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Fossil remains of the mycorrhizal fungal
Glomus fasciculatum
complex in postglacial
lake sediments from Maine'
R. Scorr ANDERSON'
Department of Botany and Plant Pathology and lrlstit~lte ,fi)r Quclternary Studies, Ur~ive~sity of Maine,
Orono, ME, U.S.A.
04469
RICHARD L. HOMOLA
Department
qj'
Botany and Plarlr Pathology, University oJ'Maine, Orono. ME, U.S.A.
04469
AND
RONALD B. DAVIS
AND
GEORGE L. JACOBSON, JR.
Department
of Botany and Plant Pathology and lrl.sriture for Quaterrlclry Studies, University uf Maine,
Orono, ME. U.S.A.
04469
Received January 3, 1984
ANDERSON,
R. S., R. L.
HOMOLA,
R. B.
DAVIS,
and G. L.
JACOBSON,
JR.
1984. Fossil remains of the mycorrhizal fungal
C1omu.s fa.sciculaturn
complex in postglacial lake sediments from Maine. Can.
J.
Bot.
62:
232552328,
Specimens of the vesicular-arbuscular mycorrhizal fungal
C1otnu.s fascicuk~rum
complex have been identified from lake
sediment cores from Gould Pond (central Maine) and Upper South Branch Pond (north central Maine). The fungus became
established with tundra vegetation on newly developing soils soon after the melting of Wisconsin ice. This is the first record
of such an occurrence for North America. The earliest specimens from Gould Pond were deposited in sediment dated at about
13 000 years old and those at Upper South Branch Pond between about 12 500 and
I
I
000 years
old.
This is at least 1000 years
prior to the arrival of trees in those areas. Sedimentologic and biologic evidence
indicates
soil instability and relatively high
soil erosion rates from the watersheds during the late-glacial period. Thus erosion probably accounts for the abundance of
G1omu.s
in late-glacial sediments. The reduced abundance of the fungus in Holocene sediments is attributed to a decrease in
the rate of soil erosion after the establishment of trees.
ANDERSON,
R. S., R. L.
HOMOLA,
R. B.
DAVIS
et G. L.
JACOBSON, JR.
1984. Fossil remains of the mycorrhizal fungal
Clornlls
fa.sciculnrum
complex in postglacial lake sediments from Maine. Can.
J.
Bot.
62:
2325-2328.
En examinant des carottes de sediments lacustres provenant de Gould Pond (Maine central) et du Upper South Branch Pond
(Maine centre-nord), les auteurs ont identifiC des spCcimens du complexe
Clotnus fir.sciculaf~cn~,
champignons formant des
endomycorhizes
a
vCsicules et arbuscules. Les champignon s'est Ctabli avec la vCgCtation de toundra sur les sols en formation,
peu aprks la disparition des glaces du Wisconsin.
I1
s'agit de la prcmikre mention d'une telle prCsence en AmCrique du nord.
Les spCcimens les plus anciens
it
Gould Pond ont CtC dCposCs dans le sCdiment
il
y a cnviron 13 000 ans, et ceux de Upper
South Branch Pond, entre 12 500 et l
l
000 ans. Ceci Ctait au moins 1000 ans avant I'arrivCe des arbres dans ces rCgions. Les
donnCes ~Cdimentologiques et biologiques indiquent la prCsence d'une instabilite du sol et d'unc erosion du sol relativement
importante dans ces bassins hydrographiques pendant la fin de la pCriode glaciaire. Ainsi
il
est probable que I'abondance des
Glomus
dans les sCdiments tardi-glaciaires soit like
a
I'Crosion. La diminution dans I'abondance du champignon dans les
sCdiments de I'holocknc est attribuCe
a
une diminution dans le taux d'krosion du sol
lie
a
['apparition des arbres.
[Traduit par le journal]
Introduction
The symbiotic relationships between plants and mycorrhizal
fungi are now recognized to be important to the health and
productivity of many plant taxa. Under certain conditions my-
corrhizae may enhance growth (Gerdemann 1968), protect the
host plant from pathogens (Marx 1972), or both. Vesicular-
arbuscular (VA) mycorrhizae (endomycorrhizae) of genera
such as
Glomus
are found in association with a great variety of
cultivated plants and plants of natural forest communities.
These relationships are undoubtedly not recent in origin (Butler
1939; Malloch et al. 1980; Wagner and Taylor 1981). How-
ever, there is little evidence that indicates how quickly mycor-
rhizal plants became established in developing postglacial soils
which initially consisted of barren glacial deposits, open to
colonization by many forms of biota.
Lake sediments usually contain fossils that provide detailed
'A contribution of the Institute for Quaternary Studies, University
of Maine, Orono, ME, U.S.A. 04469.
'Present address: Department of Geosciences, Laboratory of
Paleoenvironmental Studies, University of Arizona, Tucson, AZ,
U.S.A. 85721.
evidence about the past characteristics of the lake water, vege-
tation and soils of the watershed, and the interactions of vege-
tation and environment. We have been examining lake sedi-
ments of Upper South Branch Pond (hereafter called USBP),
located at 46'5'
N
by 68'54' W in north central Maine, and
Gould Pond, 44'44' N by 69'19' W in central Maine (Fig. I),
to deduce the characteristics of the vegetation and abiotic envi-
ronment surrounding the lakes in former times.
The sediments of both lakes contain fossil fungal hyphae
and, although fungal hyphae and spores are commonly found
in lake and wetland sediments (see for example, Bradbury
1978; Birks 198
1
;
Tolonen 1983), late-glacial and postglacial
fossil mycorrhizal fungi from North American sediments are
rarely identified and reported (for European reports see van
Gee1 1978; van Gee1 et al. 1980). Such reports from North
America have been limited to localities in Alberta (Butler
1939), Minnesota (Rosendahl 1943), North Carolina (Wolf
1969), and New York (Miller 1973). We report here on the
occurrence of specimens of the vesicular-arbuscular fungus,
Glomus fasciculatum
(Thaxter sensu Gerdemann) Gerdemann
and Trappe complex, in ancient sediments from the two local-
ities in Maine.
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2326
CAN.
J.
BOT.
VOL.
62,
1984
FIG.
I.
Locations of Upper South Branch Pond (Pd.) and Gould
Pond. ME.
Methods
Sediment cores were collected with a modified Livingstone piston
coring apparatus (Livingstone 1955). Subsamples of the sediment
cores used for macrofossil analysis were first disaggregated in dilute
KOH
solution (USBP) or in distilled water (Gould Pond). The slurry
was then washed gently through a series of sieves and collected on a
180 pm mesh sieve. Until examined, the macrofossils were stored in
either a Formalin solution (USBP) or ethyl alcohol (Gould Pond).
Specimens of the fungus selected for photography were then mounted
in 70% alcohol.
Results
Specimens of the fossil fungus from either six (USBP) or
seven (Gould Pond) levels were examined. The specimens
included both chlamydospores and hyphae (Figs. 2 and 3).
Chlamydospores occurred in loose aggregations or in small,
compact clusters of 4 to 27. The chlamydospores are sub-
globose to globose, with diameters of 52
X
67 to 100
x
125 pm. Two separate walls are evident in these specimens.
The outer thick wall is smooth, has adhering debris, occa-
sionally appears irregularly pitted, and is yellow-brown in
color. The inner membranous wall is thinner and lighter in
color. The hyphal attachment is ca. 7 pm in diameter, with a
wall near the chlamydospore attachment ca. 4 pm thick and
yellow-brown in color.
A difference in interpretation of G. fasciculatum exists
among taxonomic specialists of the Endogonaceae (Gerdemann
and Trappe 1974; Trappe 1982; S. Berch, personal commu-
nication). The fungus from the two Maine locations has been
identified as belonging to the G. fasciculatum group
(J.
Trappe,
personal communication; Trappe 1982). It is possible that more
than one species is involved. Because of this uncertainty, we
have used the term "Glomus fasciculatum complex."
The fossil Glomus fasciculatum complex material from
Gould Pond was recovered from sediment deposited between
ca. 13 000 years BP and ca. 10 900 years BP. The USBP
samples came from sediment deposited between ca. 12 500 and
ca. 10 200 years BP. Nonarboreal pollen types, such as wil-
FIG. 2. Fossil members of the
Glomus fasciculat~tm
complex,
Upper South Branch Pond, ME, lake sediment. Chlamydospores and
hyphae. Ca. 142~.
FIG. 3. Fossil member of the
Glomus fnsciculatum
complex,
Upper South Branch Pond, ME, lake sediment. Chlamydospore
enlargement. Ca. 700x.
lows (Salix spp.), sedges (Cyperaceae), grasses (Gramineae),
and other herbs, attained maximum postglacial values during
these time periods (Anderson and Davis 1980; Anderson et al.
1982). Vaccinium uliginosum and Dryas integrifolia plant mac-
rofossils are found in many of the same sediment subsamples
as the fungus, as well as contiguous subsamples. Macrofossils
of arboreal species, beginning with spruce (Picen), are found
in sediment deposited after ca. 10 800 years BP at Gould Pond
and ca. 10 400 years BP at USBP. Tree remains dominated the
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ANDERSON
ET
AL.
2327
macrofossil assemblages throughout the Holocene.
Glomus
fasciculatum
complex specimens are found only at widely
spaced intervals (ca. 9000, 7700, 6500, 4000, and 100 years
BP) in USBP Holocene sediments and are not found in
Holocene sediments from Gould Pond.
Discussion
Fossil mycorrhizal fungi recovered from late-Quaternary
sediments were first reported by Butler (1939) as species of
Rhizophagus
Dang. from glaciolacustrine and peaty sediments
("muskeg") near Edmonton, Alta., Canada. Later, Rosendahl
(1943) described two slightly differing species from Minnesota
(Bronson, Jackson, and Moorhead sites):
Rhizophagites butleri
Rosendahl (vesicles light brown, 75
X
79 to 103
x
124 pm)
and
R.
rninnesotensis
Rosendahl (vesicles dark brown, 42
X
46
to 58
x
6
1
pm). Wolf (1969) recovered
Endogone (Rhizo-
phagus)
in peaty lake sediment from New Bern, NC. Miller
(1973) described
Rhizophagites butleri
from glacial Lake
Iroquois sediments in Lockport, NY. Later, Gerdemann and
Trappe (1974) placed
R. butleri
in synonymy with
Glomus
fasciculatum.
Although none of the above specimens was found conclu-
sively in organic connection with other plant materials, infor-
mation on associated fossil plants is available for all specimens.
Only the Alberta specimens remain undated.
Picea-Larix
bog
remains (Bronson and Jackson sites) and boreal forest litter
deposits (Moorhead site) are associated with the Minnesota
fungi (Rosendahl 1948). Radiocarbon dates on these materials
have been reported as follows: Bronson site, greater than
36 000 years BP (Flint and Rubin 1955); Moorhead site,
9930
t
280 years BP (Wright and Rubin 1956). On strati-
graphic grounds, the Jackson, MN, material is no less than
13 700 years old (Denton and Hughes 1981). The Lockport,
NY, fungus, dated at about 12 000 years BP, was associated
with plants from a wide variety of habitats, and the vegetation
was interpreted as being similar to boreal forest. No associated
plants were reported for the ca. 24 000 year old North Carolina
materials (Wolf 1969), but D. A. Livingstone (personal com-
munication) suggested a forest association there also.
Members of the
Glomus fasciculatum
complex are known to
occur naturally with a wide variety of plant species character-
istic of temperate and boreal forest ecosystems (Gerdemann
and Trappe 1974); as mentioned above, the fossil specimens in
the previous studies were found in association with plants char-
acteristic of such environments. Unnamed VA endomycor-
rhizae have also been reported from modern tundra soils near
Barrow, AK (Miller and Laursen 1978). These authors found
an association with several members of the Ranunculaceae,
Gramineae, Cyperaceae, and
Saxifraga
and with at least one
moss,
Pogonatum.
However, the occurrence of members of the
G. fascirulatum
complex in the lowest sections of the two
Maine cores is significant because it is the first known occur-
rence of the fungus deposited with late-glacial tundra plant
assemblages from North America. Pollen and macrofossils
from sediment levels containing the fungus, and from adjacent
core intervals, suggest that during this period of tundra around
Gould Pond and USBP, sedges (Cyperaceae) and grasses
(Gramineae) were common along with, at USBP,
Oxyria
digyna
(L.) Hill and
Arenaria rossii
Richards. (Anderson and
Davis 1980; Anderson et al. 1982).
Saxifraga
pollen was
recovered from Gould Pond sediments (R. S. Anderson,
unpublished data). The most common dwarf shrubs included
Dryas integrifolia
M. Vahl,
Salix
spp.
(S. herbacea
L. and
S. uva-ursi
Pursh at USBP),
Vaccirzium
cf.
uliginosum
and
V.
cf.
vitis-idaea. Pogonatum alpinum
(Hedw.) Rohl. grew
around USBP, along with at least two species of arctic-alpine
mosses,
Didymodon asperifolius
(Mitt.) Crum, Steere and
Anders. and
Aulacomnium turgidum
(Wahl.) Schwaegr.
(N. Miller, personal communication). No macrofossils of trees
were found in sediments deposited during the tundra period.
From macrofossil evidence, spruce did not grow around
Gould Pond until ca. 10 800 years BP and USBP until
ca. 10 400 years BP. The fungi were present in soils for at least
1000 years before the arrival of tree species.
The only sporadic occurrence of specimens of the
Glomus
fasciculatum
complex in posttundra sediments of USBP and the
absence of the fungus in posttundra sediments of Gould Pond
are best explained by the stabilization of soils and subsequent
decrease in erosion around the lakes, following the establish-
ment of arboreal species. The lake sediments deposited during
the tundra period contain higher percentages of inorganic
matter derived from the terrestrial environment, indicating that
soils were probably less stable during the tundra period than in
later times. The 41 samples of late-glacial tundra sediments at
USBP average 97.4% inorganic matter, whereas the 153 sam-
ples of Holocene sediments average 78.0% inorganic matter.
At Gould Pond, the 19 samples of late-glacial tundra sediments
average 97.2% inorganic matter, while the 45 samples of
Holocene sediments average 77.3% inorganic matter. A link
between establishment of trees and a decrease in de~osition of
inorganic matter has also been suggested for Mirror Lake, NH,
by Davis and Ford (1982), who found that the postglacial
arrival of
Picea
in the region coincided with a precipitous
decline in rates of inorganic accumulation within the basin.
In addition to sedimentary indicators of landscape insta-
bility, early successional mosses that colonize bare rock
(Grimmia
sp.,
Arctoa- Kiaeria
sp.,
Andreaea rothii
Web.
&
Mohr) and mineral soil
(Ceratodon purpureus
(Hedw.) Brid.,
Pogonatum alpinum
(Hedw.) Rohl.,
Polytrichum
sp.) are com-
mon at USBP (N. Miller, personal c.ommunication). These
species occur as fossil only in the sediments deposited during
the tundra period. Based on pollen evidence, Spear (1981)
suggested that periglacial frost activity on the White Mountains
of New Hampshire during the late-glacial period contributed to
landscape instability. This would have provided continuously
fresh mineral surfaces near the lakes. The periglacial frost
hypothesis is strengthened if relict periglacial features are
found within the area.
Acknowledgements
We thank Dr. James M. Trappe for tentative identification of
the fossil specimens of
Glomus fasciculatum;
Dr. Norton G.
Miller for sharing unpublished information on the fossil fungi
from Lockport, NY, and fossil mosses and
Arenaria
from
USBP, ME; and Dr. Robert Stuckenrath, for the radiocarbon
dates on sediment from the two Maine lakes. This work was
supported, in part, by a Sigma Xi grant-in-aid of research to
R. S. Anderson, and by National Science Foundation grant
DEB 8004171 to G. L. Jacobson, Jr., and R. B. Davis.
ANDERSON,
R.
S., and
R.
B. DAVIS. 1980. Late- and postglacial
vegetational history at Upper South Branch Pond
in
northern
Maine. Sixth Biennial Meeting, AMQUA, Orono, ME, U.S.A.
p.
5.
(Abstr.)
ANDERSON,
R.
S.,
G.
L.
JACOBSON,
JR., and
R.
B. DAVIS. 1982.
Late- and postglacial environmental change
in
central Maine: a
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2328
CAN.
J.
BOT.
VOL.
62,
1984
record of marine, lacustrine and terrestrial fossils from Gould Pond.
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