Assemblages of myxomycetes associated with three different substrates affected by forest wildfires

Abstract

Background and aims – In late November and early December of 2016, forest wildfires occurred over portions of the Great Smoky Mountains National Park (USA) and more than 4 000 ha were affected. Previous studies have shown that myxomycete assemblages can be greatly impacted as a result of this type of disturbance; after which, the recovery of the forest determines the availability of substrates for new colonisation. The objective of the project reported herein was to assess the impact of wildfires on the recovery of the assemblages of myxomycetes associated with three different substrates (forest floor leaf litter, the bark of living trees, and woody twigs) in two areas with different fire intensity.Material and methods – Two study areas subjected to different fire intensity were selected and sampled 30 months after the wildfires. Myxomycetes were studied using the moist chamber culture technique as it applies to these organisms. Satellite imagery was used to determine forest recovery and similarity indices were used to compare experimental myxomycete assemblages among study areas and substrates. Historical data were used as a reference to contextualise the results.Key results – A total of 38 species of myxomycetes representing 17 different genera were recorded from the two study areas. Samples from the lower intensity burn area yielded more myxomycetes than samples from the higher intensity burn area, with values of 84% and 59%, respectively. This same pattern was also observed for the number of recorded specimens (133 and 93, respectively). The comparison of experimental assemblages with previous data suggested that ground litter assemblages were still in early stages of recovery, whereas the assemblages associated with bark and twigs had recovered much faster.Conclusion – The relatively higher intensity fire had more of an effect on myxomycetes than the relatively lower intensity fire. Myxomycete assemblages are resilient to wildfires and they recover differentially depending on the substrate they grow on.

Full text

Plant Ecology and Evolution 154 (1): 15–27, 2021
https://doi.org/10.5091/plecevo.2021.1762
Assemblages of myxomycetes associated with three dierent substrates
aected by forest wildres
Steven L. Stephenson1, Nazrana Payal2, Gurpreet Kaur2 & Carlos Rojas3,*
1Department of Biological Sciences, University of Arkansas, Fayetteville, Arkansas 72701, USA
2Department of Biotechnology, Shoolini University of Biotechnology and Management Sciences, Solan-Oachghat-Kumarhatti Highway,
Bajol, Solan, Himachal Pradesh 173229, India
3Department of Biosystems Engineering and Engineering Research Institute, University of Costa Rica, San Pedro de Montes de Oca 11501,
Costa Rica
*Corresponding author: carlos.rojasalvarado@ucr.ac.cr
RESEARCH ARTICLE
Background and aims – In late November and early December of 2016, forest wildres occurred over
portions of the Great Smoky Mountains National Park (USA) and more than 4 000 ha were aected.
Previous studies have shown that myxomycete assemblages can be greatly impacted as a result of this
type of disturbance; after which, the recovery of the forest determines the availability of substrates for
new colonisation. The objective of the project reported herein was to assess the impact of wildres on the
recovery of the assemblages of myxomycetes associated with three dierent substrates (forest oor leaf
litter, the bark of living trees, and woody twigs) in two areas with dierent re intensity.
Material and methods – Two study areas subjected to dierent re intensity were selected and sampled
30 months after the wildres. Myxomycetes were studied using the moist chamber culture technique as it
applies to these organisms. Satellite imagery was used to determine forest recovery and similarity indices
were used to compare experimental myxomycete assemblages among study areas and substrates. Historical
data were used as a reference to contextualise the results.
Key results – A total of 38 species of myxomycetes representing 17 dierent genera were recorded
from the two study areas. Samples from the lower intensity burn area yielded more myxomycetes than
samples from the higher intensity burn area, with values of 84% and 59%, respectively. This same pattern
was also observed for the number of recorded specimens (133 and 93, respectively). The comparison of
experimental assemblages with previous data suggested that ground litter assemblages were still in early
stages of recovery, whereas the assemblages associated with bark and twigs had recovered much faster.
Conclusion – The relatively higher intensity re had more of an eect on myxomycetes than the relatively
lower intensity re. Myxomycete assemblages are resilient to wildres and they recover dierentially
depending on the substrate they grow on.
Keywords – Disturbance; ecological recovery; forest ecology; Great Smoky Mountains National Park;
North Carolina; slime molds; Tennessee.
© 2021 Steven L. Stephenson, Nazrana Payal, Gurpreet Kaur, Carlos Rojas.
This article is published and distributed in Open Access under the terms of the Creative Commons Attribution License (CC BY 4.0), which
permits use, distribution, and reproduction in any medium, provided the original work (author and source) is properly cited.
Plant Ecology and Evolution is published by Meise Botanic Garden and Royal Botanical Society of Belgium
ISSN: 2032-3913 (print) – 2032-3921 (online)
INTRODUCTION
Myxomycetes (plasmodial slime molds or myxogastrids) are
a group of amoebozoans associated with dead plant material
in virtually every type of terrestrial ecosystem investigated
to date, with approximately 1000 species known worldwide
(Lado 2005–2020). The myxomycete biota of the Great
Smoky Mountains National Park (USA) is especially well
known, largely as a result of surveys carried out in the context
of the All Taxa Biodiversity Inventory project (Stephenson
et al. 2001; Stephenson & Landolt 2009). More than 220
species have been recorded from the park. This total is as

16
Pl. Ecol. Evol. 154 (1), 2021
high as or higher than those recorded for any comparable
area of North America.
In temperate forests, myxomycetes are associated with a
number of dierent substrates. These include relatively large
pieces of coarse woody debris (e.g. stumps and fallen logs)
on the forest oor, woody twigs typically with a diameter of
no more than a centimetre, the bark surface of living trees,
forest oor leaf litter (also known as ground litter), the dung
of herbivorous animals, and aerial portions of dead but still
standing herbaceous plants. Each of these substrates tends
to be characterised by a distinct assemblage of species
(Stephenson 1988, 1989, 2011; Stephenson & Stempen
1994). The myxomycetes associated with the bark surface
of living trees, forest oor litter, and twigs are consistently
present and often abundant, but many of the species involved
are rather inconspicuous or sporadic in their occurrence
and thus dicult to detect in the eld. However, the moist
chamber culture technique as it applies to myxomycetes
(Gilbert & Martin 1933) provides a convenient and often very
productive method for studying these substrates. Since its
introduction, the technique has been used with considerable
success by numerous researchers (e.g. Härkönen 1977;
Blackwell & Gilbertson 1980; Stephenson 1989).
In late November and early December of 2016, forest
wildres occurred over portions of the Great Smoky
Mountains National Park. More than 4000 ha were aected
by these events. Some areas were subjected to intense res
that consumed most of the vegetation, whereas the res in
other areas were less intense. Recent studies carried out in
these areas show that microbial dynamics and functional
guilds were greatly impacted (Brown et al. 2019) and that
after two years, some microbial communities were in early
stages of their recovery process (Harpe et al. 2019). It is
well known that wildres are among the most impactful
disturbance factors in forest ecosystems (Ahlgren & Ahlgren
1960) and their negative impact on the ecological dynamics
of ground-level organisms is strong (Merino et al. 2018;
Bowd et al. 2019; Dove et al. 2020). Due to the fact that some
myxomycetes occur in association with substrates on the
ground, wildres can have a strong eect on the assemblage
of species present (see Adamonytė et al. 2016).
In that context, the present study was the result of two
circumstances. The general area impacted by such wildres
in the Great Smoky Mountains had been investigated
previously for myxomycetes and there are limited studies
showing the eect of wildres on these organisms (e.g.
Novozhilov et al. 2007; Gabel et al. 2010). The overall
Figure 1 – Infrared False Color Composite image (Sentinel Bands 2,3,8 from October 2018, courtesy of the European Space Agency-ESA)
showing the areas aected by the 2016 wildres around the city of Gatlinburg, TN. The location of the two study areas relative to the burned
sections of forest is shown. This type of image allows a clearer delimitation of burned vs non-burned areas based on the reectance signatures
of living vegetation in the infrared section of the electromagnetic spectrum.

17
Stephenson et al., Myxomycetes aected by wildres
Figure 2 – Landscape-level (Landsat 8, courtesy of the U.S. Geological Survey) and ground-level images of the study areas selected for this
investigation. A. Pre-wildre satellite view of the general area obtained on 12 Nov. 2016. B. Post-wildre satellite view obtained on 21 Dec.
2016 showing the extension aected by re. C. View of the forest in Jun. 2019 in TC. D. View of the forest in Jun. 2019 in BCT. Photographs
by Will Kuhn.
objective of the project reported herein was to assess the
impact of wildre on the assemblages of myxomycetes
associated with the three dierent substrates in two areas
subjected to dierent re intensities. As such, this study
contributes to the knowledge of wildre eects on microbial
systems and their ecological recovery.
MATERIAL AND METHODS
The present study was carried out in the Great Smoky
Mountains National Park, a federally managed forest
encompassing an area of 2113 km2 in western North Carolina
and eastern Tennessee in the United States. The park is
located at 35°36ʹN, 83°29ʹW, with elevations ranging from
approximately 270 to 2000 m above sea level.
Based on satellite imagery (Sentinel-2, courtesy of the
European Space Agency, ESA) as well as public information,
two study areas in the vicinity of Gatlinburg, Tennessee,
were selected on the basis of re intensity during the 2016
events (gs 1, 2). A relatively lower intensity burn study area
(g. 2C) was designated on a slope with a west-northwest
aspect at the Twin Creeks All Taxa Biodiversity Inventory
site (35°41ʹ11ʺN, 83°29ʹ60ʺW, 595 m; referred to as
TC), and a relatively higher intensity burn study area (g.
2D) was selected on a slope with a west-southwest aspect
along the Baskins Creek Trail (35°40ʹ45ʺN, 83°28ʹ38ʺW,
865 m; referred to as BCT). Prior to the wildres, TC was
characterised by a second-growth mixed hardwood cove
forest with a great laurel (Rhododendron maximum L.)
understory and BCT encompassed a second-growth mixed
oak-hardwood forest. Both areas have been under natural
regeneration after the wildres.
Thirty months after the wildres, on 28 Jun. 2019,
samples of twigs (T), forest oor leaf litter (GL), and
bark (B) were collected at regular intervals along walking
transects that extended through the two study areas being
considered. Those substrates would have been depleted
during the wildres in both study areas, and the assemblages
of myxomycetes associated with them at the time of sampling
were considered to be the result of new colonisation. In this
manner, ten samples were collected from each of the three
types of substrates in each of the two study areas, for a total
of 60 samples. Each sample was placed in an individual
paper bag and the latter numbered and marked with the type
of substrate. The entire set of samples was returned to the
Eumycetozoan Laboratory at the University of Arkansas for
processing.

18
Pl. Ecol. Evol. 154 (1), 2021
At the University of Arkansas, the samples were used
to prepare moist chamber cultures. These were prepared in
the manner described by Stephenson & Stempen (1994) and
consisted of plastic disposable Petri dishes (90 mm diameter)
lined with lter paper. Three Petri dishes were prepared from
each sample, for a total of 30 for each type of substrate, 90
for each study area, and a total of 180 for the entire project.
Enough sample material was placed in each dish to cover
most of the bottom, and the material was moistened with
distilled water. After a period of approximately 24 hours, the
pH of each culture was determined with a portable pH meter
and then excess water in the Petri dish was poured o. Moist
chamber cultures were placed out of direct sunlight and
maintained at room temperature. Water was added to these
cultures when necessary to maintain moist conditions, and the
cultures were checked at least twice each week for evidence
(either plasmodia or fruiting bodies) of myxomycetes over a
period of approximately three months. When fruiting bodies
were observed, they were recorded, removed from the moist
chamber culture, air-dried, and placed in small pasteboard
boxes for long-term storage and identication. A specimen
was dened as a record of the occurrence of one or more
fruiting bodies of a particular species of myxomycete in a
single culture. Some specimens consisted of only a single
fruiting body, whereas others consisted of numerous fruiting
bodies. With the information obtained from the moist
chambers, a database of species, records, and pH values was
created for the three dierent substrates at the two study
areas.
Using Landsat 8 images (courtesy of the U.S. Geological
Survey) from each season from winter 2016 (the time of the
wildres) to the summer of 2019 (the time of sampling), a
forest recovery progression was recreated for 20 hectares
associated with each of the two study areas. For this, the Soil
Adjusted Vegetation Index (SAVI), an estimator of vegetation
health similar to the NDVI but with a correction based on
soil variables (Huete 1988), was used. This estimator is
useful to analyze normal seasonal vegetation dynamics as
well as abnormal events and is based on the formula:
SAVI =
(1 + L) (NIR − Red)
(NIR + Red + L)
where NIR is the spectral data between 850–880 nm, Red
represents the spectral data between 640–670 nm, and L is
a canopy background adjustment factor, for which a value of
0.5 was used herein.
SAVI values oscillate between -1 and 1. Negative values
are associated with lack of vegetation, heavy disturbance,
or non-forested areas; values close to 0 are associated with
bare soil and disturbance; and positive values are associated
with vegetation cover and biomass presence, even during
the winter. Values closer to 1 are interpreted as complex and
healthy vegetation cover, usually associated with forested
areas during the peak of the growing season.
In a similar manner, for the winter seasons of 2016,
2017, and 2018, a calculation of land surface temperature
for a 20-hectare polygon was carried out using Landsat
8 information. This approach was used to analyse if the
wildres had aected the temperature values of the study
areas during 2016 (immediately post-re) more drastically
than normal oscillations in relation with non-burned areas of
the general landscape. Based on the work of Adamonytė et
al. (2016), more intense res (that can be determined on the
basis of temperature) have a stronger impact on myxomycete
communities. The calculations of land surface temperature
are complex and were based on a scheme similar to that of
Avdan & Jovanovska (2016) using Band 10 for the thermal
infrared spectral analysis.
With the myxomycete moist chamber data, the number
of species and records as well as the Shannon’s Index
of Diversity and Evenness values were calculated for
each substrate in each of the study areas. Also, the Chao1
estimator associated with the maximum number of species
expected with the methods used in the present study was
calculated using the formula:
Chao1 = a +
b2
2c
where a is the total number of species in the dataset, b is
the number of singletons (species with a single occurrence
in the sample), and c the number of doubletons (species with
double occurrences in the sample).
Also, the assemblages of species recorded from the
dierent substrates and the two study areas were compared
by pairwise calculation of the coecient of community (also
known as Bray-Curtis Index) using the formula shown below
(Mueller-Dombois & Ellenberg 1974):
Coecient of community (CC) =
2d
e + f
where e represents the total number of species present in
the rst study area or dataset being considered, f represents
the total number of species in the second study area or
dataset being considered, and d represents the common
species present between two study areas or datasets being
considered. The value of CC ranges from 0 to 1, where 0
indicates no common species shared between two study
areas or datasets and 1 indicates that all species are common
to both study areas or datasets.
In addition, all datasets recorded herein were compared
with previous information using a species assemblage
approach. For this, a pre-wildre dataset of myxomycete
records for the Great Smoky Mountains National Park
collected as part of the All Taxa Biodiversity Inventory
(ATBI) project, was used. This task was carried out only
with the assemblages of species that were associated with the
three substrates studied as part of the present investigation.
For this, a cluster analysis using the Bray-Curtis algorithm
was performed and both the numbers of unique and common
species were calculated for each branching. This approach
was used to determine the relative complexity of species
assemblages associated with the three substrates and the two
study areas.

19
Stephenson et al., Myxomycetes aected by wildres
RESULTS
A total of 38 species representing 17 genera were recorded
from the 226 specimens appearing in the 180 moist
chamber cultures prepared in the present study (table 1,
supplementary le 1). The vast majority (> 97%) of all
specimens could be identied to species, but a few were
poorly developed or aberrant and could be referred only to
genus. The only exception was one specimen of Trichia that
was well developed but could not be placed in any species of
which the senior author is aware and thus will need further
examination. It was left as Trichia sp. in table 1. Myxomycete
data previously obtained in the general area showed about
99 species on the same substrates studied herein. With such
gure, the present study recorded about 40% of the known
biodiversity in the area. Interestingly, Didymium ochroideum
G.Lister, Perichaena pedata (Lister & G.Lister) G.Lister ex
E.Jahn, and Physarum auriscalpium Cooke had not yet been
recorded.
The most diverse genus was Physarum, with seven
species. The single most common species was Arcyria cinerea
(Bull.) Pers., which was represented by 46 specimens. It was
the only species recorded from all three substrates in both of
the study areas. Historical data (using information generated
by the rst author during the period 1999–2003 with support
from Discover Life in America and the All Taxa Biodiversity
Inventory project) showed that Didymium melanospermum
(Pers.) T.Macbr., Leocarpus fragilis (Dicks.) Rostaf., and
Physarum viride (Bull.) Pers. are fairly common on all three
substrates, but the results from the present study were not
consistent with that observation. The next most common
species was Echinostelium minutum de Bary (27 specimens),
followed by Stemonitis fusca var. nigrescens (Rex) Torrend
(22), Perichaena vermicularis (Schwein.) Rostaf. (16), and
Perichaena chrysosperma (Curr.) Lister (14). The numbers
of species associated with the three dierent substrates
ranged from only six (litter for BCT) to 20 (bark for TC).
In BCT, bark was also the most productive substrate, with
14 species. Historical data showed that ground litter was
the substrate with the highest number of both species and
records in the area. When the totals for the same type of
substrate in the two study areas were compared, the most
appreciable dierence existed for litter. Sixteen species were
recorded for TC, which was appreciably higher than the total
(six, as noted above) for BCT. These results showed that,
in comparison with the 67 historically recorded species on
litter, only 28% of them were recorded in the present study.
The coecient of community indices calculated for
the possible pairwise comparisons of the three dierent
substrates, based on pooled data from both study areas,
were 0.37 (bark and litter, 6 common species), 0.49 (bark
and twigs, 8 common species), and 0.47 (litter and twigs, 10
common species). These values suggest that the assemblages
of myxomycetes associated with bark and litter are the least
similar, whereas those associated with bark and twigs are
the most similar. With this index, litter was the most distinct
substrate. However, these patterns were not necessarily
reected in the values obtained for pairwise comparisons
calculated for substrates in just one of the two study areas.
For TC, the values were 0.39 (bark and twigs, 5 common
species), 0.42 (bark and litter, 6 common species), and 0.38
(litter and twigs, 8 common species), whereas for BCT they
were 0.32 (litter and twigs, 3 common species), 0.29 (bark
and litter, 3 common species), and 0.50 (bark and twigs, 6
common species).
Figure 3 – Violin charts and boxplots of the pH value proles recorded in the study sites (A) and substrates (B) evaluated in the present
investigation.

20
Pl. Ecol. Evol. 154 (1), 2021
Myxomycete species Historical data Baskins Creek Trail Twin Creeks
BL T BL T BL T
Arcyria anis Rostaf. 2
Arcyria cinerea (Bull.) Pers. 17 70 5 5 1 13 7 1 19
Arcyria denudata (L.) Wettst. 1 2
Arcyria major (G.Lister) Ing 1
Arcyria pomiformis (Leers) Rostaf. 1 1
Arcyria stipata (Schwein.) Lister 3
Badhamia anis Rostaf. 1 1
Badhamia goniospora Meyl. 1
Badhamia macrocarpa (Ces.) Rostaf. 1
Badhamia papaveracea Berk. & Ravenel 1
Badhamia versicolor Lister 1
Badhamiopsis ainoae (Yamash.) T.E.Brooks & H.W.Keller 3
Ceratiomyxa fruticulosa (O.F.Müll.) T.Macbr. 2
Clastoderma debaryanum A.Blytt 9 3 1 1
Collaria arcyrionema (Rostaf.) Nann.-Bremek. ex Lado 1 2 1
Collaria lurida (Lister) Nann.-Bremek. 9
Colloderma oculatum (C.Lippert) G.Lister 3 3
Comatricha elegans (Racib.) G.Lister 1 1
Comatricha laxa Rostaf. 1 1 1
Comatricha nigra (Pers. ex J.F.Gmel.) J.Schröt. 8 4 2
Comatricha pulchella (C.Bab.) Rostaf. 1 2 2
Comatricha sp. 1 1
Comatricha tenerrima (M.A.Curtis) G.Lister 3
Craterium minutum (Leers) Fr. 3 2 1
Craterium obovatum Peck 1
Cribraria cancellata (Batsch) Nann.-Bremek. 1
Cribraria confusa Nann.-Bremek. & Y.Yamam. 4
Cribraria intricata Schrad. 3 4
Cribraria microcarpa (Schrad.) Pers. 3 2 1 3
Cribraria purpurea Schrad. 1
Cribraria rufa (Roth) Rostaf. 2
Cribraria violacea Rex 1 4
Diachea subsessilis Peck 1
Diacheopsis insessa (G.Lister) Ing 1
Dictydiaethalium plumbeum (Schumach.) Rostaf. 4
Diderma eusum (Schwein.) Morgan 111 1 2 2 1
Diderma hemisphaericum (Bull.) Hornem. 2 1 1
Diderma roanense (Rex) T.Macbr. 1 6
Diderma spumarioides (Fr. & Palmquist) Fr. 2
Diderma testaceum (Schrad.) Pers. 12 3
Didymium anellus Morgan 4
Didymium clavus (Alb. & Schwein.) Rabenh. 1
Didymium diorme (Pers.) Gray 7
Table 1 – Occurrence of myxomycetes in the three dierent substrates in the two dierent study areas. Also provided is a summary of values
(with the highest shown in bold) obtained for the assemblage of species associated with each substrate.

21
Stephenson et al., Myxomycetes aected by wildres
Myxomycete species Historical data Baskins Creek Trail Twin Creeks
BL T BL T BL T
Didymium iridis (Ditmar) Fr. 1 7
Didymium melanospermum (Pers.) T.Macbr. 15 18 4
Didymium minus (Lister) Morgan 2
Didymium nigripes (Link) Fr. 4
Didymium ochroideum G.Lister 1
Didymium squamulosum (Alb. & Schwein.) Fr. & Palmquist 1 4
Echinostelium minutum de Bary 5 3 8 7 4 8
Enerthenema papillatum (Pers.) Rostaf. 3
Fuligo septica (L.) F.H.Wigg. 6 4
Hemitrichia calyculata (Speg.) M.L.Farr 5 4
Hemitrichia minor G.Lister 2 1
Hemitrichia serpula (Scop.) Rostaf. ex Lister 2 6 1
Lamproderma columbinum (Pers.) Rostaf. 3
Lamproderma scintillans (Berk. & Broome) Morgan 1
Leocarpus fragilis (Dicks.) Rostaf. 2 2 5
Lepidoderma tigrinum (Schrad.) Rostaf. 3
Licea minima Fr. 4 1 3 1 1 1 1
Licea operculata (Wingate) G.W.Martin 6 1 3
Licea parasitica (Zukal) G.W.Martin 1
Licea pedicellata (H.C.Gilbert) H.C.Gilbert 1
Licea sp. 1
Lycogala epidendrum (L.) Fr. 3 5 2
Metatrichia oriformis (Schwein.) Nann.-Bremek. 1
Metatrichia vesparia (Batsch) Nann.-Bremek.
ex G.W.Martin & Alexop. 6 3
Perichaena chrysosperma (Curr.) Lister 9 3 3 6 1 1
Perichaena depressa Lib. 6 1
Perichaena pedata (Lister & G.Lister) G.Lister ex E.Jahn 4 1
Perichaena sp. 1 1
Perichaena vermicularis (Schwein.) Rostaf. 5 6 3 1 6
Physarum album (Bull.) Chevall. 21 9 1
Physarum auriscalpium Cooke 1 1
Physarum bethelii T.Macbr. ex G.Lister 1
Physarum bivalve Pers. 16 1 2
Physarum cinereum (Batsch) Pers. 4 2
Physarum compressum Alb. & Schwein. 5
Physarum contextum (Pers.) Pers. 1
Physarum crateriforme Petch 3
Physarum decipiens M.A.Curtis 1 1
Physarum galbeum Wingate 2
Physarum cf. hongkongense Chao H.Chung 1
Physarum leucophaeum Fr. & Palmquist 1 1
Physarum nucleatum Rex 1 1
Table 1 (continued) – Occurrence of myxomycetes in the three dierent substrates in the two dierent study areas. Also provided is a
summary of values (with the highest shown in bold) obtained for the assemblage of species associated with each substrate.

22
Pl. Ecol. Evol. 154 (1), 2021
Table 1 (continued) – Occurrence of myxomycetes in the three dierent substrates in the two dierent study areas. Also provided is a
summary of values (with the highest shown in bold) obtained for the assemblage of species associated with each substrate.
Myxomycete species Historical data Baskins Creek Trail Twin Creeks
BL T BL T BL T
Physarum penetrale Rex 1
Physarum psittacinum Ditmar 1
Physarum pusillum (Berk. & M.A.Curtis) G.Lister 2 1
Physarum viride (Bull.) Pers. 12 9 3 1 1 2
Stemonitis axifera (Bull.) T.Macbr. 2 4
Stemonitis avogenita E.Jahn 1 3
Stemonitis fusca var. nigrescens (Rex) Torrend 3 2 4 9 9
Stemonitis herbatica Peck 1
Stemonitopsis hyperopta (Meyl.) Nann.-Bremek. 2 2 1
Stemonitopsis typhina (F.H.Wigg.) Nann.-Bremek. 1
Symphytocarpus herbaticus Ing 1
Trichia botrytis (J.F.Gmel.) Pers. 1 1 1
Trichia contorta (Ditmar) Rostaf. 1
Trichia decipiens (Pers.) T.Macbr. 2 3
Trichia erecta Rex 4 2
Trichia favoginea (Batsch) Pers. 8 15 1 2 1
Trichia munda (Lister) Meyl. 1 1
Trichia persimilis P.Karst. 3
Trichia sp. 1
Trichia subfusca Rex 3 4 1 4
Tubifera ferruginosa (Batsch) J.F.Gmel. 3 1
Willkommlangea reticulata (Alb. & Schwein.) Kuntze 3 2
Number of species 58 67 15 14 6 13 20 16 16
Number of records 198 340 31 41 7 45 50 26 57
Chao1 Maximum number of species 96 77 60 17 11 34 23 34 25
Shannon Index of Diversity 3.6 3.5 2.4 2.4 1.8 2.1 2.8 2.6 2.1
Shannon Evenness 0.6 0.5 0.7 0.8 0.9 0.6 0.8 0.8 0.5
Many of the species represented by three or more
specimens displayed a preference for one type of substrate.
For example, 18 of the 22 records of Stemonitis fusca var.
nigrescens were from twigs, whereas all ve records of
Willkommlangea reticulata (Alb. & Schwein.) Kuntze were
from this substrate. Species displaying an apparent anity
for bark were Comatricha tenerrima (M.A.Curtis) G.Lister,
Cribraria violacea Rex, Licea minima Fr., L. operculata
(Wingate) G.W.Martin, and Stemonitis avogenita E.Jahn.
Only a very few species were represented by at least four
records on litter, but two of these (Perichaena pedata and
Trichia subfusca Rex) appeared to display an anity for
this substrate. In the historical records, there was not a clear
preference of species for twigs or bark, but Clastoderma
debaryanum A.Blytt, Cribraria microcarpa (Schrad.) Pers.,
Diderma hemisphaericum (Bull.) Hornem., D. testaceum
(Schrad.) Pers., Perichaena chrysosperma (Curr.) Lister,
P. depressa Lib., Physarum bivalve Pers., Ph. cinereum
(Batsch) Pers., and Perichaena vermicularis had only been
recorded on litter.
Collectively, 72% of all moist chamber cultures yielded
some evidence (either fruiting bodies or plasmodia) of
myxomycetes. Two or more species appearing in a single
culture was not unusual, and a few cultures yielded four or
more species. However, the set of cultures prepared with
samples from TC (84%) was more productive than the set
prepared with samples from BCT (59%). Moreover, the
former set of cultures yielded a total of 133 specimens,
whereas the latter produced only 93 specimens.
The median value of pH for all 180 moist chamber
cultures was 5.3, with a maximum value of 7.5 and a
minimum value of 3.9 (g. 3). Cultures prepared with twigs
were the most acidic (median = 5.1 with a range of 4.4 to
6.2), whereas those prepared with bark were the least acidic
(median = 5.9 with a range of 3.9 to 7.5). Values of pH also

23
Stephenson et al., Myxomycetes aected by wildres
diered between study areas, with a median value of 5.0
(range of 3.9 to 6.4) for BCT, and a median of 5.7 (range of
3.9 to 7.5) for TC.
Satellite imagery showed that both study areas were
aected by the wildres and that the rst growing season,
in the spring of 2017, was the most disturbed in terms of
deviation from normal dynamics (g. 4). The average SAVI
value for the springs of 2017 and 2018 was 0.48, and the
non-burned control plot during the spring of 2017 showed
an average value of 0.34. In contrast, the average value for
the burned study areas was -0.08, thus demonstrating the
strong eect of the wildres. Even the winter seasons of
2017 and 2018 showed a higher average value of 0.13. This
imagery also showed that the forest was in recovery during
the 30 months after the wildres, with a cover of vegetation
already present in the summer of 2017. Such recovery
has been more accelerated in TC than in BCT, which also
tended to have milder winter surface temperatures (see g.
5), presumably aecting myxomycete dynamics indirectly
by allowing vegetation to recover faster and modifying the
Figure 6 – Cluster analysis of similarity (Bray-Curtis algorithm)
showing the relationships among assemblages of myxomycetes
recorded on substrates (GL, T, and B) and in study sites (TC and
BCT). The historical records (HIST) obtained for the same general
area of the Great Smoky Mountains National Park for the same
substrates were included. The numbers of unique and shared species
are shown in the branches and the Shannon’s Index of Diversity as
well as Evenness values are shown next to each subdataset label.
Figure 5 – Maximum (Max), minimum (Min), and average (Avg)
temperature values (in °C) calculated for the winters of 2016–2018
from Landsat 8 images for 20 hectares associated with both study
areas (TC and BCT) as well as non-burned (NB) control areas. 2016
is abbreviated as Post, for being the immediate post-re season.
Figure 4 – Soil Adjusted Vegetation Index (SAVI) values for each
season and study area (TC and BCT) between the winter of 2016
and the summer of 2019. The values for TC and BCT during the
spring of 2017 were compared with a control non-burned (NB) area.
WI = winter, SP = spring, SU = summer, FA = fall.
microhabitat conditions. Interestingly, of the three winter
seasons evaluated, only during the winter of 2016, surface
temperatures associated with the two study areas were
signicantly dierent from those from non-burned areas.
This analysis also showed, as expected based on the reported
re intensity, that temperatures in BCT were higher than in
TC, demonstrating a stronger eect from the re.
Finally, when the species assemblages associated with
the three substrates and two study areas were compared with
previous data from the Great Smoky Mountains National
Park, the results did not show many dierences between
study areas (g. 6). All beta diversity values using the
Whittaker estimator were higher than 0.65. Albeit the latter,
approximately 65 species previously recorded in the area
were not observed in the present survey. Interestingly, clear
dierences in assemblage recovery were observed for ground
litter, with less complex assemblages (i.e. simpler in terms
of diversity and evenness) in comparison with historical
data. This observation also supports the results obtained
from the coecient of community comparison discussed
earlier. For litter, the remarkably high evenness values
demonstrated numerically the simplicity in the structure
of the assemblage in comparison with the other substrates,
instead of assemblage maturity. Both twigs and bark, in both
study areas, showed an intermediate stage of recovery in
relation to previous information. Taking the Shannon’s Index
of Diversity and Evenness values, it is also apparent that
historical data represented more complex and established
assemblages, as expected.
Annotated list of species
In the list that follows, species of myxomycetes recorded in
the present study were arranged alphabetically by genus and
then species. Information was provided on the number of
specimens recorded, the range of pH values of the culture(s)
in which the specimen appeared, and the substrate(s)

24
Pl. Ecol. Evol. 154 (1), 2021
with which records were associated. Numbers given in
parentheses are collecting numbers of the rst author.
Arcyria cinerea (Bull.) Pers.
Represented by 46 specimens (including 33761, 33808, and
34102) and associated with all three substrates in both study
areas. However, it was most common (32 specimens) on
twigs. The cultures in which this species appeared ranged in
pH from 3.9 to 7.1.
Clastoderma debaryanum A.Blytt
Represented by ve specimens (including 33960 and
33962) and recorded from both study areas. The majority of
specimens (four) were recorded on bark (pH 5.4 to 6.1).
Collaria arcyrionema (Rostaf.) Nann.-Bremek. ex Lado
Represented by three specimens (including 34008 and
34010), all of which were recorded on bark (pH 5.7 to 6.7).
This species was recorded from both study areas.
Comatricha elegans (Racib.) G.Lister
Represented by a single specimen (33770) recorded on bark
(pH 6.4) from BCT.
Comatricha nigra (Pers. ex J.F.Gmel.) J.Schröt.
Represented by two specimens (including 33771), both on
bark (pH 7.3 and 7.5) from TC.
Comatricha pulchella (C.Bab.) Rostaf.
Represented by four specimens (including 33840 and 33988),
all on bark (pH 5.0 to 5.9). Two specimens were recorded
from both BCT and TC.
Comatricha tenerrima (M.A.Curtis) G.Lister
Represented by three specimens (34131 and 34194) on bark
(pH 7.1 and 6.9), both from TC.
Cribraria microcarpa (Schrad.) Pers.
Represented by six specimens (including 33812 and 33856),
three on bark (pH 4.1 to 5.7), two on litter (pH 4.1 and 5.0),
and one on twigs (pH 5.1). This species was recorded from
both study areas.
Cribraria violacea Rex
Represented by four specimens (including 33809 and
34137), all on bark (pH 6.1 to 6.7) from TC.
Diderma eusum (Schwein.) Morgan
Represented by six specimens (including 33762 and 33939),
with three on litter (pH 5.8 to 6.6), two on bark (pH 5.7 and
6.7) and one on twigs (pH 5.6). This species was recorded
from both study areas. It is typically associated with litter
(Martin & Alexopoulos 1969).
Diderma hemisphaericum (Bull.) Hornem.
Represented by two specimens, the rst (33946) recorded
on litter (pH 6.1) and the second on bark (pH 5.0). Both
specimens were recorded from TC.
Diderma testaceum (Schrad.) Pers.
Represented by three specimens (34040 and 34041) on litter
(pH 5.9 for both) from TC.
Didymium ochroideum G.Lister
Represented by a single specimen (33963) recorded on litter
(pH 6.6) from TC.
Echinostelium minutum de Bary
Represented by 27 specimens (including 33769, 33855 and
33954), with 15 recorded on twigs (pH 4.4 to 5.9) and 12
recorded on bark (pH 3.9 to 5.9).
Hemitrichia minor G.Lister
Represented by a single specimen (34180) on bark (pH
6.7) from TC. In earlier taxonomic treatments of the
myxomycetes, this species was listed as Perichaena minor
(G.Lister) Hagelst.
Hemitrichia serpula (Scop.) Rostaf. ex Lister
Represented by a single specimen (34211) on twigs (pH
4.9) from BCT. This species is only occasionally recorded
from moist chamber cultures, but the specimen on which this
record is based was well-developed.
Lamproderma scintillans (Berk. & Broome) Morgan
Represented by a single specimen (observed but not
collected) recorded on bark (pH 5.9) from BCT.
Licea minima Fr.
Represented by seven specimens (including 33841 and
34228), recorded from both study areas. Four specimens
were recorded on bark (pH 4.1 and 5.8), one on twigs (pH
5.0 and 5.2) and two on litter (pH 4.8). Litter is an unusual
substrate for this species.
Licea operculata (Wingate) G.W.Martin
Represented by three specimens (including 33846 and
34095), all recorded on bark (pH 5.0 to 5.1) from TC.
Lycogala epidendrum (L.) Fr.
Represented by two specimens (including 33848) recorded
on bark (both pH 5.0) from TC. This species is very rarely
recorded from moist chamber cultures. The two aethalia
obtained in the present study were unusually small but
otherwise typical in overall morphology.
Perichaena chrysosperma (Curr.) Lister
Represented by 14 specimens (including 33961, 34109, and
34117) recorded on bark (pH 5.6 to 7.0), litter (pH 6.1), and
twigs (pH 5.2 to 5.6) from both study areas.

25
Stephenson et al., Myxomycetes aected by wildres
Perichaena depressa Lib.
Represented by a single specimen (34278) consisting of
only three sporocarps on bark (pH 5.8) from BCT. This is
a distinctive species because of the consciously attened
sporocarps.
Perichaena pedata (Lister & G.Lister) G.Lister ex E.Jahn
Represented by ve specimens (including 34097 and 34108),
with four on litter (pH 5.9 to 6.1) on one on twigs (pH 5.2)
from TC.
Perichaena vermicularis (Schwein.) Rostaf.
Represented by 16 specimens (including 33814 and 33942),
with six from bark (pH 5.1 to 6.4), nive from twigs (pH 5.2 to
6.2), and one from litter (pH 5.9). This species was recorded
from both study areas.
Physarum album (Bull.) Chevall.
Represented by a single specimen (34229) on twigs (pH
5.4) from TC. This specimen consisted of a single but well-
developed fruiting body.
Physarum auriscalpium Cooke
Represented by two specimens (34207 and 34233), one from
a twig (pH 5.0) from BCT and one from litter (pH 6.1) from
TC.
Physarum bivalve Pers.
Represented by three specimens (34138 and 34139), both
from TC. One was on litter (pH 5.9) and the other two were
from twigs (pH 5.2).
Physarum cinereum (Batsch) Pers.
Represented by two specimens (34091 and 34192) recorded
on litter (pH 6.0 and 6.4) from TC. This species typically
occurs in large fruiting, but the specimens recorded in the
present study consisted of only a few sporocarps.
Physarum cf. hongkongense Chao H.Chung
Represented by a single specimen (34321) on twigs (pH 5.4)
from TC. This species was described originally from Asia but
has since been reported from scattered localities throughout
the world. It is not common.
Physarum nucleatum Rex
Represented by two specimens (34007 and 34132), with one
on twigs (pH 5.2) and one on bark (pH 5.7).
Physarum viride (Bull.) Pers.
Represented by four specimens (including 34302), with three
on twigs (pH 4.9 and 5.2) and one (pH 5.9) on bark. This
species was recorded from both study areas.
Stemonitis avogenita E.Jahn
Represented by three specimens (including 33903 and
33941) on bark (pH 5.0 for all three) from TC.
Stemonitis fusca Roth
Represented by 22 specimens (including 33890 and 33948),
with 18 on twigs (pH 4.5 to 5.9) and four from bark (pH 5.7
to 5.9). This species was recovered from both study areas.
Specimens obtained from moist chamber cultures, as was
the case in the present study, appear to represent the variety
nigrescens, which is sometimes recognised as the distinct
species Stemonitis nigrescens Rex (Martin & Alexopoulos
1969).
Trichia botrytis (J.F.Gmel.) Pers.
Represented by two specimens (including 34107), both from
BCT. One was on litter (pH 4.2) and the other was on bark
(pH 4.6).
Trichia favoginea (Batsch) Pers.
Represented by four specimens (including 33780 and
34184), with two on bark (pH 5.6 and 5.7) and one each on
litter (pH 6.7) and twigs (pH 5.0). This species was recorded
from both study areas.
Trichia subfusca Rex
Represented by ve specimens, all recorded from TC. Four
(including 34090 and 34129) were on litter (pH 5.4 to 6.2)
and one was on bark (pH 52 and 6.7).
Trichia sp.
Represented by a single specimen (34197) on litter (pH 6.1)
from TC. This specimen could not be assigned to any species
of which the rst author is aware. However, it was clearly
dierent from the other species of Trichia obtained in the
present study.
Willkommlangea reticulata (Alb. & Schwein.) Kuntze
Represented by ve specimens (including 34053 and 34210),
all of which were recorded on twigs (pH 4.9 to 5.1) from both
study areas. It has been suggested (Stephenson et al. 2008)
that twigs represent the primary substrate for this species.
DISCUSSION
When a forest community is subjected to re, the rst thing
that burns is the litter layer (consisting mostly of dead leaves)
on the forest oor. Mixed in with this litter layer are woody
twigs that have fallen from trees. Both the litter layer and the
twigs can be consumed almost completely by surface res.
These same res can cause charring of the outer bark on the
lower trunks of trees in the forest. However, the outer bark
is already dead, and unless the tree is a species with very
thin bark, it is likely to survive. More intense res can kill
the tree and consume even relatively large pieces of coarse
woody debris (e.g. fallen logs).
Many of the woody twigs and probably all of the dead
leaves processed in the present study would be expected to
have fallen to the forest oor well after the res that took
place 30 months earlier. None of the samples of litter and
only a few of the samples of twigs displayed any evidence
that they had been subjected (at least directly) to re. This
was not the same for the samples of bark, many of which

26
Pl. Ecol. Evol. 154 (1), 2021
did show some evidence of charring. This was the case for
both the TC and BCT study areas and was expected based on
the clear strong eect of the wildres, as quantied through
satellite imagery.
There have been very few studies that have examined
the eects of re on myxomycetes. Novozhilov et al. (2007)
compared the assemblages of myxomycetes associated with
bark and litter in a series of burned and control (i.e. unburned)
sites in a study area located north of Fairbanks in central
Alaska. They reported that the numbers of species recorded
for these two substrates were fairly comparable. It should be
noted that no distinction was made between fallen leaves and
small woody twigs in this study. Gabel et al. (2010) collected
myxomycetes both before and after what they described
as a devastating re at a study site in South Dakota. They
recorded 13 species before the re and 14 species after the
re, with four species represented in both sets of collections.
Adamonytė et al. (2016) examined the eects of crown re
and surface re on the myxomycetes associated with pine
plantations in Lithuania. Their study extended over a period
of three years, and they reported that both types of re had a
major impact on the assemblages of myxomycetes present.
Both the assemblages present in plantations subjected to re
and those which were not proved to be equally diverse, but
they were quantitatively very dierent with respect to the
species of myxomycetes present. Interestingly, particular
species tended to switch to dierent substrates during the
course of post-wildre succession. The results obtained in
both of these studies suggest that myxomycetes associated
with particular localities recolonise the area rather quickly
after a re, but the structure of the assemblages present may
undergo appreciable changes. In the present study, such
detailed documentation was not possible because the exact
locations had not been examined with the same techniques
and eort prior to the res, but when data was compared with
historical records, results also suggest changes in species
assemblages.
It has been reported (Mataix-Solera et al. 2009) that an
intense re accompanied by high temperatures can result
in complete sterilisation of the soil microbiota. There is no
reason to suggest that the same would not be the case for the
microbiota (including myxomycetes) associated with litter,
woody twigs, and bark. In the same general study areas as
the present study, strong eects on that microbiota have been
reported (Brown et al. 2019), and at least based on surface
temperatures during the 2016 winter, signicant dierences
existed between the study areas and non-burned portions
of the forest. However, many species of myxomycetes are
known to have a high dispersal potential (Stephenson 2011),
so once substrates suitable for their growth and development
have become available again following the re, a new
assemblage of species would become established. This
certainly appears to be the case in the present study, since
the structure of the assemblages associated with the three
substrates is somewhat dierent from previous data. In
particular, it is remarkable that the number of shared species
among substrates/study areas was very low, suggesting that
such colonisation process was still very active.
The coecient of community values from both
study areas indicated that the assemblages of species of
myxomycetes associated with bark and litter were the least
similar, whereas the assemblages associated with litter
and twigs were the most similar. Based on what is known
about the ecological distribution of particular species of
myxomycetes, these results might have been anticipated.
The species associated with bark (which are commonly
referred to as corticolous species) tend to be dierent than
those associated with litter, based on the results obtained
in numerous studies (e.g. Stephenson 1989). Moreover, the
litter substrate and the twig substrate occur together on the
forest oor, so some overlap in species might be expected.
At least some of the overlap in species composition that
occurred between bark and twigs is almost surely the result
of the fact that most twigs still had their bark (albeit very
thin) still intact. However, it is interesting that ground litter
showed the most distinct (and simple) species assemblage
in the present study but the most complex assemblage in
the historical data. This result suggests a stronger eect
of the re on that substrate, in comparison with twigs and
bark. However, it should be noted that the historical records
considered herein were obtained in the larger region of the
Great Smoky Mountains National Park and are simply a
point of reference for analysis.
It has long been known that pH is a major factor
contributing to the distribution of myxomycetes in nature
(Stephenson 2011). However, pH did not appear to be
important for the three types of substrates examined, since
there was a high degree of overall similarity in the values
recorded for the sets of cultures. Nevertheless, the wide
range of values (3.9 to 7.5 when all cultures are considered)
probably has some impact on the occurrence of certain
species. The dierence in pH values between study areas
was interesting, showing more acidic substrates in BCT
than in TC. Although these were not signicant dierences,
they may have accounted for some dierences in species
composition between the two sites. For instance, of the 13
recorded species within the genera Physarum, Diderma, and
Didymium, all of which produce fruiting bodies with lime
(calcium carbonate) present, 62% were not recorded from
BCT. Records from these genera at TC were observed at an
average pH of 5.9 in comparison with a value of 5.3 for BCT.
In summary, the wildres that took place in 2016
appear to have had a strong eect on the assemblages of
myxomycetes associated with the three substrates in the
two study areas, since several levels of ecological analysis
diered from previous data. This eect seems to have been
less pronounced in TC than in BCT, as expected. As such,
our data suggest that the relatively higher intensity burn had
more of an eect on myxomycetes than the relatively lower
intensity burn. Remarkably, even though the assemblages of
species present have largely recovered 30 months after the
two study areas were subjected to burning, such recovery
is still an ongoing process. It is likely that the assemblages
of species associated with substrates in the areas aected
by the wildres were dierent from those found before this
event. Historical data pointed in that direction. It would seem
worthwhile to carry out future studies in the same study areas
in order to follow up on the present investigation.

27
Stephenson et al., Myxomycetes aected by wildres
SUPPLEMENTARY FILE
Supplementary le 1 – Complete list of myxomycete
specimens recorded in the present study along with
information on substrate, pH values, and the study site where
they were observed.
https://doi.org/10.5091/plecevo.2021.1762.2355
ACKNOWLEDGEMENTS
The research reported herein was supported in part by the
Slime Mold Project at the University of Arkansas and a grant
from Discover Life in America. Appreciation is extended
to Hana Hess for her assistance in collecting the samples.
Technical support was received from University of Costa
Rica through project 731-B7-721.
REFERENCES
Adamonytė G., Motiejūnaitė J. & Iršėnaitė R. 2016. Crown re
and surface re: eects on myxomycetes inhabiting pine
plantations. Science of the Total Environment 572: 1431–1439.
https://doi.org/10.1016/j.scitotenv.2016.02.160
Ahlgren I.F. & Ahlgren C.E. 1960. Ecological eects of forest res.
The Botanical Review 26: 483–533.
https://doi.org/10.1007/BF02940573
Avdan U. & Jovanovska G. 2016. Algorithm for automated mapping
of land surface temperature using LANDSAT 8 satellite data.
Journal of Sensors 2016: 1480307.
https://doi.org/10.1155/2016/1480307
Blackwell M. & Gilbertson M.L. 1980. Sonoran Desert
myxomycetes. Mycotaxon 11: 139–149.
Bowd E.J., Banks S.C., Strong C.L. & Lindenmeyer D. 2019. Long-
term impacts of wildre and logging on forest soils. Nature
Geosciences 12: 113–118.
https://doi.org/10.1038/s41561-018-0294-2
Brown S.P., Veach A.M., Horton J.L., Ford E., Jumpponen A. &
Baird R. 2019. Context dependent fungal and bacterial soil
community shifts in response to recent wildres in the Southern
Appalachian Mountains. Forest Ecology and Management 451:
117520. https://doi.org/10.1016/j.foreco.2019.117520
Dove N.C., Saord H.D., Bohlman G.N., Estes B.L. & Hart S.C.
2020. High‐severity wildre leads to multi‐decadal impacts
on soil biogeochemistry in mixed‐conifer forests. Ecological
Applications 30(4): e02072. https://doi.org/10.1002/eap.2072
Gabel A., Ebbert E., Gabel M. & Zierer L. 2010. A survey of
myxomycetes from the Black Hills of South Dakota and the
Bear Lodge Mountains of Wyoming. Proceedings of the South
Dakota Academy of Science 89: 45–67.
Gilbert H.C. & Martin G.W. 1933. Myxomycetes found on the bark
of living trees. University of Iowa Studies in Natural History
15: 3–8.
Härkönen M. 1977. Corticolous myxomycetes in three dierent
habitats in southern Finland. Karstenia 17(1): 19–32.
https://doi.org/10.29203/ka.1977.121
Harpe V.R., Moorhead L.C., Moore J.A.M. & Kivlin S.N. 2019.
Fungal response to wildre in southeastern forests: eects at
the urban-forest interface. Middle Atlantic States Mycological
Conference 2019. Available from
https://trace.tennessee.edu/masmc/14 [accessed 19 May 2020].
Huete A.R. 1988. A soil-adjusted vegetation index (SAVI). Remote
Sensing of Environment 25(3): 295–309.
https://doi.org/10.1016/0034-4257(88)90106-X
Lado C. 2005–2020. An online nomenclatural information system
of Eumycetozoa. Real Jardín Botánico, CSIC. Madrid, Spain.
Available from https://eumycetozoa.com [accessed 16 Sep.
2020].
Martin G.W. & Alexopoulos C.J. 1969. The myxomycetes.
University of Iowa Press, Iowa City.
Mataix-Solera J., Guerreno C., Garcia-Orenes F., Barcenas-Moreno
G. & Torres M.P. 2009. Forest re eects on soil microbiology.
In: Cerdá A. & Robichaud P.R. (eds) Fire eects on soils and
restoration strategies: 133–175. Science Publishers, Inc, Boca
Raton, Florida.
Merino A., Fonturbel M.T., Fernández C., Chávez-Vergara B.,
García-Oliva F. & Vega J.A. 2018. Inferring changes in soil
organic matter in post-wildre soil burn severity levels in a
temperate climate. Science of The Total Environment 627: 622–
632. https://doi.org/10.1016/j.scitotenv.2018.01.189.
Mueller-Dombois D. & Ellenberg H. 1974. Aims and methods of
vegetation ecology. John Wiley & Sons, New York.
Novozhilov Y.K., Stephenson, S.L., Overking M., Landolt J.C. &
Laursen G.A. 2007. Studies of Frostre myxomycetes including
the description of a new species of Diderma. Mycological
Progress 6: 45–51. https://doi.org/10.1007/s11557-007-0527-z
Stephenson S.L. 1988. Distribution and ecology of myxomycetes in
temperate forests. I. Patterns of occurrence in the upland forests
of southwestern Virginia. Canadian Journal of Botany 66(11):
2187–2207. https://doi.org/10.1139/b88-302
Stephenson S.L. 1989. Distribution and ecology of myxomycetes
in temperate forests. II. Patterns of occurrence on bark surface
of living trees, leaf litter, and dung. Mycologia 81(4): 608–621.
https://doi.org/10.1080/00275514.1989.12025792
Stephenson S.L. 2011. From morphological to molecular: studies
of myxomycetes since the publication of the Martin and
Alexopoulos monograph. Fungal Diversity 50: 21.
https://doi.org/10.1007/s13225-011-0113-1
Stephenson S.L. & Stempen H. 1994. Myxomycetes: a handbook of
slime molds. Timber Press, Portland, Oregon.
Stephenson S.L. & Landolt J.C. 2009. Mycetozoans of the Great
Smoky Mountains National Park: an All Taxa Biodiversity
Inventory project. Southeastern Naturalist 8(2): 317–324.
https://doi.org/10.1656/058.008.0210
Stephenson S.L., Schnittler M., Mitchell D.W. & Novozhilov Y.K.
2001. Myxomycetes of the Great Smoky Mountains National
Park. Mycotaxon 78: 1–15.
Stephenson S.L., Urban L.A., Rojas C. & McDonald M.S. 2008.
Myxomycetes associated with woody twigs. Revista Mexicana
de Micología 27: 21–28.
Communicating editor: Jérôme Degreef.
Submission date: 20 May 2020
Acceptance date: 16 Sep. 2020
Publication date: 23 Mar. 2021