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Mycologia,
82(4), 1990, pp. 452-459.
? 1990, by The New York Botanical
Garden,
Bronx,
NY 10458-5126
NEW SPECIES OF PHYLLOPORUS AND TRICHOLOMOPSIS
FROM COLOMBIA, WITH NOTES ON
PHYLLOPORUS BOLETINOIDES
ROLF SINGER
Department
of Botany,
Field Museum
of Natural
History, Chicago,
Illinois 60605-2496
CLARK
L. OVREBO
Department
of Biology, Central
State University,
Edmond,
Oklahoma
73034
AND
ROY E. HALLING
The New York
Botanical Garden,
Bronx, New York 10458-5126
ABSTRACT
Phylloporusfibulatus
and Tricholomopsis
humboldtii from Colombian
oak forests are
newly
described,
and a new section of Phylloporus,
sect. Fibulati,
is proposed.
Phylloporus
boletinoides is redescribed
from recently
collected material
and placed in sect. Manausensis.
A comparison
of characters
distin-
guishing
Paxillus and Phylloporus
is given.
Key Words:
Paxillus, Phylloporus,
Tricholomopsis,
Quercus,
Colombia,
Andes Mountains.
The study of recent collections made by Hall-
ing and Ovrebo in Colombian oak forests has
resulted in the discovery of two new fungus taxa.
One, a boletinoid, lamellate fungus, Phylloporus
fibulatus, is interesting not only because the data
indicate an unpublished taxon, but the species
also contributes to a better understanding of the
infrageneric taxonomy of Phylloporus and to a
further clarification of the differences between
Phylloporus and Paxillus. A new section is pro-
posed to accommodate it. The second, Tricho-
lomopsis humboldtii, is the first species of that
genus known to occur on wood of Quercus hum-
boldtii Bonpl., a substratum of widespread but
apparently diminishing supply in Andean Co-
lombia.
Finally, recent collections of Phylloporus bole-
tinoides Smith & Thiers made in Florida by R.
S. Williams and in Texas and Alabama by D. P.
Lewis provide the basis for a redescription of
that species. Further study of these specimens
indicates that P. boletinoides has affinities with
species in Phylloporus sect. Manausensis.
In the descriptions below, colors are indicated
in terms of Komerup and Wanscher (1978),
Munsell (1976), and some, marked by "M & P,"
are the terms of Maerz and Paul (1930). The
symbol "Q" represents the basidiospore length/
width ratio and "Q" represents the mean length/
width ratio.
Phylloporus fibulatus Singer, Ovrebo, & Halling,
sp. nov. FIGS. 1, 6, 7
Pileo flavo, subsordescente,
ad marginem
brunnes-
cente, subtomentoso,
opaco, convexo, dein applanato
vel applanato-depresso,
11-35 mm lato. Lamellis
flavis
dein flavoviridantibus vel flavido-griseis,
subdistan-
tibus, furcatis
et moderatim
intervenosis,
decurrenti-
bus. Stipite flavo, ad apicem rubido-cincto,
lentis as-
perulato-fibrilloso,
aequali
vel basin versus
attenuato,
15-45 x 2.5-9 mm; velo nullo; mycelio basali flavo.
Came flava interdum
fractu subcaerulescente;
odore
fragrante;
sapore
debili, haud notabili.
Sporis 8-11 x
3-4.5 ,m, plerumque
8.5-10 x 3-4 ,um,
oblongis,
fu-
siformibus,
cylindraceis, depressione suprahilari
debili
instructis,
levibus, colore in cumulo brunneolo-oliva-
ceo, sub microscopio
Xerocomorum
gaudentibus,
in-
amyloideis. Basidiis 23-41 x 6-8.5 ,um,
tetrasporis.
Cystidiis (29-)50-104 x 6-10.4, m, subventricosis vel
subcylindriceis,
interdum subcapitatis
vel subclavic-
ulatis vel apiculatis,
hyalinis,
interdum
guttulis
minutis
aureo-flavis
repletis,
tenui-tunicatis.
Hyphis multis fi-
bulis
omatis, in tramate
hymenophorali
bilaterali
(typi
Phyllopori)
haud
gelatinosis, hyalinis.
Epicute
pilei ex
hyphis intertextis
(trichodermium
haud palisadicum)
452
SINGER ET AL.: COLOMBIAN FUNGI
'~~~~~~~~.1
?.
?
I I^SI
EiSW*
.46pl r -4-
FIGS. 1-2. 1. Phylloporus fibulatus (holotype), carpophores, x 1. 2. Tricholomopsis humboldtii (holotype),
carpophores, x 1.
453
r
le - --
MYCOLOGIA
formata,
sublaxa
sed non gelatinoso,
hyphis elongatis
3.3-11 gm crassis. Ammoniaci (NH40OH)
ope super-
ficies pilei reactionem brunneolam nec caeruleam
provocat.
TYPUS in Colombia,
Antioquia,
sub Quer-
cus humboldtii
a C. L. Ovrebo 2546 lectus (HUA).
Pileus 11-35 mm wide, convex to plane or
plano-depressed, with downcurved to incurved
and often wavy edge; surface dry, dull, appearing
woolly-felted, yellow (3A6) overall, becoming
somewhat duller yellow (3A6-4A6, 4B8) in age,
brownish tinged near margin when older. Con-
text 4-5 mm thick, yellow (3A5), not discoloring
or with a hint of blue in places; odor fragrant;
taste slight, nondescript. Lamellae 2-3 mm wide,
decurrent, subdistant (2-3 mm apart for through-
lamellae), distantly anastomosing at half height
or when young, moderately low intervenose, not
distinctly boletinoid, with the lamellulae nu-
merous but not in distinct tiers and often con-
necting with the neighboring through lamellae,
yellow when young, yellowish green (4C5-4D6)
to yellowish gray (5D3-5E3) when mature, not
discoloring. Stipe 15-45 mm long, 2.5-9 mm
thick, equal or tapering downward, surface slightly
roughened-fibrillose, yellow (3A5-4A5) with a
reddish zone at the apex; context solid, yellow,
not discoloring; basal mycelium pale yellow to
yellow.
Macrochemical color reactions: 2.5% KOH
quickly ochre on all parts; 10%
FeSO4 olivaceous
on context and lamellae; NH4OH brownish on
surface of pileus, deep purple brown at junction
of pileus surface and trama.
Basidiospores brownish olive in fresh deposit,
8-11 x 3-4.5 ,um, most frequently 8.5-10 x 3-
4 Am (Q = 2.5-4, Q = 2.7), generally with a slight
suprahilar depression, fusiform to elliptical in
face view, smooth, rather thin-walled, dull gold-
en in KOH, inamyloid. Basidia 23-41 x 6-8.5
Am, narrowly clavate, 4-sterigmate, hyaline to
pale yellowish. Hymenial cystidia (29-)50-104
x 6-10.4 Am, cylindric with the mid-portion
swollen to narrowly fusiform-ventricose, with the
apex sometimes subcapitate, with rounded tip,
smooth, thin-walled, hyaline. Hyphae of hyme-
nophoral trama 3.5-14 ,m wide, hyaline, slightly
divergent when young and of the Phylloporus-
type, slightly interwoven when mature, cylindric,
many refractive and similar in appearance to
oleiferous hyphae, yellow in KOH, inamyloid or
amyloid reaction disappearing after 5-10 min-
utes. Subhymenial hyphae 2.8-3 Am wide,
scarcely distinctive as a layer. Hyphae of pileus
surface 3.5-9.2 ,m wide, interwoven to radially
interwoven, ?cylindric and all strongly elongat-
ed, smooth, thin-walled, with or without a gran-
ular content, light yellow in KOH, forming a
trichodermium when young, with the end cells
rather loosely arranged but not gelatinized. Hy-
phae of the pileus context 3.5-17 um wide, cy-
lindric, hyaline to light yellow, inamyloid. Hy-
phae of the stipe surface 3.5-5.8 ,um wide,
appressed to slightly interwoven, cylindric,
smooth, thin-walled, hyaline. Hyphae of the stipe
context 4.6-17 Mm
wide, parallel and compacted,
cylindric, pale yellow, inamyloid. Clamp con-
nections present, numerous.
Solitary or gregarious in Quercus humboldtii
forests during November.
TYPE: Colombia. Departamento
Antioquia:
along
road
from Santa
Rosa de Osos to San
Jose de la Mon-
tafia, 25.XI.1986, Ovrebo 2546 (HOLOTYPE:
HUA;
ISOTYPE: NY).
PARATYPES: Colombia. Departamento Antioquia:
Municipio Santa Rosa de Osos, + 11 km N of Santa
Rosa de Osos, road to Aragon,
near Llanos de Cuiva,
6?45'N, 75?30'W,
+2500 m elev, under
Quercus
hum-
boldtii,
5.XI.1986, Halling 4983 (HUA, NY). Depar-
tamento Narifio:
Municipio Pasto, 11
km E of Chacha-
gui, bosque "El Comfin," 22.XI.1988, Halling 6132
(NY, PSO); km 17, road from Pasto to Chachaguii,
vereda
"La
Josefina,"
23.XI.
1988, Halling 6153 (NY,
PSO).
In all species of Phylloporus known to date,
the base of the basidia and cystidia as well as the
other hyphae have simple septa, and only very
exceptionally in a few species can a clamp-bear-
ing septum be found. In all species of sect. Phyl-
loporus (in contrast to sect. Manausensis), the
fresh spore print has an olive hue. Further, in all
species except those in sect. Manausensis, a char-
acteristic and distinct (not fleeting) blue or green-
ish blue ammonia reaction is observed similar
to that seen in Xerocomus sect. Pseudophyllopori
Sing. Until the discovery of sect. Manausensis,
the absence of clamps, the olive tinted spore print,
and the blue ammonia reaction were considered
to be characteristic for the whole genus. Thus,
Pegler and Young (1981) considered that paxil-
loid species without clamp connections belonged
to Phylloporus and consequently transferred
Paxillus defibulatus Sing. [? = Paxillus statuum
(Speg.) Horak] to Phylloporus. The species de-
scribed above is an exception in Phylloporus. Be-
cause of the presence of clamp connections in P.
fibulatus, we propose the following new section
for this interesting species:
454
SINGER
ET AL.: COLOMBIAN
FUNGI
Phylloporus sectio Fibulati Singer, Ovrebo, &
Halling, sect. nov.
Hyphis fibulatis.
TYPUS: Phylloporus
fibulatus Sing., Ovrebo, &
Halling (see above).
PHYLLOPORUS
BOLETINOIDES
Smith & Thiers,
Contr. N. Amer. Sp. Suillus, 105. 1964.
Phylloporus boletinoides has been recollected
with fresh notes by Robert S. Williams in Florida
and by David P. Lewis in Alabama and Texas.
These recent collections indicate that P. boleti-
noides has most characters in common with sect.
Manausensis. The following description is based
on these recent materials and observations (mi-
croscopical data by the senior author).
Pileus some shade of brown [5 YR 3.5/4; 5
YR 2/4; 5 YR 3/4; YR 4/4 (Munsell, 1976); 7D5
to 8E6 (Kornerup and Wanscher, 1978)], some-
times showing pallid yellow in a few shallow pits,
velvety, glabrescent, later sometimes subfibril-
lose-subsquamulose, convex to plano-convex
with somewhat involute margin, then partly ap-
planate with straight margin, 23-75 mm broad,
circumference circular, rarely broadly elliptical.
Context whitish, then often dingy, unchanging
on bruising, sometimes more bluing observed
(DPL 4015), rather thick. Odor mild to fungoid.
Taste mild. Hymenophore lamellose-subporoid,
often more poroid near stipe and more lamelloid
and forked near margin, also vice versa, but pores
formed by lower anastomoses and rather wide
(2-5 x 2-3 mm), 5-8 mm deep, mostly close or
subclose when young and soon becoming sub-
distant, often with short lamellulae at margin,
2.5 Y 4/4 or 5 Y 6/4 or 10 YR 2/2 or 4B5 to
4B5, later colored like spores, unchanging when
bruised. Stipe 2.5 Y 8/3 at apex, towards the base
concolorous with pileus or more reddish, 9D6,
glabrous but with some decurrent lines below
hymenophore, these lines sometimes anasto-
mosing, solid, becoming cavernous in age, cen-
tral or subcentral, straight or curved; veil none;
basal mycelium whitish or "putty" (M & P).
Chemical color reactions:
NH4OH on surface
of pileus reddish purple, or wine color on context
and slightly darkening or little reaction; on hy-
menophore darkening. Five percent KOH slight-
ly darker or little reaction, 10% KOH dirty yel-
low on surface of pileus, on context little reaction,
slowly becoming brownish. FeSO4 on context
slowly and slightly grayish green.
Basidiospores "cognac" in fresh deposit, with-
out any olive tint; when dried "cochin" (M & P)
or more sepia or 10 YR 3/2; 10.5-14.5 x 3.5-
6.5 ,m, most frequently 10.5-12 x 4.5-5 ,im,
fusoid with obtuse ends to oblong-fusoid, more
rarely cylindric, rarely ovoid or ellipsoid, (Q =
1.5-3.3, most frequently 2.6-3, Q = 2.7), smooth,
most with suprahilar depression, with firm to
slightly thickened wall, without germ pore, mel-
leous brown or stramineous (from spore print,
RSW 326), inamyloid. Basidia 27-35 x 7.5-9.7
Am, 4-spored; basidioles in some specimens
tending to be transformed later to cystidioles,
16-23 x 6.5-7 Am, clavate, thin to thick-walled
(up to 1 um thick). Cystidia numerous on and
near edges, less numerous on sides of lamellae,
40-65(-75) x 10-19 gm, mostly fusoid or am-
pullaceous and then with 5-8 ,um broad apex,
with mostly obtuse or mucronate tips, without
resinous incrustation, thin-walled, hyaline, but
at edge often brownish inside. Hymenophoral
trama inamyloid, bilateral of the Phylloporus-
type, with the mediostratum rather dense, of sub-
parallel to slightly interwoven hyphae, 1.5-5 ,tm
broad, forming a 20-25 um wide layer, hyaline
or melleous-hyaline, non-gelatinous; lateral stra-
tum concolorous and 75-80 gm wide, slightly
gelatinous and slightly divergent, only in places
slightly curved outward, of hyphae 4-8 ,um
broad
and moderately spaced. Pileus surface composed
of mostly upright, not fully parallel hyphae, the
terminal ones 28-56 x 6-11 Am, not encrusted
in KOH, with rounded, rarely subacute tip, in-
amyloid and not orange-rufous in Melzer's re-
agent; hypodermium present, interwoven, ex-
posed in places where surface is glabrescent.
Clamp connections occasional and then only on
the non-encrusted, hyaline hyphae of the basal
mycelium.
Often gregarious or cespitose on the ground
among litter in mixed Pinus-Quercus woods.
Material examined: FLORIDA. Sarasota Co.:
30.XII.
1986, R. S. Williams 326 (F). ALABAMA.
Bald-
win Co.:
Orange
Beach,
21 .VII.
1982,
D. P. Lewis
3196
(F).
TEXAS.
Orange
Co.:
Vidor, 17.IX.
1986,
D. P. Lewis
4015 (F).
This species shares the non-bluing NH4OH
reaction with P. fibulatus. It enters sect. Ma-
nausenses but differs from that section in the lack
of blackening on the hymenophore and stipe. It
would seem that a special subsection with species
that do not blacken on injury may eventually be
455
MYCOLOGIA
proposed. However, it may be prudent to wait
for chemical data on the other blackening species
from Malaya (Comer, 1970) now tentatively in-
serted in sect. Phylloporus.
Both P. boletinoides and P. fibulatus differ from
Paxillus in several ways. In the first place, they
are cystidiate, showing the same type found in
other species of Phylloporus. Cystidia which are
similar in form can be found only in Paxillus
sect. Paxillus, but in that section the spores are
short and ellipsoid unlike the elongate-boletoid
spores of Phylloporus. Furthermore, the brown
staining of the context and lamellae (oxidation
of involutin) is quite different from the unchang-
ing or bluing context (oxidation of pulvinic acid,
etc.) of Phylloporus. Even macroscopically the
differences are obvious: the lamellae in Paxillus
sect. Paxillus are close and the pileus margin is
involute for a long time. In Phylloporus, the la-
mellae are generally more distantly spaced and
the pileus margin is incurved only in youth.
Chemically, both species treated here are not yet
analyzed, and P. manausensis does not have, at
least in the carpophores, any identified pigments.
On the other hand, the chemistry of P. rhodo-
xanthus (sect. Phylloporus) is sufficiently differ-
ent from the only clampless Paxillus [P. defi-
bulatus Sing. = P. statuum (Speg.) Horak sensu
Horak] (Garrido, 1988, p. 123).
Tricholomopsis humboldtii Sing., Ovrebo, &
Halling sp. nov. FIGS. 2, 3-5
Pileo nigro, dein plerumque
inter tomentum dif-
fractum
flavo-tincto,
applanato
inrepandis,
sicco, 25-
100 mm lato. Lamellis
flavis, subdistantibus,
adnexis
vel adnatis.
Sporis
in cumulo albis. Stipite supra
fun-
dum flavum nigrello,
lanoso, mox punctato vel sub-
sericello,
20-65 x 5-20 mm. Came
dilute
flava; sapore
miti;
odore subnullo.
Sporis
6.5-7.5 x 3-4 Am,
levibus,
inamyloideis, hyalinis. Basidiis 33-43 x 5.8-7 ,im,
tetrasporis.
Cheilocystidiis
32-63 x 5.8-15 ,um,
cylin-
draceis vel subcylindraceis,
obtusis, tenuitunicatis,
hy-
alinis vel intus obscuris pigmento intracellulari,
dis-
soluto. Pilei et stipitis tegumento continuo vel
discontinuo, trichodermiali,
cellulis hyalinis vel pig-
mento obscuro intracellulari
repletis,
fibulatis derma-
tocystidiis
prasentibus.
Ad lignum
emortuum
Quercus
humboldtii
in Colombia,
HOLOTYPUS
Ovrebo
2554
(HUA).
Pileus 25-100 mm wide, irregularly plano-
convex when young, expanding to nearly plane
with a wavy-irregular margin, often depressed at
the center, edge incurved when young, straight
or uplifted when older, surface dry, dull, felt-like
on young buttons, when mature still having felt-
ed appearance but surface layer separating some-
what to reveal yellow ground color, appearing
minutely punctate under a 10 x lens, when older
becoming rimose, black when young, when ma-
ture dark brownish black with faint yellow ground
color showing through near the edge, occasion-
ally yellowish with a blackish overtone; context
3-10 mm thick, light yellow, odor and taste none.
Lamellae 3-15 mm wide, adnexed to adnate,
when young bright yellow (4A8), fading some-
what in age to a dull light lemon yellow (duller
than 3A4), not discoloring, subdistant (5-7 mm
apart for through-lamellae), slightly rugulose-in-
tervenose at stipe apex, entire; lamellulae nu-
merous but not arranged in distinct tiers. Stipe
20-65 mm long, 5-20(-30) mm thick, equal te-
rete or flattened slightly, the base rounded, sur-
face on buttons blackish-woolly with yellow
ground color evident, when mature blackish
punctate to at times almost silky fibrillose, the
yellow ground color evident beneath the blackish
surface fibrils or punctate vesture; context fleshy-
fibrous, hollow, light yellow.
Macrochemical color reactions: negative with
2.5% KOH and 10% FeSO4.
Basidiospores white in deposit, 6.5-7.5 x 3-
4 ,um, subreniform to cylindric in profile, cylin-
dric to oblong in face view, smooth, thin-walled,
hyaline, acyanophilous, inamyloid. Basidia 33-
43 x 5.8-7 ,um, 4-sterigmate, clavate, hyaline,
inamyloid. Cheilocystidia 32-63 x 5.8-15 Aim,
cylindric to somewhat clavate-inflated, smooth,
thin-walled, hyaline to dark translucent with
blackish intracellular content; content most ev-
ident on young carpophores. Hyphae of lamellar
trama 3.5-17 uim wide, parallel, cylindric to
slightly inflated, hyaline to light yellow, inamy-
loid. Hyphae of subhymenium 3.5 Aim
wide, cy-
lindric, hyaline, not distinctive as a layer. Hy-
phae of pileus surface forming a continuous to
discontinuous trichoderm which is most evident
at the center; hyphae often in fascicles, more or
FIGS. 3-5. Tricholomopsis
humboldtii
(holotype).
3. Basidiospores.
4. Cheilocystidia.
5. Dermatocystidia.
FIGS.
6-7. Phylloporus
fibulatus
(holotype).
6. Basidiospores.
7. Hymenial
cystidia. Scale line = 10 Atm,
long
scale line is for basidiospores,
short one is for cystidia.
456
SINGER ET AL.: COLOMBIAN FUNGI
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MYCOLOGIA
less interwoven in places; dermatocystidia 25-
135 x 3-8.5 Aim,
cylindric to subgeniculate, with
tips often slightly swollen, smooth, thin-walled,
nearly hyaline or with light to dark fuscous to
blackish, translucent intracellular content; con-
tent opaque and black in Melzer's. Hyphae of
pileus context 3.5-17.3 Am wide, cylindric to
slightly inflated, hyaline. Hyphae of stipe surface
4-5.8 Atm
wide, forming on young carpophores
a more or less continuous trichoderm and on
older ones a discontinuous trichoderm inter-
mixed with more or less repent hyaline hyphae;
trichodermal hyphae hyaline or with blackish,
opaque content like hyphae of pileus surface. Hy-
phae ofstipe context up to 17.3 ,um
wide, parallel,
compacted, cylindric to slightly inflated, hyaline.
Clamp connections present.
Gregarious on well decayed log (probably al-
ways Quercus humboldtii).
TYPE: Colombia. Departamento
Antioquia: Mu-
nicipio San Jos6 de la Montaiia,
along road from San
Jos6 de la Montania to Labores,
26.XI.1986, Ovrebo
2554 (HOLOTYPE:
HUA; ISOTYPES:
CSU, NY)
PARATYPES: Colombia. Departamento Antioquia:
Municipio
San Jos6 de la Montafia,
along road from
San
Jose de la Montania
to Labores,
26.XI.
1986, Ovre-
bo 2563 (HUA, NY).
The coloration of T. humboldtii is quite strik-
ing with its bright yellow lamellae contrasted
against the dark brownish black to black pileus
and stipe. The yellow ground color is evident
occasionally on the pileus, especially on older
carpophores, due to separation of surface fibrils
during expansion. The yellow ground color of
the stipe is generally more evident and so its
coloration usually is lighter than the pileus.
Microscopically, surface cells of the pileus and
stipe as well as cheilocystidia have a dark pig-
ment dissolved in the cell sap (FIGS.
4, 5). The
intensity of the pigment is most pronounced on
younger carpophores. The cheilocystidia are not
inflated but are mainly cylindric to only slightly
inflated. Although inflated cheilocystidia are more
typical in Tricholomopsis, slender ones are not
unusual for species in the genus. Both the pileus
and stipe surfaces have end cells of the tricho-
derm which we interpret as dermatocystidia. The
trichoderm is more or less continuous on youn-
ger carpophores, but becomes discontinuous on
expansion. Also, the spores are relatively narrow.
The yellow coloration of T. humboldtii places
this species in sect. Tricholomopsis where it ap-
pears to be comparable to T. araucariae Sing.
(Singer, 1953). The latter has bronze-colored la-
mellae, a different host [Araucaria angustifolia
(Bertol.) 0. Kuntze] and geographic distribution
(southern Brasil and northeastern Argentina), has
numerous pleurocystidia, broader cheilocystidia,
and the pileus has a different structure causing
the surface to be more squamulose to squarru-
lose. The Colombian species also comes rather
close to the common temperate T. decora (Fr.)
Sing. which is more yellow on the pileus and
stipe, has relatively broader spores and only oc-
casionally has blackish scales. Also, T. decora
grows exclusively on conifer wood. There are
also some other blackish species of Tricholo-
mopsis, e.g., T. nigra (Petch) Pegler (Pegler, 1986)
and T. tricholomatoides Sing., (Singer, 1989) but
these have white lamellae and context, grow on
bamboo clumps or are terrestrial, and have either
subglobose spores or differentiated pleurocys-
tidia. Tricholomopsis atrogrisea Pegler is smaller
(up to 3 5 mm broad) with pale gray lamellae and
broader spores (Pegler, 1983). Our species is the
first Tricholomopsis known to occur on wood of
Q. humboldtii. Further collecting will possibly
show that it occurs on other substrates or that
other species of Tricholomopsis may share this
substrate with T. humboldtii.
ACKNOWLEDGMENTS
Field work
in Colombia was supported by National
Science Foundation grant #BSR-8600424 to R. E.
Halling.
Material received
by R. Singer
from
NY (Du-
mont collections),
from
R. S. Williams
(Sarasota,
Flor-
ida) and D. P. Lewis (Vidor,
Texas) has also contrib-
uted to conclusions
expressed
in the discussions. This
assistance is herewith
gratefully acknowledged.
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Accepted
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459
