Primed in Grear Brirain

TAXONOMIC REVISION OF SEPTORIA-LIKE FUNGI

PARASITIC ON BETULACEAE
By 0 . CONSTANTINESCU
Institute of Systematic Botany, P.O. Box 541, S-751 21 Uppsala, Sweden

As a result of a rather broad generic concept, and of a basically morphological approach to

species delimitation, 6 species of Septaria are accepted on Betulaceae: S . alni, S . alnijolia, S .
betulae, $ , betulae-odoratae, S . ostryae and S. weiriana . These are fully described and
illustrated and a key is provided. Cercoseptoria septoriopsidis and Phloeospore//a borealis
comb.nov., are proposed and a list of 12 related, doubtful and excluded taxa is appended.
This study deals with a group of coelomycetous
fungi which produce leaf-spot diseases of Betu-
laceae (including Carpinaceae and Corylaceae).
They are biotrophic on leaves, occasionally on
catkins . The damage caused to the hosts is gene-
rally minor, but sometimes the attack results in a
severe and premature defoliation, particularly de-
structive to the seedlings in tree nurseries. Besides
the fungi originally described under Septaria, a
search to uncover species placed in other genera
such as Cylindrosporium and Phloeospora was also
made.
MATERIAL AND METHODS
Some 371 herbarium specimens, including the
types of 23 out of 27 taxa dealt in this paper were
examined. Leaf fragments bearing conidiomata
were wetted with 70 % ethyl alcohol, rehydrated
with distilled water, and 8-12 p,m thick sections
were cut with the aid of a freezing microtome.
Squash preparations of the conidiomata were also
made for most of the specimens. Sections and
conidiomata were routinely mounted in cotton blue
in lactic acid. On occasion, other stains such as 3 %
erythrosin in 10 % NH 40H (Su tton, 1980), Azur A
(Shoem aker, 1964), Chlorazol Black (Arm itage,
1944) and laetofuchsin (Carmichael, 1955) were
used. The slides were examined in transmitted
light. Phase- and interference-contrast microscopy
were used in critical cases to reveal the annellations
of the conidiogenous cells. The figures given for the
lengths of conidia represent the maximum distance
between the ends. In most cases, due to the
conidium curvature, this is an underestimate of the
real length. The widths of both conidia and coni-
diogenous cells were measured at their widest part.
Herbarium abbreviations are those used by Holm-
gren, Keuken & Schofield (1981 ). In some cases
only selected specimens examined are cited, but a
complete list is deposited with UPS herbarium.
GENUS DELIMITATION
The demarcation between Septaria Sacco and re-
lated genera is not clear (Teterevnikova-Babayan,
1961; Jorstad, 1962, 1965; Sutton, 1980; von Arx,
1981, 1983). The characters mostly used in dis-
tinguishing these fungi are the conidiomatal type,
conidiogenesis, and the conidial shape, width and
septation. Septoria is most closely related to Phloeo-
spora Wallr., and these two genera are either treated
as separate taxa (Sutton , 1980), or considered
congeneric (Jorstad, 1961, 1965; von Arx, 1981 ,
1983).
Conidiogenesis, extensively used as a taxonomic
character in recent studies on anamorphic fungi,
apparently cannot elucidate the problem of delimi-
ting these taxa. Our study confirmed previous
reports, mentioned by Sutton (1980), that Septaria
may show different types of conidiogenesis. Unfor-
tunately, in many specimens it was difficult to
detect by light microscopy the annellidic condition
of the conidiogenous cells. Phase- and interference-
contrast microscopy, as well as different stains
used, did not always clearly reveal the annellations,
which in some specimens are at the limit of light
microscope resolution.
Although fungi with different modes of coni-
diogenesis have been commonly placed in separate
genera, this attitude seems to change. Recently,
Samuels & Hallett (1983) consider conidiogenesis
type as an infrageneric character in Microdochium
and mention four coelomycetous genera having
both sympodulae and annellides, whereas Zhang,
Kendrick & Brubacher (1983) use conidiogenesis
only to distinguish subspecific taxa. Moreover, as
shown by Minter, Kirk & Sutton (1982) and Deigh-
ton (1983), some fungi exhibit intermediate stages
between holoblastic sympodial arid annellidic
mechanisms of conidiogenesis. These apparently
contradictory features are also present in some
Septoria-like fungi parasitic on Betulaceae. Thus,
384 Septoria spp . on Betulaceae
S . alni and S . betulae are typical Septaria sens u
Sutton (1980), S . alnifolia, S . ostryae and S.
weiriana differ mainly by the presence of both
sympodulae and annellides, whereas S . betula e-
odoratae fits Phloeospora, except that the coni-
diomata are pycnidial. Moreover, no teleomorph is
known in these six species, and their taxonomy
depends exclusively on the anamorph characters.
Until the delimitation of Septaria from related
Coelomycetes becomes more precise it is preferable
to maintain in Septoria those fungi havingpycnidial
conidiomata, conidiogenous cells sympod u lae
and/or annellides, holoblastic conidial ontogeny,
and hyaline scolecosporous, septate conidia pro-
vided with unthickened scars.
ported by cross-inoculation experiments which
proved that M. betulinum can infect both Alnus and
Betula (R oll- H an sen & Roll-Hansen, 1981 ). The
recently described M elampsoridium asiaticum
Kaneko & Hiratsuka (1983) parasitizes both Car-
pinus and Ostrya.
No pathogenicity test was carried out for this
study and only morphologically distinguishable
units (sp ecies) at the host family level are recog-
nized, irrespective of their presumed host speci-
ficity, As Sutton (1980) pointed out in the case of
Ascochyta , ' a taxonomy based on ho st families
rather than host genera or species may provide the
circumscription b y which some order may evolve
out of the present apparent confusion ' .

SEPTORIA ALNI Sacc., Michelia 1: 177 (1878).

SPECIES CONCEPT
The supposed affinity to a certain host genus
or species has been the main character used in
delimiting Septaria species. As a result, morpho-
logically indistinguishable fungi were given specific
rank solely because they have been found on hosts
not previously reported. Long ago Beach (19 19)
questioned this approach of Septaria taxonomy. In
inoculation experiments with 15 Septaria species
parasitic on plants from various families he found
that some ha ve a host range limited to a few closely
related host species, whereas in other Septorias two
or even three related host genera can be infected .
Relatively recent studies on Betulaceae-parasitic
fungi show that more than one host genus can
harbour the same fungus. Thus French, Hodges &
Froyd (1969) consider that strains of Hypoxylon
mammatum (W ah l.) Mill., from alder and birch are
indistinguishable. According to Smerlis (1969),
strains of Godronia cassandrae Peck from Alnus and
Betula are not host-specific at the generic level. By
detailed studies of urediniospore morphology,
Kaneko & Hiratsuka (19 81) showed that M elamp-
soridium betulinum Kleb . is parasitic on Alnus,
Betula and Ostrya, and M. carp ini (F uckel) Dietel
on Carpinus and Ostrya. They concluded that the
host genus is not reliable for differentiating these
Melampsoridium species and their results are sup-
(Fig . 1)
Rhabdospora alni (Sacc.) Kuntze, Rev. Gen. Pl.
3(3 ) : 509 (1898) .
S. carisolensis Kabat & Bubak, Ost. bot. Z. 55: 184
(190 5) .
Leaf spots brown to reddish brown, irregular,
vein-limited, 0 '5-5 mm diam, up to 14 mm when
coalesced, margin not distinct, or narrow and some-
what darker; reverse paler. Conidiomata mostly
hypophyllous, appear as black pustules, scattered,
pycnidial, subcuticular or more or less immersed,
separate, subglobose when young but often acer-
vular-like when mature, yellowish to pale brown,
dehiscence by pore or irregular, 60-110 Jlm diam
and 60-80 lim high ; wall 10-15 Jlm thick, textura
angularis. Conidiogenous cells sympodulae, hyaline,
lageniformorampulliform, 5-12 x 2'5-5 '5 usn.Con-
idia hyaline, obclavate, more or less curved, (1- )3- 4
(-7-9)- sep tat e, truncate at base, tapered to a
subacute apex, (25-)33- 55(-65) x 1-2(-2 '5 ) Jlm.
Tel eomorph unknown.
Hosts and distribution: On Alnus firma Siebold &
Zucc. , A. glutinosa (L. ) Gaertner, A. hirsuta
Turcz. , A . incana (L.) Moench, A. [aponica
(T h u n b .) Steudel, A . pendula Matsum., A . siebol-
diana Matsum., and A. tenuifolia Nutt., in Eurasia
and N . America.

KEY TO THE SPECIES

1. Conidia mostly 3-3 '5 flm w ide, one-septate. . , . . , . . . S . betulae-odoratae
1 . Conidia mostly les s than 3 I'm wide, with 3 or more septa . 2
2 . Conidia predominantly 3-septate . . . . . . . . . .... · 3
2. Conidia predominantly wi th more than 3 sep ta .. . . . . 4
3 . Conidia usually over 60 I'm long . , . S . weiriana
3. Conidia usually under 50 I'm long . . S .ostryae
4 . Conidia alm ost cylindrical , mostly more than 21,m wide, with rounded apex . S . alnifolia
4. Conidia obclavate, mostly I-21,m wide, apex usually subacute, . . . . . . 5
5, Conidia on average 33-55 x 1-2I'ffi, parasitic on Alnus . , . . , . . . S . alni
5, Conidia on average 40-70 x f ' 5-2 I'm, parasitic on Betula and C arp inus . . . . . . . . , . . S. betulae
 O. Constantinescu

Fig. 1. Septoria alni. (A ) From the holotype ; (B) from the holotype of S . carisolensis; (C) from TFM 2767;
(D) from BUCM 32501.
386 Septaria spp. on Betulaceae
S elected specimens examined: on Alnus firma, Japan ,
Meguro, Tokyo, 30 July 1952, K . It o & T . Koba yashi,
TFM 2768 ; Suzuka, Mie Pref. , 30 Nov . 1949, K. It o,
TFM 0152 ; Tawara-mura, Kawachi-gun, Tochigi Pref.,
3 1 Oct. 1950, K. Shibukawa, TFM 2955; Kamabuchi,
Yamagata Pref., 28 Sept. 1948, K . It o, TFM 0020. On
Alnus glutinosa, Czechoslovakia, Miihr.-Weisskirchen ,
Hrabuvka, 1 Sept . 1914, F. Petrak, Flora Bohem. et
Morav . exs. 1309, BR, K, S, WIS; Italy , Conegliano,
June 1876, C. Spegazzini, holotype of S. alni, PAD ;
Carisolo, Val Rendena, 28 July 1904, E. Kabat, holotype
of S. carisolensis, BPI; Tortona, autumn 1875, Passerini,
Thiim., Myc, univ. 693, S, UPS; S. Teresa, near Papiam ,
Cavara, Fungi Longob. exs. 190, K, S; Poland, Stani slan,
Bystrzyca Solottwinska, 8 Oct. 1917, F . Petrak, Fungi
polonici exs. 210, K, S ; Romania, Distr. Nearnt, Varatec ,
4 Sep t. 1952, C. Sandu-ViIle, BUCM 32501; USSR,
Latvia, ProvoVidzeme, Madona, Vestiena , 25 Aug. 1931,
K. Stares, BUCM 32502, K, LE, S; USA, Wis., Dane
Co ., Madison, 4 Sept. 1944, H . C. Greene 824, WIS. On
Alnus hirsuta, Japan, Koma, Iwate Pref., 23 Aug. 1950,
H. Chiba, TFM 2949; Meguro, Tokyo, 30 July 1952,
K. Ito & T. Kobayashi, TFM 2767; ditto, 11 Oct. 1950,
K. Shibukawa, TFM 2769. On Alnus incana, Canada,
N.S., Kings Co., Casey's Corner, 3 Sept . 1952,
D . B. O. Savile, WIS; USA, Wis., Arcadia , 5 Sept. 1917,
J . J. Davis, WIS; Madison, 4 Sept. 1944, H . C. Greene
766 and 842, WIS. On Alnus japonica, Japan, Meguro,
Tokyo, 30 Aug. 1952, K. Ito & T . Kobayashi, TFM
2771 ; Nakahata, Shirakawa-gun, Fukushima Pref.,
15 Sept. 1954, K. Ito, TFM 0796. On Alnus pendula,
Japan, Meguro, Tokyo, 30 July 1952, K . Ito &
T . Kobayashi, TFM 2780. On Alnus sieboldiana, Japan,
Ibaraki Pref., Kushigata, Taga-gun, 23 June 1951,
Y. Hosaka, TFM 2954. On Alnus tenuifolia, USA, Mont.,
Lincoln Co., 24 July 1924, J . Dearness, WIS .
An economically important leaf-spot d isease pro-
duced by S. alni on different A/nus species was
reported from Japan by Ito & Shibukawa ( 1956).
The disease has resulted in foliage losses of seed-
lings in tree nurseries, but it was effectively con-
trolled (Sato, Ora & Shoji, 1955).
The supposedly S. alni-Mycosphaerella alni-
viridis (d e Not.) Tomilin connexion was not con-
firmed in inoculation experiments (I to &
Shibukawa, 1956).
SEPTORIA ALNIFOLIA Ell. & Ev., Proc . Acad. nat.
Sci. Phi/ad., Part 3: 366 (1894). (F ig . 2)
Rhabdospora alnifolia (Ell. & Ev. ) Kuntze, Rev.
Gen . Pl. 3(3): 509 (1898) .
Leaf spots ochreous to pale brown, circular to
irregular, 1-20 mm diam or covering large areas
when coalesced, margin narrow, brown to vio-
laceous brown, somewhat raised ; reverse paler.
Conidiomata epiphyllous, rarely amphigenous,
appear as yellowish to dark pustules, usually agglo-
merated in the spot centre, sometimes with circular
disposition, rarely scattered, p ycnidial, first irnmer-
sed, later erumpent, more or less globose, yellowish
to pale brown, dehiscence irregular, 60-150 Jlm
diam; wall 10-20 JIm thick, textura angularis . Con-
idiogenous cells sympodulae, rarely annellides, hya-
line, lageniform, 10-18 x 3-6 tun. Conidia hyaline,
cylindric to obclavate, more or less curved, (2-)3-6
d
(-9)- sep t ate , base truncate, apex rounded,
(3 2-)3 7- 56(- 74 ) x ( 1' 5)2- 2 ' 5(- 3) usn , Teleomorph
unknown.
Hosts and distribution: On Alnus rhombifolia
Nutt., and A. rubra Nutt., in N. America.
Selected specimens examined: On Alnus rhombifolia,
USA, Cal. , San Diego Co., Cuyamaca, 12 Nov . 1934,
O . A. Plunkett, CUP 56856, NY ; Pasadena, Aug . 1894,
McClatchie, NY ; San Bernardino, 28 Nov . 1920,
M . F. Barrus, CUP 45581; Santa Clara Co., Stanford
Univ., 1 Nov. 1902, coll . E. B. Copeland, det. Earle ,
C. F . Baker , Pacific Slope Fungi 1849, K, NY, PC, S,
UPS, WIS; Del None Co ., Smith River , Sept . 1934,
H. E. Parks 5184 (H , S, WIS two spec.). On Alnus rubra,
USA, Cal., Monterey Co., Big Sur, 14 Aug. 1937,
L. Bonar, Calif. Fungi 563, CUP, H, K, NY two spec. , S,
UPS, W, WIS ;Wash. ,Seattle,Aug.1893,C. V. Piper 82,
holotype of S. alnifolia, NYS; isotype, NY; ditto, Aug .
1894, C. V. Piper Ell. & Ev., N. Amer. Fungi 3367, S,
UPS, WIS ; Port Orchard, 19 Aug. 1912, E. Bartho-
lomew, Fungi Columbo4476, CUP, K, WIS; Challam Co .,
near Forks, 3 July 1926, J . S. Boyce, Sydow, Fungi exot,
exs. 745, S, WIS.
SEPTORIA BETULAE Pass., Comment. Soc . crittog . ital.
2: 441 ( 186 7), non (Lib .) Westend. ex Sacco
(1878) . (F ig. 3 )
S . betulina Pass., Atti Soc. crittog: ital. 2 : 40
(1879) ·
Rhabdospora betulina (P ass.) Kuntze, Rev. Gen .
Pl. 3(3 ) : 510 ( 189 8) .
S . betulico/a Peck, Rep. N. Y. St. Mus. nat. Hist.
34: 44 ( 1883), non Lobik ( 1928) .
Rhabdospora betulicola (Peck) Kuntze, Rev. Gen .
Pl. 3(3) : 510 ( 18 9 8).
S. betu/icola Peck val'. margina/is Peck, Rep.
N . Y. St. Mus . nat. Hist. 34: 44 (1883).
S . carpinea J. Davis, Trans. Wis. Acad. Sci. Arts
Leu. 18 : 88 (19 15).
Leaf spots brown to reddish brown, rarely with
ochreous centre, polyangular to irregular, vein-
limited, 0 ' 5-4 mm diam, up to 9 mm when coa-
lesced, margin narrow, darker; reverse paler.
Conidiomata mostly hypophyllous, appear as brown
to black pustules, scattered all over the spots,
pycnidial, more or less immersed, globose to subglo-
bose, 50-160 pm diam, osteolate or dehiscence
irregular ; wall 10-15 pm thick, textura angularis.
Conidiogenous cells sympodulae, hyaline, lageni-
form to ampulliform, 7-18 x 3-7 psix. Conidia hya-
line, obclavate to acicular, more or less curved,
(1- )3- 5(- 8)- sep tat e, base truncate, apex subacute
 O. Constantinescu

Fig. 2. Septaria alnifolia. (A) From the holotype; (B) from Ell. & Ev., N. Am. Fungi 3367 in UPS; (C) from
USA, H. E. Parks 5184 in S.

to rounded, (27-)40-70(-86) x (1-)1'5-2(-3) I"m.
Teleomorph unknown.
Hosts and distribution: On Betula lenta L., B. lutea
Michx, B. papyrifera Marsh., B. pendula Rothm.,
B. populifolia Marsh., B. pubescens Ehrh., B. pumila
L. and Carpinus caroliniana Walt. in Eurasia and N.
America.
Selected specimens examined: On Betula lema, USA,
Mass., Arnold arboretum, Jamaica Plain, 24 Aug. 1892,
J. G. Jack & A. B. Seymour, WIS; ditto in Seymour &
Earle, Econ, Fungi 169, BUCM, CUP, K, NY two spec.
On Betula lutea, Canada, N.S., Kings Co., White Rock, 9
Sept. 1952, D. Creelman & I. V. Hall, WIS. Ont.,
Algoma, 27 Aug. 1906, J. Dearness, E. Bartholomew,
Fungi Columbo 2375, CUP, K, NY, S, WIS; USA, N.Y.,
 Septoria spp . on Betulaceae

I lO/lm

c
Fig. 3. Septoria betulae. (A) From the isotype in UPS; (B) from the holotype of S . betulicola; (C) from the
holotype of S. carpinea .
 O. Constantinescu
Catskill Mts., Aug., C. H . Peck, holotype of S. betulicola
Peck, NYS ; dino, holotype of S. betulicola Peck var .
marginalis Peck, NYS ; Wis., Neopit, J . J. Davis, 14 July
1914, WIS. On Betula papyrifera, Canada, Man.,
Brandon, 29 Aug. 1944, W. L. Gordon 2709 , DAOM
152125 ; N .S., Diamond Lake, Temagami, 1 Sept. 1935,
H. S. Jackson, BPI; Ont., London, Komoka swamp, 20
Aug. 1889, J. Dearness, NY; Whimey Lake, T. F . R., 30
July 1931, coil. S. M. Pady, det . H . S. Jackson, DAOM
82046 ; USA, Mass., Jamaica Plain, Arnold Arboretum,
24 Aug. 1892, Seymour & Earle, Econ. Fungi 525, BUCM,
CUP, K, NY two spec ., S, WIS two spec.; N .Y., Lake
Pleasant, Aug ., C. H. Peck, NYS, WIS ; W is., Dane ce .,
Madison, 10 Aug. 1943, H. C. Greene 472, WIS ; Three
Lakes, 15 June 1892, J. J. Davis 9214, NY. On Betula
pendula, Czechoslovakia, Miihren, Usti near Miihr.-
Weisskirchen, Aug . 1940, F . Petrak, Myc. gen. 776, S,
UPS ; Germany, Westfalen, near Siegen, 11 Sept. 1921,
A. Ludwig, Sydow, Myc. germ. 1975, UPS; Zahlendorf
near Berlin, Sept. 1888, P. Sydow, Myc. march. 2378, S;
Italy, Parma, Collechio, Sept. 1873, Passerini, Thum.,
Herb. myc . oee. 242, isotype of S. betulae Pass., UPS;
Sweden, Vasterborten, Umea, Sept. 1909, J . Vleugel, S;
USSR, Kursk, Vishinovo, 25 Aug. 1907, LE. On Betula
populifolia, Canada, N.S., Yarmouth, Co., Pleasant Lake,
11 Sept. 1952, D . B. O. Savile, WIS; USA, Kan., Man-
hattan, Agric. College Nursery, July 1887, W. T.
Swingle, NY ; dino in Ell. & Ev., N . Amer. Fungi 2166,
UPS, WIS ; Wis ., Madison, 4 July 1957 , H . C . Greene
2088, WIS. On Betula pubescens, Austria, Siidtirol, St
Peter, Vilnostahl, 4 Aug . 1905, J. E . Kabat, Kabat &
Bubak, Fungi imperf, exs. 416, S, W ; Norway, Halse &
Hakrmark, Ro, 29 July 1964, H. B. Gjaerum, Herb. Inst.
PI. Prot., As ; Sweden, Norrboten, Karlsvik near Lulei,
Sept. 1912, J. Vleugel, UPS; Vastergotland, Oxnered,
near Vanersborg, 27 Sept. 1905, A. G . Eliassen, S, UPS ;
Skane, N. Mellby parish, Mellby mosse, 11 Aug . 1982,
O. Constantinescu & E. Gunnerbeck 4205, UPS; Viist-
manland, Sura parish, Finnmossen, 21 Jul y 1983,
E. Gunnerbeck 4513 , UPS; USSR, West Siberia, Omsk,
Aug. 1926, K . E. Murashkinsky, Petrak, Myc.gen. 1279,
UPS, W. On Betula pumila, USA, Wis., Gaslyn, 4 Aug.
1911, WIS. On Carpinus caroliniana , USA, Wi s., Gaslyn,
8 Aug. 1911 , J . J . Davis, holotype of S . carpinea, WIS.
Septaria betulae is a variable species, particularly
regarding the size of conidia and number of septa.
In some collections most of the conidia are short
(35-50 p,m), narrow (1'5 p,m) and 3-septate,
whereas in others they are longer (55-70 p,m), wider
(2-2 '5/lm) and 5-septate. These characteristics
could not be correlated with the host range or
distribution area and, moreover, material from
different hosts forms an overlapping series. Con-
sequently, I prefer to maintain all these specimens
in a single, variable species. Some collections of S .
betulae can hardly be distinguished from S . alni and
it is tempting to consider these two taxa conspecific.
However, the conidia of S . alni are shorter, nar-
rower, have fewer septa and are far less variable
than those of S . betulae, Moreover, S . betulae is
frequently associated with a spermatial Asteromella
state. The morphology and physiology of this
fungus (under S . chinensis) was studied by Ono
(196 1), who also proved experimentally that it can
parasitize different Betula species.
In most of the European herbarium specimens
and literature records Septoria on Betula is labelled
S. betulae (Lib.) Westend., a nomen nudum which
cannot take precedence over the later but validly
published homonym S. betulae Pass. Septoria betu-
licola Pass., proposed to replace S. betulae Pass., is
thus superfluous. For the taxonomic and nomen-
clatural status of S. betulae (Lib.) Westend. ex
Sacc., see the list of related, doubtful and excluded
taxa.
Two American collections preserved in herb.
WIS, labelled S. betulicola Peck (on Betula lema,
N .Y ., Petersburg, Aug., colI. C. H. Peck, and on B.
lutea, Wisc., White Lake, 20 Aug. 1921, coll.
J. J. Davis), apparently belong to S. ostryae. It is
not impossible that S. ostryae can attack Betula, but
such a hypothesis should be supported by more
similar specimens.
SEPTORIA BETULAE-ODORATAE Bubak & Vleugel,
Svensk bot. Tidskr. S: 348 (1911). (Fig. 4)
Cylindrosporium utahense Solheim, Univ. Wyo .
Pubis 36 : 62 ( 1970) .
Leaf spots ochreous to pale reddish brown,
irregular to more or less round, 1-10 mm diam,
margin narrow, darker; reverse paler. Conidiomara
epiphyllous, rarely amphigenous, appear as
brown to blackish pustules, usually agglomerated,
pycnidial, immersed, becoming erumpent, sub-
globose to almost cupulate, pale brown,
115-350 x 100-200 p,m, dehiscence irregular; wall
10-15 p,m thick, textura angularis. Conidiogenous
cells annellides, rarely sympodulae, hyaline, lageni-
form to ampulliform, 10-21 x 3'5--6 p,m. Conidia
hyaline, cylindrical, more or less curved, 1(-3)-
septate, base truncate, apex rounded,
(32-)40-55(- 78) x (2' 5-)3- 3'5(- 4) p,m. Teleomorph
unknown.
Hosts and distribution: On Betula occidentalis
Hook, B. papyrifera Marsh., and B. pubescens Ehrh.
var. carpatica (Willd.) Ascherson & Graebner in N.
America and Northern Europe.
Specimens examined: On Betula occidentalis, USA,
Wash., Waitsburg, Aug. 1900, R. M. Horner,
E. Bartholomew, Ell . & Ev ., Fungi Columbo 1586, CUP,
K, NY, S, WIS two spec.; Mont., Darby, July 1916,
J . R. Weir, NY ; Ut., Cache Co., East of Smithfield, 23
Aug . 1961, W . G . & R. Solheim, 6106, Solheim, Mycoft.
Saximont. exs. 1375, holotype of Cylindrosporium utahense,
RMS 42897 ; isotypes RMS 287624 and UPS; along
Logan River, below Wood Camp Hollow, Cache National
Forest, 13 July 1974, C. T. Rogerson, NY. On Betula
papynfera, Canada, Ont., Bear Island, Lake Temagami,
390 Septoria spp. on Betulaceae

I IOllm I

9
A

Fig. 4. Septoria betulae-odoratae, (A) From the holotype; (B) from the holotype of Cylindrosporium utahense ;
(C) from Canada, R. F. Cain 1263 in NY.
 O. Constantinescu
23 June 1931, coli. G. E. Thompson, WIS ; 1 Jul y 1931,
R. F. Cain, 1263, NY ; USA, Wis ., Racine , 4 Oct. 1893,
J. J . Davis , WIS. On Betula pubescens var. carpatica,
Sweden, Vasterbonen, Umea, Sept. 1909, J. Vleugel,
holotype of S. betulae-odoratae, BPI; isotype S.
S eptoria-odoratae has characters of both
Phloeospora sensu Sutton (1980) and S eptoria, The
annellidic condition of most of the conidiogenous
cells and the cylindrical, mostly one-septate conidia
with truncate base suggest a close affinity with
Phloeospora, but the thin-walled pycnidial conidio-
mata are typical for Septaria .
SEPTORlA OSTRYAE Peck, Rep. N . Y . St. Mus . nat.
H ist . 33 : 25 (1883) · (Fig. 5)
Rhabdospora ostryae (P eck) Kuntze, Rev. Gen .
Pl. 3(3) : 512 (189 8).
S. corylina Peck, Rep. N. Y. St. Mus. nat . Hist.
34 : 44 (188 3).
Rhabdospora cory/ina (Peck) Kuntze, Rev. Gen.
Pl. 3(3 ): 510 (189 8).
G/oeosporium rostratum Ell. & Ev., J . Myco/. 3:
129 (1887).
Cylindrosporium dearnessii Ell. & Ev., Proc. Acad.
nat. Sci. Phi/ad.: 83 (1891 ).
Phloeospora dearnessii (Ell. & Ev.) Hohnel, Ann/s
myco/ . 22 : 200 ( 1924).
Leaf spots epiphyllous, ochreous to pale brown,
round to irregular, 0 '5-12 mm diam, up to 30 mm
when coalescing, margin narrow, reddish to vio-
laceous brown; reverse paler. Conidiomata epi-
phyllous appear as yellowish to brown pustules,
usually agglomerated in the spot centre, pycnidial,
immersed, separate or gregarious, globose to
cupulate, yellowish to pale brown, dehiscence
irregular, 50-10 flm diam, sometimes up to 160 flm
wide; wall circa 10 flm thick, textura angularis.
Conidiogenous cells sympodulae, sometimes annel-
lides, hyaline, lageniform, rarely ampulliform,
6-14 x 3-7 flm . Conidia holoblastic, hyaline,
obclavate, curved, (1-)3(- 7)- sep tat e, truncate at
base, tapered to a subacute apex, (26-)35- 50
(-60) x (2-)2 '5- 2 ' 8(- 3 '5) usn. Te/eomorph un-
known.
Hosts and distribution: On Carpinus caro/iniana
Walt., Cory/us americana Walt., C. cornuta Marsh.
and Ostrya virginiana (Mill.) Koch in N. America.
Selected specimens examined: On Carpinus caroliniana,
Canada, Ont., London, July 1890, J. Dearness, Ell. & Ev.,
N . Amer. Fungi 2664, isotype of Cylindrosporium dear-
nessii, UPS. On Corylus americana : Canada, Ont.,
London, Sept. 1889, J. Dearness, Ell . & Everh ., N . Amer.
Fungi 2858, NY, UPS; USA, Mich., Saginaw, 4 Sept.
1889, M . B. Waite, NY; Neb., Nebraska City,June 1889,
H. J. Weber 53, NY; N .Y., Green ce ., Albany, 7 Aug.
1948; Wis., Dane Co ., Madison, 19 Oct . 1942; Uni v.
Wise. Arboretum, 19 Aug 1943, all in WIS, collected by
H. C. Greene. On Corylus cornuta var. califo rnica,
39 1
Canada, B.C. , May 1887, J . Macoun, holotype of Gloeo-
sporium rostratum, NY; Ont., London, July 1893,
J . Dearness, Ell . & Ev., Fungi Columbo366, CUP, K , NY ,
WIS ; USA, Ca!., Del Norte Co., Darlingtonia, Smith
River , June 1937, H . E. & S. T. Parks 6011, Calif. Fungi
717, BR, CUP, H , NY, S, W, WIS; Humbold Co. ,
Lawrence Creek, 6 July 1934, H . E. Parks & J . P . Tracy
11101, Calif. Fungi 875, S; Berkeley, 12 June 1906,
C. O. Smith,E. Bartholomew,FungiColumb.2376,CUP,
K, S, WIS; Mass ., Granville, 4 Aug. 1885, A. B.
Seymour, Seymour & Earle, Econ. Fungi 175, BUCM,
CUP, K ; N . J., Powerville, Sept. 1892, F. L. Stevens,
Ell. & Ev., Fungi Columbo 438, CUP, K, WIS two spec. ;
N.Y., Millerton, June 1880, C. H . Peck, holotype of
Septoria corylina, NYS ; isotype, WIS ; Ore., Portland, 23
Aug. 1915, E . Bartholomew,Fungi Columb o4772, CUP, S,
WIS; Lane Co., Rujada, 17 June 1923, J. S. Boyce,
Sydow , Fungi exot . exs. 747, S, UPS, WIS; Wash., Kitsap
Co., Bainbridge Is!., 26 Aug . 1909, E. Bartholomew,
Fungi Columbo 3186, CUP, S, WIS. On Corylus comura
var. comuta, Canada, N .S., Kings Co., Black River, 24
June 1953, D . Creelman, WIS; USA, N.Y., Essex Co. , St
Huberts, 17 June 1958, C. T. Rogerson 28102, NY,
NYS; Wis., Madison, Univ. Wise. Arboretum, 6 Aug.
1960, H. C. Greene 2290, WIS; Sawyer Co., Flambeau
State Forest, near Oxbow, 4 Sept. 1963, H. C. Greene,
WIS. On Ostrya v irginiana, Canada, Ont., near London,
20 Aug. 1910, J . Dearness, E . Bartholomew, Fungi
Columbo 3485, BPI , CUP, K, S, WIS; ditto, 24 Aug. 1910,
Kabat & Bubak, Fungi imperf, exs. 821, BPI, K , S two
spec., W ; Que. , West of Danford Lake, 4 July 1947,
D. B. O. Savile, S; USA, N .Y., Helderberg Mts, Jul y,
C. H . Peck, holotype of Septoria ostryae, NYS; Wis.,
Dane Co., Madison, 6 Aug . 1947, H. C. Greene, BPI, S,
WIS; Kenosha Co., 23 Aug. 1933, WIS; Madison, 20
Aug. 1893, WIS ; Racine, 7 July 1892, BPI, NY , WIS ;
Shieton, 15 Aug. 1917, WIS, all collected by J . J. Dav is.
In some specimens on Ostrya, S. ostryae
produces atypical symptoms, i.e, brown, often
coalescing spots, located at the leaf margin. On
these spots the conidiomata appear as hypo-
phyllous, scattered, dark brown to blackish pus-
tules. This form of attack sometimes occurs on the
same leaf with the common type. On Carpinus and
Ostrya, S. ostryae is frequently associated with an
Asteromella, and on Cory/us, less often, with
Monostichella cory/i (D esm .) V. Hohn., sensu von
Arx (1970).
SEPTaRIA WEIRIANA Sacc., Nuovo G. bot. ital., N .S.
27: 83 (1920). (Fig. 6)
S. boycei Dearn., Mycologia 20: 236 (1928) .
Leaf spots ochreous, pale 6rown to reddish brown,
somewhat raised and coriaceous, irregular, often
vein-limited, 1-8 mm diam but covering large areas
when coalesced, margin reddish to violaceous
brown, up to 0 '5 mm wide ; reverse paler with no
distinct margin. No spots are produced on catkins.
Conidiomata epiphyllous, appear as black, scattered
pustules, pycnidial, immersed, separate, more or
less globose, dehiscence irregular, 90-140 flm
39 2 Septaria spp. on Betulaceae

IOllm

:'
c
@ ,,'

Fig. 5. Septoria ostryae. (A) From the holotype; (B) from the holotype of S. corylina; (C) from USA, Big
Hollow, 13 July 1893, C. L. Shear in NY; (D) from the isotype of Cylindrosporium dearnessii in UPS.
 o. Constantinescu 393

, IDO I'm I

Fig. 6. Septoria weiriana. (A) From the ho!otype ; (B) from the holotype of S . boycei.
394 Septoria spp. on Betulaceae
diam; wall dark brown, ca 20 Jlm thick, textura
angularis; on catkins conidiomata appear as black,
raised pustules, pycnidial, more or less superficial,
separate or aggregate, globose, 160-320 Jlm diam,
wall dark brown, 20-50/lm thick, outer layer tex-
rura globosa, inner layer textura angularis. Conidio-
genous cells sympodulae, sometimes annellides,
hyaline, lageniform, rarely ampulliforrn,
8-12 x 3'5-6 psn. Conidia hyaline, acicular to
almost obclavate, more or less curved or sigmoid,
(1-)3(-4)-septate, truncate at base, tapered to an
acute apex, (40-)60-75(-95) x 2-2'5(-3) Jlm.
Teleomorph unknown.
Hosts and distribution: On Alnus crispa (Aiton)
Pursh, A. tenuijolia Nutt., and Betula occidentalis
Hook, in N. America.
Specimens examined: On Alnus crispa, Canada, B.C.,
Mara, 5 Aug. 1964, colI. J. Grant, det. D. B. O. Savile,
DAOM 107718; Que., Great Whale River, 5 Aug. 1944.
D. B. O. Savile603,DAOM75099;H, WIS;USA, Wis.,
Brule, 10 Aug. 1924; Hayevard, 27 Aug. 1924; Herbster,
11 Aug. 1923; Minocqua, 21 Aug. 1919; ditto, 6 July
1910; Vilas Co., 25 July 1902, all in WIS, coli. by
J. J. Davis. On Alnus tenuifolia, USA, Ida., St Maries, 13
May 1919, J. R. Weir 10160, det. Saccardo, holotype of
S. weiriana, PAD; isotype, BPI; Wash., S.E. of North-
port, 30 Aug. 1928, G. G. Hedgcock, CUP 46482, UPS,
WIS; Metalline Falls, 28 Aug. 1934, colI. G. G.
Hedgcock, CUP 464481, UPS, WIS, On Betula occi-
dentalis, USA, Ore., Clackamas Co., 8 July 1924, coli.
J. S. Boyce, det. J. Dearness 5670, holotype of S. boycei,
DAOM; isotype BPI.
Septoria weiriana was originally described as
parasitic on catkins of Alnus tenuifolia and it was
only known from the type collection. During our
study numerous specimens of a similar but leaf-
parasitic fungus, labelled S. alnijolia and inhabiting
Alnus tenuifolia and A. crispa, became available.
Although the fungus parasitic on leaves has smaller
pycnidia, with thinner walls and smaller conidia, its
morphology is essentially the same as in the catkins'
form. The differences are most probably due to the
fact that the latter is parasitic on a substratum with
physiologically more active tissues than the leaves.
According to Ehrlich (1942), the leaf-spot disease
produced by this fungus (under S. alnifolia) was
abundant during the summer of 1942 in Central
Idaho, USA.
RELATED, DOUBTFUL AND EXCLUDED SPECIES
Cercoseptoria septoriopsidis (Dearn. &
Overh.) comb.nov. (Fig. 7A)
Septoria septoriopsidis Dearn. & Overh., Myco-
logia 20: 238 (1928).
Leaf spots reddish brown, irregular, 5-20 mm
diam, margin effuse, yellowish; reverse the same.
Conidiomata epiphyllous, agglomerated in the spot
centre, pseudostromatic, lower part immersed and
yellowish, upper part hyaline, emerging through
the leaf tissue or stomata, up to 55 Jlm wide.
Conidiogenous cells on short conidiophores or arising
from the upper part of the pseudostrorna, syrn-
podulae, hyaline, lageniform, 7-22 x 2-4 Jlm.
Conidia hyaline, acicular to obclavate, slightly
curved, 3-5-septate, base truncate, apex rounded to
subacute, 25-60 x 2-2'5 psn.
Specimen examined: On Betula lema L., USA., Pa,
Center Co., Snow Shoe, 28 July 1924, L. O. Overholts &
R. S. Kirby 5705, holotype, DAOM.
Except for the hyaline conidia, this fungus fits
Cercoseptoria Petrak sensu Deighton (1976). Never-
theless, species with hyaline conidia have been
included in Cercoseptoria by von Arx (1983).
Already Dearness & Overholts (loc. cit.) have stated
that Septoria septoriopsidis resembles Cercoseptoria.
Their decision to place it in Septoria was apparently
due to the pycnidia of an Asteromella, very abun-
dant in the type specimen, and which they con-
sidered part of the same fungus.
CYLINDROSPORIUM BETULAE J. Davis, Trans. Wis.
Acad. Sci. Arts Lett. 18: 99 (1915). (Fig. 8B)
Leaf spots brown to reddish brown, scattered,
irregular, vein-limited, 0'5-3'5 mm diam, margin
somewhat darker; reverse paler. Conidiomata hypo-
phyllous, hardly visible, acervular, subcuticular,
45-190/lm wide, hyaline, thin-walled, textura
angularis; dehiscence irregular. Conidiogenous cells
phialides, hyaline, lageniform, 10-17 x 2-3 /lm.
Conidia hyaline, cylindrical, more or less curved
(o-jr-septate, base and apex rounded,
30-50 x r- 5-2 /lm.
Specimen examined: On Betula pumila L., USA, Wis.,
Jefferson Co., Sullivan, 17 July 1906, J. J. Davis, holo-
type, WIS.
The septate conidia and the absence of coni-
diophores exclude this fungus from Cylindro-
sporium. It could not be accommodated in any of
the Phialostromatineae genera treated by Sutton
(1980). According to Martin (1929), C. betulae
caused heavy defoliation of Betula sp. in Delaware,
USA.
CYLINDROSPORIUM OSTRYAE Woronikhin, Trudy
bot. Mus. Acad. Nauk SSSR (21): 203 (1927).
This species was described on Ostrya carpinifolia
Scop., from USSR. No type or other material was
available at LE and LEP herbaria. The host and the
description suggest Septaria ostryae Peck, and the
author himself considered it close to Cylindro-
sporium dearnessii Ell. & Ev., which is conspecific
 O. Constantinescu 395

IO/.lffi

Fig. 7. (A) Cercoseptoria septoriopsidis from the holotype; (B) Asteroma alneum from the isorype of Septoria
a/nico/a in UPS; (C) Phloeosporella borealis from the holotype,

with S. ostryae. However, S. ostryae seems to be
restricted to N. America.
Phloeosporella borealis (Lind & Vleugel) comb.
nov. (Fig. 7C)
Phloeospora borealis Lind & Vleugel, Svensk bot.
Tidskr. 2: 383 (1908). (as 'Phleospora')
Leaf spots reddish to chocolate brown, irregular,
1-6 mm diam; reverse paler. Conidiomata hypo-
phyllous, gregarious, acervular, becoming erum-
pent, hyaline to pale brown, textura angularis,
90-180 pm wide, 35-60 pm high. Conidiogenous
cells sympodulae, hyaline, cylindrical to lageni-
form, 6-15 x 2-3 pm. Conidia hyaline, cylindrical,
straight or slightly curved, (1-)3(-7)-septate, base
truncate, apex rounded, 17-52 x 2' 5-3 pm. Teleo-
morph unknown.
Specimen examined: On A/nus irn:ana (L.) Moench
subsp. ko/aensis (Orlova) A. & D. Love, Sweden, Viister-
botten, Urnea, Oct. 1906,]. Vleugel, holotype, S.
 Septoria spp. on Betulaceae
The presence of sympodulae and multiseptate
conidia place this fungus in Phloeosporella ,
SEPTORIA ALNICOLA Cooke, J. Bot., L ond. 4: 114
(1886) . (F ig. 7B)
Rhabdospora alnicola (C ooke) Kuntze, R ev. Gen.
Pl. 3(3) : 509 (1898) .
Leaf spot ochreous to pale brown, irregular,
2-8 mm diam, up to 20 mm when coalesced, margin
not defined or narrow, brown. C onidiomata appear
as brown to blackish superficial pustules, agglo-
merated or scattered, acervular, subepidermal,
lenticular. Con idiogenous cells phialides, hyaline,
lageniform, 6-10 x 2-4 pm. Conidia hyaline, cylin-
drical to fusiform, sometimes slightly curved, con-
tinuous, 9-13 x 2'5-3'5 p m .
Specimens exam ined: On Alnus glutinosa (L. ) Gaertner,
France, Seine et Marne, Fontainebleau, Oct. 1883, coil.
Feuilleaubois, 577, Rourneguere, Fungi gall. exs. 3274,
UPS ; place and date not indicated, coil. Fautrey, UPS;
Germany, Berlin, Hippodrom, Oct. 1887, P. Sydow,
Syd., Myc. march. 1758, UPS; UK: Surrey, Shere, Oct.
1866, M . C . Cooke, Fungi brito exs. 203, isotypes,
UPS,W.
Septoria alnicola Cooke is conspecific with
Asteroma aln eum (P ers.: Fr.) Sutton.
SEPTORIA ALNIGENA Sacc., Michelia 1: 179 (1878) .
Rhabdospora alnigena (Sacc .) Kuntze, R ev . Gen .
Pl. 3 (3) : 509 (1898) .
This species was described on Alnus glutinosa (L. )
Gaertner, from Italy. No type material is available
at PAD and TO herbaria. According to Saccardo's
description, S. alnigena has h yaline, continuous,
20 x 0'75 pm conidia, which suggest A steroma
rather than S eptaria.
SEPTORIA AVELLANAE Berk. & Broome, Rabenh.,
Fungi eur. 1958 (1875) Also de scribed in Ann.
Mag. nat. Hist ., Ser 4,17: 141 (F eb . 1876 ) and
Grevillea s: 56 (D ec. 1876).
Rhabdospora avellanae (Ber k. & Broome) Kuntze,
Rev . Gen. Pl. 3(3): 510 ( 189 8).
Specimens examined: On Corylus avellana L ., Germany,
Oberammergau, road to Ma11estein, Aug. 1891, Allescher,
Allescher & Schnabl, Fungi bavarici 181, S two spec.,
WIS; Westfalen, Distr. Meschede, Deutrneke, 24 Aug.
1924, A. Ludwig, Sydow, Myc. germ. 2400 , LE, S, WIS;
ditto, Weringhausen, 3 Sept. 1924, A . Ludwig, Wirtgen,
Herb. PI. crit., select. hybrid, Fl. Rhenanae, ed . nov ., 11
Abt., 74, LE ; UK, Bathford, Nov. 1874, C. E . Broome,
Rabenh., Fung i eur. 1958, isotype, LE two spec ., UPS ;
Goosehill Wood, Hanbury, Worcestersh ire , 28 Oct. 1927 ,
W . B. Grove & P . M. Rhodes, 3302, LE.
Septaria avellanae is conspecific with Asteroma
coryli (F uckel) Sutton and it is welI illustrated by
Sutton ( 1980) . It produces a common leaf-spot
disease of Cory Ius in Europe and Asia. Severe
attacks were recorded by Ribaldi (1953) in Italy
(u nd er Cylindrothyrium cory li (F uckel) R ibaldi)
and by Akhundov (1958, 1961 ) in Azerbajdzhan
SSR (und er Cylindrosporium cory li Ibragimov &
Akhundov, and the invalid name Cylindrosporium
av ellaneum (Ber k. & Broome) Akhundov &
Ibragimov).
SEPTORIA BETULAE (Lib .) Westend. ex Sacc. ,
Michelia 1 : 177 (1878). (F ig. 8A)
Ascochyta betulae Lib., apud Westend., L es
Cryptogames classees d' apres leurs stations
naturelles : 156 ( 1854), nom. nud.
S. betulae (L ib. ) Westend., I.e., nom. nud.
Rhabdospora betulae (Lib .) Kuntze, R ev . Gen . Pl.
3(3 ) : 5 10 (1898) as ' «Lib.) Westend.)' , nom.
nud.
Leaf spots pale yelIowish to ochreous, more or less
circular, 1-2'5 mm diam. Conidiomata hypo-
phylIous, appear as minute reddish spots, scattered,
pycnidial, immersed, globose, yelIowish, osteolate,
40-130 pst: diam; wall ca 5 pm thick, textura angu-
laris. Conidiogenous cells phialides, hyaline, lageni-
form or ampulliform, 6-14 x 2-3 ' 5 usx». Conidia
hyaline, falcate, aseptate, provided with a chromo-
philic spherical body just below the middle,
30-40 x (1- ) 1' 5(- 2) lIm . Teleomorph unknown.
Spe cimen examined : On Betula pendula Rothm. , Italy,
Conegliano, Oct. 1876, coli . C . Spegazzini, authentic for
the name S. betulae (L ib. ) Westend. ex Sacc., PAD .
Ascochya betulae Libert was apparently never
published. Westendorp made the combination in
Septoria but produced no description. No voucher
specimen seems to exist for these two names .
Saccardo's description was based on a different
specimen preserved now at PAD. This specimen is
here described and depicted. The aseptate conidia
exclude this fungus from Septaria. I could not find
a coelomycetous genus to accommodate it but
refrain from introducing a new generic name till
more similar fungi become available.
SEPTORIA BETULICOLA Lobik, Bolez. Rast. 17 : 179
(1928).
Described on Betula pubescens Ehrh., from USSR.
This is a later homonym of S. betulicola Peck. No
type material was available at LE and LEP her-
baria. The description suggests S. betulae-odoratae,
which also parasitizes this plant.
SEPTORIA CHINENSIS Miura, Flora ofManchuria and
East Mongolia, III Cryptogams, Fungi : 447
(19 2 8).
Described on Betula chinensis Max., from N .E.
China. The type of this species could not be located
at PE, TAl and TFM herbaria. The description
agrees with S . betulae Pass., except for the conidia,
 O. Constantinescu 397

20/lm
IO/lm I L----...J

O,..,C/C> ~\)Gtz::> C7 v Q Or

v ~
~~i~'~
(~~
c

r-; /',
Fig. 8. (A) Septaria betulae (Lib .) Westend. ex Sacc., from the authentic specimen in PAD ; (B)
Cylindrosporium betulae from the holotype; (C ) Asteroma microspermum from the holotype of Septoria
microsperma.

which are narrower. In one specimen from Japan,
labelled S. chinensis (TFM 0779), the fungus is
similar to S. betulae Pass.
SEPTORIA CORYLl Desm. apud Sydow, Myc. march.
1762 ( 1889) , nom. nud.
Sp ecimens examined: On Corylus avellana L. , Germany,
Berlin, Thiergarten, Oct. 1887, P . Sydow, S, UPS .
In both specimens no fungus could be detected .
One leaf in UPS shows yellowish spots charac-
teristic for Asteroma coryli (F u ckel) Sutton.
SEPTORIA MICROSPERMA Peck, Rep. N . Y. St . Mus.
nat. Hist . 34 : 44 (1883). (Fig. 8C)
Rhabdospora microsperma (Peck) Kuntze, Rev.
Gen. Pi. 3(3) : 512 (1898) .
Specimens examined : On Betula lenta L. , USA,
Knowersville, Oct., C. H. Peck, holotype, NYS ; Nuttall-
burg, West Va ., Oct. 1894, L.W. Nuttall, Ell. & Ev .,
N . Amer. Fung i 3276, UPS; ditto in Ell. & Ev., Fungi
Columbo 674, CUP, K .
Septoria microsperma Peck is the basionym of
Asteroma microspermum (P eck) Sutton.
 Septoria spp. on Betulaceae
I am in debted to the directors and curators of the
herbaria BPI, BR, BUCM, CUP, DAOM, H, K,
LE, NY, PAD, PC, RMS, S, TFM, UPS, W, WIS
and Inst. Plant Protection As, Norway for loan of
material, to Professor O. Hedberg for working
facilities provided, to E . Gunnerbeck for advice
concerning the nomenclature of S . betulae Pass.,
and to Dr B. C. Sutton for comments on the
manuscript. This work was financed by the
Swedish Natural Science Council grant 065-108 .
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