropics,

has

obviously pigmented paraphyses

and

_{covering hyphae,}

while its

spores are

only

2—3(—4)

pm

wide,

short-setuloseatthe baseand

shaped

like thoseof H. serotinus.

3.

_{Hymenoscyphus}

menthae

_{(W. Phill.)}

Baral—

Fig.

5

Helotium menthae W.Phill.,Elv. brit. (4) (1881)n. 188 [invalid: nomennudum];W.Phill. in

W. Phill.&Plowr.,Grevillea 10(1881)69.(basionym).—Hymenoscyphascutulavar.menthae(W. Phill.)

W. Phill.,Man. Brit.Discomyc. (1887) 137. — Phialea scutula _var. menthae (W. Phill.) Sacc., Syll. Fung. 8 (1889)266. —Helotium scutula var.menthae(W. Phill.)Rehm in Rabenh. Krypt.-Fl. ed. 2,

1 (3) (1893)793. —Helotium scutula forma menthae(W. Phill.) Massee,

Brit._Fung.-fl. 4(1895)254.

—Hymenoscyphusmenthae (W. Phill.)Baralin Baral & Krieglsteiner,

Beih. Z. Mykol.6(1985) 131 [bibliographicerrorof citation ofbasionym].

Fig.5.Hymenoscyphusmenthae(fromMaas Geesteranus 9046),a.habit (x 20),b. asci(x 750),c.ascus

Hengstmengel:Notes onHymenoscyphus-II 203

Apothecial morphology

—

Apothecia stipitate-cupulate,

variable insize,

upto 6mm

high

when

_rehydrated,

concolorous

light

ochraceous

_yellow

when

_{dry, scattered,}

gre-garious

or clustered,

rarely

mutually

grown

together along

whole

_length

of the

_stipe,

erumpent

through

substratal

_epidermis

or

superficial

ondecorticated_{parts, scanty}to_very

numerous

(up

to_morethan_a thousand

_specimens

over a

length

of 10cmof the substratal

stem). Cupule

cup- to

_{saucer-shaped,}

_uptomore than 1.5 mmin diameter when

rehy-drated,when young withmore or less raised

_{margin. Hymenium}

concave to

flat,

young

orange-yellow

when fresh.

_Receptacle

concolorous with

hymenium,

smoothto

subprui-nose.

Stipe cylindrical

to

obconical,

uptoabout 5 x0.6mm when

rehydrated,

aboutas

long

asor

(much) longer

than the diameter of the

cupule,

more orless

pruinose,

at the base

_occasionally

surrounded

_by

asmall,often dark-coloured collaret of

_epidermal

tissue.

Anatomy

—Asci

[k

=

1,

n=

_{10] (cylindric-)obconical}

to

cylindric-clavate,

(72-)76-90

pm

long [82.8

± 5.1

pm],

7-9 pm wide[8.3 ± 0.5

jurn],

with

length/width-ratio

of

(8.4—)9.1—11.1 (—11.6) [10.0

±

_{0.7], 8-spored,}

_originating

from croziers of which the arches remain visibleatthe base of theasci; apex more orless truncated

conical,

thick-walled around the pore; annulus

turning

blue in Melzer's_reagent,

_especially

in the middle

part.

Ascospores [k

=_{1, n}=

_{20] axially}

to

_{bilaterally symmetrical (in}

the lattercase asym-metrical in

side-view), fusiform-ellipsoidal, ellipsoidal,

obovoidalor

ellipsoid-fusiform

to

obovoid-fusiform, straight

to

_{slightly curved,}

_{14-21 pm}

_{long [16.5}

± 1.7

_{pm], 3-4(-5)}

pm wide

[3.7

± 0.2

_pm],

with

_{length/width-ratio}

of

_{(3.4—)3.6—5.5(—5.9)}

_[4.4

±

_0.5J,

1-celled, only

afew

_{2-celled, hyaline, guttulate, obliquely biseriate,}

thin-walled, smooth,

without obvious setulae but

_{occasionally apiculate}

at_apexorbase;apexbluntorrounded

to

_{oblique-angulate,}

_occasionally

rather acute; base blunttosubacute.

_{Paraphyses [k}

= 1, n =

_10]

obconical,

equalling

theasci, below 1.0-2.0 pm and above 2.0-3.0 pm wide,

provided

with

_(1—)2—4

_septain the_{uppermost80 pm, often with}one ortwofurcations in the lower

_half;

terminal cell

_{(26—)31—50(—65)}

_pm

_{long [41.3}

± 6.2

_pm],

1.2-2.5 timesas

long

asthe subterminal cell[1.7 ±

_0.4],

with rounded

tip;

subterminal cell _{18-31 pm}

_long

[24.3

± 3.7

pm]. Subhymenium

in the central_{part up}toabout 65 pm thick.

Exciple

2-layered.

Medullanot

sharply

defined from

_subhymenium

and cortex,nearthe

_stipe

about 35-50

pm

thick, consisting

oftextura porrectawith 2-4 pm

wide,

thin-walled

_hyphac.

Cortex about 40-60 pm

thick,

atextura

_prismatica,

without

_clearly

differentiated

_covering

layer; hyphae

4-13 pm wide,

parallel

with ororientedatalow

_{angle (at}

most

45°)

tothe

excipular surface,

notembedded in a

gelatinous

matrix,with thinor

slightly

thickened

walls;

separatecells about 8-40 pm

long.

,

Occurrence—

Saprotrophic

onstems of

_{Polygonum cuspidatum}

and other herbs and

on canesof Rubus sp.;

September-October.

Collections examined. NETHERLANDS: _prov. Drente, Ruinen,WijkenvanEleveld,ondead caneof Rubus _sp„2 Oct.1983,L. Jalink & M.M.Nauta 229 (WBS);prov.Utrecht, Baarn, Lage-Vuursche,on

dead herbaceous stems, 1_Sept. 1957,J.Daams306 (L);prov.Zuid-Holland, Warmond,estate Huyste

Warmont',on dead stems of_{Polygonum cuspidatum.} 24_Sept. 1952,R.A. Maas Geesteranus 9046 (L).

With the

_{understanding}

that in relevant literature

nothing

is said about the presenceor

absence of

_croziers,

the above

_description

_{agrees well with the}

_{original description by}

Phillips (in Phillips

&

_Plowright,

1881:

₆₉₎

and with the

_description

and

_figures

ofan

Cooke,

is

'evidently

the

typecollection' but this seems inconsistent with the fact that

Helotium menthae is

_formally

basedonElvellacei britannici 188. At thesame

place

Dennis

has_putHelotium

_{(Hymenoscypha)}

scutulavar. menthae

'(Phill.)

Boud.' _{into the}

syn-onymy of Helotium scutula ‘var. solani Karst. ...

1870'.6 I doubt whether this is

justi-fied.

_Firstly,

the asci ofvar. menthaeareupto90 pm

long,

whereas theascus ofvar.

solani,

as

depicted by

Dennis

_{(1956: fig. 71B)}

from material in herb.Karsten, is more

than _{110 pm}

_{long. Secondly,}

the annulus of_var. menthae

_always

turnsblue in

iodine,

whereasvar.solani has 'thecae ...

apice

iodonon tinctae'

_(Karsten,

1870:

_234).

After-wards Dennis has

_possibly

abandoned the _{forementioned synonymy,}

_for,

in his

re-arrangementof the genus

Hymenoscyphus

(Dennis,

1964:

_73-78)

he does mention H. scutulavar. menthae whereasvar.

solani,

whose varietal

_epithet

has

_priority

in case

of synonymy, has been omitted. In my

opinion

var. solanisensuKarsten is

_quite

similar to, ifnot identical with H. consobrinus

(Boud.) Hengstm.,

also because of its

'fusoid-elongate'

spores whichare

generally uniseptate according

toDennis

_{(1956: fig. 71B).}

Assuming

that alloftheBritish collectionsmentioned

_by

Dennis

_{(1956: 78)}

represent

'var. menthae’sensu

stricto,

then thistaxon has been recorded in Great Britainonstems of Mentha sativa

_[=

M. x

verticillata],

Teucrium

scorodonia,

Solanum

tuberosum,

Cam-panula

latifolia

and

_Polygonum

sp.

(cf. Dennis,

1978:

136).

The recordon Solarium

tuberosum_seemsalsotohave been referredtoin Ellis & Ellis

_{(1985: 425-426, pi. 161,}

fig.

1672).

Kirk &

_{Spooner (1984:}

₅₃₂₎

have

_reported

on two

findings

of H. ‘scutulus’

var. solani in 1980onUrtica dioica and unidentified herbaceousstemsonArran, Scot-land. Dr. B.M.

_{Spooner (in litt.)}

has

kindly

informed

me,that 'the

interpretation

followed in the Arranaccount is thatofDennis

_(1956),

as

figured (fig. 71B)

from Karsten's mate-rial'

_[of

var. solani

s.str.!].

He added,that the Arran collection

_'may

differ from var.

solaniasdescribed

by

Dennis

_{[p. 78;}

i.e._var. menthae

s.str.!]

in

_having

_arather

white-tomentose

stipe

base

_{[characteristic}

of H.

consobrinus,

as

already

mentioned

_by

Dennis

(1956: 79)]

and in

_being

on Urtica.’ All in

all,

atleast the Arran collectionseemsto show

moreresemblancetoH. consobrinusthantoH. menthaeasdescribed in this article. Baral

_(in

Baral &

_{Krieglsteiner,}

1985:

_{131-132) reported}

several

findings

of H.

men-thae in

_{Baden-Wiirttemberg, Germany,}

viz._on

_Polygonum

_cuspidatum

and

_?piperatum,

Scrophularia

nodosa,

Lysimachia vulgaris, Lycopus

europaeus and Rubus idaeus.

Strange-ly enough, according

toBaral

_{(in litt.)}

H. menthae

_always

hasawhite

_hymenium,

where-as

Phillips

(1887:

137) speaks

ofa

bright yellow

disc. In 1914 and 1917 '

Helotium scutulavar.menthae’wasfound

_by

P.

_Vogel

in Mark

_Brandenburg,

_Germany,

on stems of Mentha

_piperita

and has been distributed within twoGerman exsiccata

_series,

viz.

Sydow's Mycotheca germanica (as

No.

₁₃₅₀₎

and

_Vogel's

Flora der Mark

_{(s. n.)}

respec-tively.

However,the examinedtwo

_copies

of each of these exsiccata

_(L)

all_representvar.

scutula.

_Exactly

thesame

misapplication

occursin Petrak's Flora BohemiaeetMoraviae

exsiccata, II.

Serie,

I.

_{Abteilung, Lfg.}

5, Nr. 243, collected

by

F. Petrak in 1911 on

G Helotium scutula _subspec. [!]solani P. Karst.,Symb. Mycol.fenn. [1] (1870)234 =Helotium

scu-tula var.solani (P. Karst.)P. Karst.,Mycol.fenn. 1 ₍₁₈₇₁₎ 111 =Helotium scutula forma solani (P. Karst.)Rehmin Rabenh.,Krypt.-Fl.ed.2,1(3) (1893)793[invalid:unintentional stat. nov.;only (erroneous)citation of Karsten's 'f. Solani’ from1871]=_{Hymenoscyphus}scutulavar.solani (P. Karst.)

Hengstmengel: Notes onHymenoscyphus-II 205

Mentha

_{longifolia (examined specimen: L;}

cf.

_{Samuels, 1985:46).}

It is

_evident,

that

_Vogel

and Petrak

_wrongly

used the substrate asanessential

distinguishing

feature. Likewise Oudemans

_{(1890: 315)}

at first

_thought

todeal with H. scutula var. menthae when he

examineda

Hymenoscyphus

found in 1889on stemsof Mentha

_aquatica

in the botanical

garden

of Amsterdam. Examination of authentic

_specimens

ofvar.

menthae,

however,

gave him

certainty

that the

_fungus

of

_Phillips

differed from hisone, not

only by

the

ab-sence of cilium-like

_appendages,

but also

_by

the size and

_shape

_{of the spores and the}

quantity

of

_guttules.

From the Netherlands

_only

the three indicated collections could be ascribedtoH. men-thae. Yet I examined several collections labelledas

Helotium/Hymenoscyphus

scutula

var. menthaeor var. solani

(herb. Swart-Velthuyzen 210,

357;Lexherb. Emste

_937/82,

949/82),

but these all

_belong

to H.

_{consobrinus, just}

liketwo

_Belgian

collections

origi-nally

determined_asHelotium scutula_var. menthae

_{(herb. Swart-Velthuyzen 272;}

herb. Batten

₈₃₉₎

anda

Belgian

collectionof'‘Helotiumscutula f. solani’

_'

(BRcoll. V. Mouton).

Outside

_{(Western) Europe}

H. menthae

_{probably only}

has been recorded under the

name

'

H. scutulavar. solani’ in thesenseof Dennis

(1956: 78).

Ahmad

_{(1978: 207-208)}

has collected thistaxon in 1953 and 1959 in Pakistan. Thind & Sharma

_(1980:

128-129,

figs. 3, 4)

found it

_{'growing luxuriantly}

_on

_Polygonum

stems

[i.a.

P.

_{amplexicaule]}

in the North-Western

_Himalayas',

India. Korf collected it in 1981onunidentifiedstemsand

on

Polygonum

cuspidatum

inSichuan,China

_(Korf

&

_Zhuang,

1985:

_{500). According}

to Lizon

_{(1992: 45), however,}

the latter collection_represents

_{(the typical variety of)}

H. scu-tula.Furthermore, Thind & Sharma

_(1980:

₁₂₉₎

mentioned theoccurrence of H. ‘scutula

var. solani’ in i.a. North

_America,

but without

_giving

_{any reference}

_although

this taxon hasnotbeen listed

by

Farretal.

(1989).

It neednot be said that

_only

acareful re-examination of relevant collectionscan

give

more

certainty

about the realoccurrenceand distributionofthis littleknown

_species.

ACKNOWLEDGEMENTS

Theauthor is indebted toMr. H.O. Baral_{(Tubingen, Germany),}Dr. E. Batten(Eefde, Netherlands), Mrs. C.M._{Swart-Velthuyzen}(Bennekom, Netherlands)and the curators of BRand WBS for the loan of relevant collections,andtoDr. B.M._Spooner(Kew, United Kingdom)for_sendinginformation onhis interpretationof H. scutulavar.solani andonPhillips' Elvellacei britannici,fasc. 3. He also wishes to

thankDr. R.A. Maas Geesteranusforchecking the Latindiagnosisand description,and Mr. J.J.A.M. Wessendorp,whopreparedthefiguresforprinting.

REFERENCES

Ahmad,S.1978.Ascomycetesof Pakistan. Part 1. Lahore. (Biol.Soc.Pakistan,Monograph7.) Arendholz,W.R.1979.Morphologisch-taxonomische Untersuchungenanblattbewohnenden Ascomyceten

ausderOrdnungder Helotiales.Hamburg.

Arx,J. A. von.1967. Pilzkunde. Ein kurzerAbrissder_Mykologieunterbesonderer_{Beriicksichtigung} der Pilzein Reinkultur. Lehre.

Baral,H.O. & G.J.Krieglsteiner.1985. BausteinezueinerAskomyzeten-Florader_{Bundesrepublik} Deutsch-land: In Suddeutschland _{gefundene Inoperculate Discomyzeten}mittaxonomischen,okologischen, chorologischenHinweisen und einer Farbtafel.Beih. Z._Mykol.6: 1-160.

Boidin,J. 1971. Nuclear behavior in themyceliumand the evolution of theBasidiomycetes.In: R.H.

Peter-sen(ed.),Evolution in thehigher Basidiomycetes,An international symposium: 129-148. Knoxville. Breitenbach,J. & F. Kranzlin. 1981. Pilzeder Schweiz. Beitr.Kenntn. Pilzfl. Schweiz. Band 1:

Asco-myceten(Schlauchpilze).Luzern.

Chadefaud,M. 1943. Sur les divers typesd'elements_dangeardiensdes_{Ascomycfetes,}etsurlaformation

des_asqueschez la Pdzize Pustularia catinus.Rev. scient.

81:

77-80.

Dennis,R. W.G.1956.Arevision of the British Helotiaceae in the herbariumof the_RoyalBotanic Gar-dens, Kew,with notes onrelated European species. Mycol. Papers62.

Dennis,R.W.G.1964. Remarksonthe_genus HymenoscyphusS.F.Gray,withobservations on sundry speciesreferred _bySaccardo and othersto the_generaHelotium,Pezizella orPhialea. Persoonia 3: 29-80.

Dennis,R.W.G.1978. British_Ascomycetes. Revised edition.Vaduz. Dorfelt,H.(ed.). 1989. Lexikon der_{Mykologie. Stuttgart}and New York.

Dumont,K.P. 1976. Sclerotiniaceae XI.On Moellerodiscus (=Ciboriopsis). Mycologia68:233-267. Dumont,K.P. 1981. Leotiaceae III. Notes onselected temperatespeciesreferred toHelotium and

Hyme-noscyphus. Mycotaxon13: 59-84.

Dumont,K.P. & S.E.Carpenter. 1982. Los hongosde Colombia VII. Leotiaceae IV. _{Hymenoscyphus} caudatus and relatedspeciesfrom Colombia andadjacent regions.Caldasia 13: 567-602.

Ellis,M.B. &J.P. Ellis. 1985. Microfungionland plants,An identification handbook. London and Sydney.

Engel,H. 1993. Pilzneufunde in Nordwestoberfranken 1989,I.Teil. Auflistung, Kurzbeschreibungen, Hinweise. Pilzfl. Nordwestoberfrankens 14/15: 3-24.

Engel,H.& B. Hanff. 1993. Pilzneufunde in Nordwestoberfranken 1989/1990,II. Teil.Ascomyceten. Pilzfl. Nordwestoberfrankens 14/15: 39-50.

Farr, D.F.,G.F.Bills,G.P. Chamuris & A.Y.Rossman. 1989.Fungionplantsand_{plant products}in the United States.St.Paul,Minnesota.

Gaumann,E. A. & C. W.Dodge. 1928.Comparative morphologyof_fungi.NewYorkand London. Haines,J.H. 1989. Studiesin theHyaloscyphaceaeIV:Fuscolachnum,a newgenus forDasyscyphus

pte-ridis. Mem. New York Bot. Garden49: 315-325.

Hawksworth, D.L. 1987. Theevolution and adaptationof sexual reproductivestructures in the Asco-mycotina.In: A.D.M._Rayner, C.M. Brasier & D. Moore(eds.),Evolutionary biologyof thefungi:

179-189._Cambridgeetc.

Hawksworth, D.L.,B.C. Sutton & G.C. Ainsworth. 1983. Ainsworth &_Bisby'sdictionaryof the _fungi Seventhedition.Kew,Surrey.

Huhtinen,S. 1990. A monographofHyaloscyphaandallied_genera. Karstenia 29: 45-252.

Karsten, P. A. 1870. Symbolaead Mycologiamfennicam [1]. Notis. Sallsk. FaunaFl. fenn. Forh. 11: 211-268.

Kirk, P.M. & B.M.Spooner. 1984. Anaccountof theFungiofArran,Gighaand Kintyre.Kew Bull.38: 503-597.

Korf,R.P. & P. Lizofi. 1994. Lambertellinia scutuloides (Sclerotiniaceae),a new_genus andspeciesfora

Discomycete previouslyconfusedwith_{Hymenoscyphus}caudatus._Mycotaxon50: 167-174. Korf,R.P. & W.-Y.Zhuang.1985. Somenewspeciesand_newrecords of_DiscomycetesinChina.

Myco-taxon 22: 483-514.

Lizon,P. 1992. The_genus Hymenoscyphus(Helotiales)in Slovakia,Czechoslovakia. _Mycotaxon45: 11-59.

Martens,P. 1932.L'originedu «crochet» etde l'«anse d'anastomose» chez lesChampignons superieurs. Bull. Soc. mycol.Fr. 48: 259-281.

Moreau,F. 1950. Le_myceliumsecondaire chez lesAscomycetes.Bull. Soc.mycol.Fr. 66:114-118. Moreau,F. 1954. Les_Champignons.Tome 2. Paris.

Moreau,F.& Mme. 1922. Le_mycdliumaboucles chez les_Ascomycetes. C.R. Acad.Sci., Paris, 174: 1072-1074.

Oudemans,C.A.J.A.1890._Micromycetesnouveaux.Versl. Meded. Kon. Akad.Wetensch.,Afd.

Hengstmengel: Notes on Hymenoscyphus-11 207

Phillips, W. 1887. A manual of the British_{Discomycetes.} London.

Phillips,W. & C.B._Plowright. 1881. NewandrareBritishfungi.Grevillea 10: 65-74.

Samuels,G.J. 1985. An annotated indextothe_{mycological writings}of Franz Petrak. Vol.4,H-L. Wel-lington.

Svrfiek, M. 1985. Ataxonomic revision of_{inoperculate Discomycetes} described _byJ. Velenovsky in the_genus Helotium,preservedin National _{Museum, Prague.}Sb. ndr. Muz.Praze40 B: 129-215 ('1984').

Thind,K.S. & M.P. Sharma. 1980. The_genus HymenoscyphusS.F. Grayin India. Nova Hedwigia32: 121-132.

Vaasma, M„K.Kalamees & A. Raitviir. 1986.Macrofungi of the Caucasian State Nature Reserve. Tal-linn.

Weber,E. 1992.UntersuchungenzuFortpflanzungund Ploidie verschiedener Ascomyceten.Berlin and Stuttgart.Biblthca _mycol. 140.

White,W.L. 1943. Studiesin the _genus Helotium,III._Historyand _diagnosisof certain_Europeanand NorthAmerican foliicolous _species.Farlowia 1: 135-170.