Mycetozoan Biodiversity in the Great

Smoky Mountains National Park

Great Smoky Mountains National Park

Great Smoky MountainsNational Park

• National Park in 1934

• International Biosphere Reserve in 1976

• World Heritage Site in 1983

• ATBI in 1998

Great Smoky MountainsNational Park

• >200,000 ha in extent

• located in western North Carolina and eastern Tennessee

• elevation range of 270 m to 2000 m

• 95% of the Park is forested

• >130 species of trees

Major Types of Communities

• Spruce-fir forests (>1525 m)

• Northern hardwood forests (1065 to 1525 m)

• Pine-oak forests (<1065 m, xeric sites)

• Cove hardwood forests (<1065 m, mesic sites)

• Hemlock forests (generally along streams)

• Other types of communities

Old-Growth Forests

Members of the Slime Mold TWIG at Purchase Knob in August 2003

Participants in the ATBI Slime Mold TWIG

• Grazina Adamonyte• Denise Binion• Dawn Black• James Cavender• Randy Darrah• Uno Eliasson• Tom Gaither• Harold Keller• Tatyana Krivomaz• John Landolt

• Roland McHugh• David Mitchell• Yuri Novozhilov• Carlos Rojas• Adam Rollins• Wayne Rosing• Martin Schnittler• John Shadwick• Fred Spiegel• Ted Stampfer

Hemitrichia calyculata

The Mycetozoans Consist of Three Distinct Groups of Organisms

• Myxomycetes (or plasmodial slime molds)

• Dictyostelids (or cellular slime molds)

• Protostelids (or protostelid slime molds)

Myxomycetes[ca 875 species]

Dictyostelids[ca 120 species]

The myxomycetes are the largest and best known group,

and primary emphasis has been placed on collecting and

studying these organisms.

Primary Microhabitats

• Coarse woody debris

• Ground litter

• Bark surface of living trees

Other Microhabitats

• Dung of herbivores (e.g., deer)

• Soil (upper humus-rich layers)

• Aerial litter (dead but attached plant parts above the ground)

• Twigs and dead aerial wood

Sources of Records

• Field collections (fruiting bodies that developed under natural conditions)

• Moist chamber culture collections (fruiting bodies that developed under laboratory conditions)

Hemitrichia serpula

Moist chamber culture prepared with samples

of bark

Moist chamber culture prepared with samples

of leaf litter

Perichaena vermicularis

Licea biforis

Clastoderma debaryanum

Perichaena chrysosperma

Ecological Distribution

• Found throughout the Park• Generally restricted to hardwood

forests• Generally restricted to coniferous

forests• Limited to particular microhabitats

Arcyria cinerea

Physarum viride

. . . if considered to be a myxomycete!

Ceratiomyxa fruticulosa

Metatrichia vesparium

Enerthenema papillatum

Leocarpus fragilis

Worldwide distribution of Leocarpus fragilis

Southern AppalachianSpruce-fir Forests

• Great Smoky Mountains National Park

• Mount Rogers in southwestern Virginia

• Blister Run/Gaudineer Scenic Area in eastern central West Virginia

Field Collections

• More than 1200 total collections

• At least 89 species representing 34 genera

• Number of species recorded from a particular study site ranged from 63 (GSMNP) to 39 (Mount Rogers)

Ecological Distribution

• >95% wood or bark

• 3% forest floor litter

• 1% other (e.g., living plants)

• >30% associated with bryophytes

Most Common Species

• Licea minima

• Trichia decipiens

• Lepidoderma tigrinum

• Lycogala epidendrum

• Physarum viride

“Special” Species (spruce-fir)

• Barbeyella minutissima

• Lepidoderma tigrinum

• Colloderma oculatum

• Lamproderma columbinum

• Elaeomyxa cerifera

Barbeyella minutissima

Colloderma oculatum

Elaeomyxa cerifera

Lepidoderma tigrinum

Since the slime mold survey component of the ATBI began in 1998, the number of species of myxomycetes known from

the Park has increased from 88 to more than 220.

Recent new records (GSMNP & NA)

• Comatrichia pinicillata

• Lamproderma granulosum

• Licea microscopica

• L. rufocuprea

• L. sambucina

Stephenson et al. 2001, Snell et al. 2003

• Diachea arboricola A species new to science; Keller et al. 2004

Dictyostelium discoideum in the wild (on deer scat)

(photo courtesy of Thomas Platt, Rice University)

Dictyostelids are microscopic at virtually allstages of their life cycle, including the fruiting body.

Oval spores showing polar granule characteristic

A large aggregation

Migrating pseudoplasmodia

Stalked fruiting bodies developing

Fruiting bodies of several types of cellular slime molds

(from Am. J. Bot)

Typical leaf litter habitat

Remove Intact leaves

Scraping soil/litter

into sterile bag

10-50 g soil/litter

in a sample

bag

Weighing out soil sample for dilution

Surveys for Dictyostelids

• Sampling was carried out during the period of 1993 to 2004

• A total of 412 samples from study sites throughout the Park

• Study sites included examples of all major forest types along with the more common types of non-forest vegetation

• >2300 clones of dictyostelids recovered

Collection SitesHigh Elevation

Clingman’ Dome

Indian Gap

Andrews Bald

Double Springs

Balsam Mountain Mid Elevation

Purchase Knob* Snakeden Ridge

Ramsey Cascade

Chimneys Campground

Foothills Parkway-West Low Elevation

Little River Ravensford

Madron Bald Trail Oconoluftee

Rich Mountain Fontana Dam Twin Creeks Deep Creek

Gregory’s Cave Perimeter

Gregory’s Cave Interior

Foothills Parkway-East

Cades Cove

Tremont

CELLULAR SLIME MOLDS FROM THE GSMNP – Species names in black are reported prior to the current survey; names in red are additions made (mostly since 2001). Many of the added species came from “extremes” of elevation, soil dryness, soil acidity or within caves. Most unassigned forms in this table have now been formally described (Cavender et al. 2005)

1 Acytostelium leptosomum 13 Dictyostelium aureostipes 24 D. purpureum 2 A. subglobossum 14 D. aureostipes var. helvetium 25 D. septentrionalis 3 A. sp. A 15 D. discoideum 26 D. sphaerocephalum 4 A. sp. B 16 D. firmibasis ? 27 D. sp. A 5 A. sp. C 17 D. giganteum 28 D. sp. B 6 A. sp. D 18 D. implicatum 29 D. sp. C 7 A. sp. E 19 D. lacteum 30 D. sp. D 8 Polysphondylium candidum 20 D. "leptosomum" 31 D. sp. E 9 P. pallidum 21 D. minutum 32 D. sp. F

10 P. pseudocandidum 22 D. mucoroides 33 D. sp. G 11 P. tenuissimum 23 D. polycephalum 34 D. sp. H 12 P. violaceum

The following images were obtained using an auto-montage set-up which allows better visualization of the minute, usually pigment free fruiting bodies and other life stages.

Special thanks to Andy Swanson

A Guide to the Common Dictyostelid Slime Molds of Great Smoky Mountains National ParkAndrew R. SwansonDepartment of Biological SciencesUniversity of Arkansas

Dictyostelium mucoroides Brefeld syn. Dictyostelium brefeldianum Hagiwara

A) Sorocarps growing on a hay infusion isolation plate. B) migrating sorogens & young sorocarps (note stalked migration). C) mature sorocarps. D) tips (note morphological range from clavate to more typical capitate). E) spores (note lack of distinct polar granules. F) aggregation.

A

B C

DEF

Dictyostelium sphaerocephalum (Oud) Sacc. et March. syn. Hyalostilbum sphaerocephalum Oud.syn. Dictyostelium mucoroides Brefeld - sensu Hagiwara

A) Mature sorocarps (note ‘L’-shaped form agar surface). B) aggregations. C) spores (note lack of prominent polar granules). D) tip (note adherent material at apex and collar). E) mature sorocarps.

A

B

CDE

Dictyostelium discoideum Raper A) mature sorocarp (note discoid base). B) base. C) tip (note closely adherent spore mass). D) spores (note lack of

prominent polar granules). E) aggregation. F) migrating slug (note stalkless migration). G) detail, discoid base.

A B

C

G EF

D

Dictyostelium minutum Raper A) Colony (note size in comparison with P. pallidum). B. developing and mature sorocarps. C) mature sorocarps. D) tip and

spores. E) tip. F) spores (note polar granules – often not as prominent as pictured). G) aggregations.

A

B C

DEFG

Dictyostelium purpureum Olive A) mature sorocarps. B) mature sorocarp (note pigmented stalk and sori). C) lower stalk (note purple pigmentation). D)

rounded base. E) base with supporter. F) stalk tip. G) spores (note lack of prominent polar granules) . H) aggregation.

AB

C D

EFGH

Polysphondylium pallidum Olive A) mature sorocarps. B) mature sorocarp (note whorls). C) branch tip. D) spores (note characteristic unconsolidated polar

granules). E) aggregation. F) developing sorocarps.

AB

C

D

EF

Polysphondylium violaceum Brefeld A) mature sorocarps (note violet pigmentation). B) mature sorocarp. C) tip. D) branch tip & spores E) spores (note prominent

polar granules). F) aggregations.

BA

C

F DE

B

Dictyostelium aureo-stipes Cavender, Raper et Norberg A) developing sorocarps. B) mature sorocarps (note yellow pigmentation at branch confluence). C) matue sorocarps.

D) mature sorocarp (note irregular crowded branches). E) stalk and spores (note yellow pigmentation at stalk edges). F) tip and spores (note obtuse tip and consolidated polar spore granules). G) aggregations. H) rising sorogens.

A B

C

D

EFGH

Dictyostelium polycephalum RaperA) Aggregation and developing slug. B) migrating slug and developing sorocarp. C) rising slug (note size of nearby

sorocarp). D) developing sorocarp. E) mature sorocarp F) mature sorocarp. G) spores (note unconsolidated polar granules).

A

B

C

D

EF

G

Acytostelium subglobosum CavenderA) numerous developing and mature sorocarps. B) sorogens, sorocarps, and spore masses. C) mature sorocarps (note

development toward activated charcoal). D) base. E) tip. F) tip & spores. G) spores. H) aggregation, sorogens & spore masses.

A B C

DEF

GH

Ecological Assemblages of Dictyostelids

• Higher elevations, soil pH <5.0

• Lower elevations, soil pH >5.0

• Wide range of elevations and

soil pH conditions

Higher elevations and soil pH <5.0

• Dictyostelium discoideum

• Dictyostelium septentrionalis

• Polysphondylium tenuissimum

• Several newly described species

Lower elevations and soil pH >5.0

• Dictyostelium giganteum

• Dictyostelium lacteum

• Dictyostelium purpureum

• Dictyostelium minutum

Wide range of elevationsand soil conditions

• Dictyostelium mucoroides

• Dictyostelium aureostipes

• Polysphondylium pallidum

• Polysphondylium violaceum

Recently Described Dictyostelid Species from

GSMNP Collections

(Cavender, J. C. et al. 2005. Mycologia 97:493-512)

A. anastomosans FP5A

A. longisorophorum DB10A

A. magnisorum 08A

A. serpentarium SAB3A

A. singulare FDIB

D. amphisporum BM9A

D. naviculare SAB5A

D. oculum dB4B

D. potamoides FP1A

D.stellatum SAB7B

Dictyostelid General Summary

• Total number of species known from the Park has increased from 12 to >30.

• Overall species richness (18 to 20) is similar at all elevations (with some differences in assemblages).

• Dictyostelid abundance generally decreases with increasing elevation/decreasing pH.

• Many of the new records and new species are from high elevation/low pH/low plant diversity “marginal” habitats.

There were no reports of protostelids from the Park

prior to the ATBI.

Surveys for Protostelids• Sampling was carried out in October

2000 and November 2003• Study sites included examples of all

major forest types along with the more common types of non-forest vegetation

• Substrates examined included aerial litter, ground litter, bark of living trees, and coarse woody debris

Some of the collections of myxomycetes and dictyostelids from the Park are being used in studies to determine the “genetic relatedness” of collections of the same species from the same and different regions of the world.

Robert Hagelstein (“found in August [1939] near Newfound Gap, at an altitude of 4200 feet”)

Prototrichia metallica

Appreciation is extended to Discover Life in America for their support of mycetozoan research in the Great Smoky Mountains

National Park and also to the Shepherd University Foundation and Alumni

Association. We also thank a number of volunteers who assisted in sampling efforts.