Cumberland Mountain Naturalist Vol 1, Issue 1, March/April 2020

Checklist of the Freshwater Sponges

(Porifera: Spongillida) of Tennessee

John Copeland

2

Cumberland Mountain Naturalist Published by Lincoln Memorial University

Issued by the Cumberland Mountain Research Center

The Cumberland Mountain Naturalist is a quarterly journal devoted to publishing

articles of varied interests related to ecology, conservation, taxonomy, organismal

biology, as well as many other topics for a general audience. Each issue consists of

a single article focused on a single topic highlighting written by experts in the field.

All articles are selected through direct solicitation of the authors by the editorial

staff. However, if you are interested in publishing an article, please contact a mem-

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librarians, youth leaders, conservation organizations, and Lincoln Memorial Uni-

versity Alumni. If you are any of the above, to receive your complimentary, printed

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address, or indicate that you are an LMU alum with the year that you graduated, to

the Editor in Chief. Electronic versions are available free of charge to all others

worldwide via the Lincoln Memorial University Digital Commons (https://

digitalcommons.lmunet.edu/cumberland_mountain_naturalist/). The Cumberland

Mountain Naturalist is published by Lincoln Memorial University, Cumberland

Mountain Research Center within the Department of Biological Sciences. Address

print correspondence to Cumberland Mountain Naturalist, Department of Biology,

Lincoln Memorial University, 6965 Cumberland Gap Parkway Harrogate, TN

37752. All electronic correspondence can be addressed to the Editor in Chief. Pub-

lication of the Cumberland Mountain Naturalist is underwritten by the Cumberland

Mountain Research Center. If you would like to support this journal and its contin-

ued publication, please contact the Editor in Chief. Opinions and perspectives ex-

pressed in publication are those of the authors and/or editor(s) and do not reflect the

official position or endorsement of Lincoln Memorial University.

If you would like to receive notifications as new issues are published, please send your name and email address to the Editor in Chief. This issue authored by Dr. John E. Copeland is

Professor Emeritus of Biology at Lincoln Memorial

University (LMU). While actively teaching at LMU

he occupied the Louis Lutz Chair in Biology. He is

a twice recipient of the Houston Award for teaching

excellence. His education includes B.S. in Wildlife

Management and M.S. in Wildlife Biology from

Tennessee Technological University and a Ph.D. in

Animal Science from the University of Tennessee.

He is a certified wildlife biologist.

3

Inaugural Issue: Letter from the Editor

Dear Reader,

The release of this inaugural issue of the Cumberland Mountain Naturalist

marks a milestone for the Cumberland Mountain Research Center and the 40+ year

history of the Lincoln Memorial University Wildlife and Fisheries Management

program. This history is in no small part the result of the contributions and efforts

of Dr. John Copeland, Professor Emeritus Wildlife and Fisheries. Although Dr.

Copeland retired in 2016, it is his work to establish, build, and maintain the LMU

Wildlife and Fisheries Management Program that has laid the foundation from

which the Cumberland Mountain Naturalist is launched.

We owe a great debt to Dr. Copeland as he not only began our Wildlife and

Fisheries Program, but he has mentored many students into becoming successful

biologists in their own right. The success of graduates from the LMU Wildlife and

Fisheries program is a testament to Dr Copeland’s dedication to ensuring many,

many, LMU students were well trained to pursue careers within the discipline. Fur-

thermore, as the nature of wildlife biology has become more of landscape and adap-

tive system management, Dr. Copeland was instrumental in the maturation of the

LMU Wildlife and Fisheries program into our new Conservation Biology program

to promote a much broader approach to resource and ecosystem management.

During his 40 years of service to Lincoln Memorial University, Dr. Copeland

also played a crucial role in the creation of the Cumberland Mountain Research

Center (CMRC). Since its creation in 1990, the CMRC has been essential in

providing leadership, mentoring, and resources to facilitate wildlife, conservation,

ecological, and environmental research and educational activities. These efforts

continue today as our Conservation Biology program continues to grow.

With these things in mind, we felt that it most appropriate to have Dr. Copeland

as the author of the first issue of the Cumberland Mountain Naturalist. His contin-

ued, post-retirement work on freshwater sponges in Tennessee is breaking new

ground as he has not only developed the checklist of sponges published here, but in

his efforts he has discovered a new species of sponge. We certainly look forward to

following Dr. Copeland’s research activities as he continues to demonstrate to fac-

ulty and students, how to be a professional, a scientist, and a wildlife biologist

through all his endeavors. Thank you! Dr Copeland for all the you have done and

continue to do.

Sincerely,

LaRoy Brandt

Editor in Chief

Director, Cumberland Mountain Research Center

Checklist of the Freshwater Sponges (Porifera: Spongillida)

of Tennessee

by John E. Copeland

Introduction

Sponges are sessile multicellular animals lacking tissues and organs. They

function through the actions of specialized cells. Totipotent cells are stem cells

capable of changing morphology and physiology to become other types of cells.

Totipotent cells occur throughout the animal kingdom but are uniquely different in

sponges. The totipotent cells of other animals change in only one direction from

stem cell to the derived cell. In sponges these cellular changes can occur naturally

in both directions. Interestingly, human stem cells can be manipulated after differ-

entiation has occurred to convert back to the stem cell state; but this has not been

observed to occur naturally.

Sponges are an ancient and successful life form. They may have been the first

multicellular animal to appear on Earth. Fossil evidence indicates sponges were

present before the Cambrian explosion which occurred over 540 million years ago.

This explosion of new life forms happened over a relatively short period of time,

20 – 25 million years, and resulted in the

appearance of most of the major animal

phyla. Sponges have survived previous

mass extinctions and have adapted to liv-

ing in a variety of aquatic environments.

Freshwater sponges are members of

the benthic (bottom) community of fresh-

water ecosystems. They are typically

found living on hard substrates. In physi-

cal appearance they resemble cushions or

encrustations on the surface of substrates

but can be branched and finger-like. Col-

ors range from white to buff, yellow to

brown, gray to black, and some are green.

The green coloration results from a symbiotic relationship between sponges and

green algae (Figure 1). This situation is common and may be one of the reason

sponges were at one time thought to be plants.

Approximately 8,500 sponge species are known to exist. Freshwater species

account for about 3% of this total. Although sponge invasion of freshwater oc-

curred by the late Carboniferous Period only 6 freshwater families having less than

250 valid species are known worldwide. Currently there are 33 described species

of freshwater sponges in the Nearctic Biogeographical realm. Thirteen species have

Figure 1. Freshwater sponge having zoo-

chlorellae alga symbiont.

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have been documented in

Tennessee waters. Species of

three families, Metanidae,

Potamolepidae, and Spon-

gillidae, are known to occur

in the United States. Species

belonging to Potamolepidae

and Spongillidae, have been

collected from Tennessee

waters.

Sponges of the family

Potamolepidae are character-

ized by usually producing

sessile gemmules that are

adapted to persisting in situ

(remaining attached to their substrate) under extreme conditions such as low water

levels to harsh flooding and heavy silting. Such gemmules typically lack a pneu-

matic layer and are not well adapted for dispersal by water currents. (Figure 2).

Many sponges of the family Spongillidae are characterized by gemmules hav-

ing pneumatic layers. Gemmules having a pneumatic layer float in water and are

dispersed downstream by water currents. (Figure 3)

Importance of Gemmules:

In physical appearance gemmules are small spherical shaped structures which

have an opening called the foramen. Gemmules have two functional roles as resting

Figure 2. Cross section of a gemmule without pneu-

matic layer.

Figure 3. Cross section of gemmule. A, outer membrane; B, pneumatic layer; C, gem-

mulosclere; D, inner membrane; E, archaeocytes.

3

bodies, gemmules are survival structures which allow sponges to survive environ-

mental conditions detrimental to the adult sponge. Because they are resistant to

freezing, desiccation, and anoxia gemmules are produced in preparation for hiber-

nation or aestivation. (Figures 4 and 5).

As propagules, gemmules are asex-

ual reproductive structures that contain

cells known as archaeocytes and amoe-

bocytes. These cells are surrounded by

a protective coating of interlocking

spicules known as gemmuloscleres

(Figure 3.) Gemmule formation begins

when archaeocytes and nurse cells

know as trophocytes aggregate together

creating a mass. Archaeocytes obtain

polyols from trophocytes which are

used to form vitelline platelets. Archae-

ocytes filled with vitelline platelets be-

come binucleated thesocytes. Amoebocytes surround the mass of developing

thesocytes and secrete an inner and outer membrane composed of spongin. Sclero-

cytes secrete gemmuloscleres between the two membranes. When fully developed

gemmules are spherical to ovoid in shape and have an opening called the foramen.

When gemmules germinate, thesocytes are released through the foramen. Once

released thesocytes differentiate into the various cell types needed for the develop-

ment of a sponge. A single

gemmule has the potential to

give rise to several young

sponges.

Gemmules played a criti-

cal role in the evolution of

freshwater sponges. Freshwa-

ter sponges evolved from

marine ancestors. The inva-

sion and colonization of

freshwater environments pre-

sented difficult challenges for

marine sponges to overcome.

Differences in salinity being

one of them. The average

salt concentration of seawater

is about 3.5% (about 35g of

salt per liter) whereas average

Figure 4. Gemmule cluster in freshwater

sponge body.

Figure 5. SEM of gemmule of Radiospongilla crateri-

formis. A, outer membrane; B, foraminal aperture; C,

distal end of gemmulosclere.

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salinity concentration in freshwater is about 0.1%. This difference is huge and

forced sponges attempting to invade freshwater to overcome osmoregulation

(maintaining an internal balance between water and dissolved materials) problems.

Other barriers had to be solved. Marine environments are continuous with little

probability of becoming dry land. In freshwater environments, there is a much

higher probability of droughts causing low water levels or complete dry-up result-

ing in desiccation of aquatic organisms. Additionally, freshwater ecosystems have a

discontinuous spatial distribution which makes it more difficult for sponges to dis-

perse from one body of water to another. Gemmules proved to be the solution to

overcoming these problems.

Skeleton:

Freshwater sponges have a skeleton composed of protein fibers and siliceous

spicules. Collagen fibers,

known as spongin, form the

fibrous portion of the skele-

ton. These fibers function to

bind spicules together. Spic-

ules which are composed of

silica dioxide form the

ridged mineral portion of

the skeletal system. The

rigidity or stiffness of the

skeleton varies greatly

among species and is deter-

mined by the degree of in-

terlocking or fusion of spic-

ules and by the availability

of silica. Spicules provide

structural support to the

sponge body and protection

to totipotent cells located within gemmules. Additionally, it is thought spicules may

act to prevent or reduce predation. The beauty of a freshwater sponge can be seen

in its spicules. Some spicules are quite ornate in design.

There are three classes of spicules: megalscleres, microscleres and gemmulo-

scleres (Figure 6). Megascleres form the primary skeletal support of the sponge.

Microscleres provide secondary reinforcement. Gemmuloscleres form the protec-

tive armor-like coat of gemmules. Because they are produced in a variety of sizes

and forms, spicules, especially gemmuloscleres, are important for sponge identifica-

tion and taxonomy. Spicules are measured in microns (µm) and may be viewed

using a light compound microscope but are best observed with a scanning electron

microscope.

Figure 6. Spicules of Heteromeyenia tubisperma. A,

gemmuloscleres; B megasclere; C, microsclere.

5

Life Cycle:

The life cycle of a freshwater sponges consists of three stages: adult, larvae,

and gemmule; and four phases: active and rapid growth, vegetative, degeneration of

the adult and gemmulation, and quiescence. Reproduction can occur either asexual-

ly or sexually. Asexual reproduction occurs by budding, fragmentation, or gemmule

production. Sexual reproduction results in the formation of flagellated larvae. A

larva freshwater sponges is known as a parenchymula. The active and rapid growth

phase of the life cycle occurs as larvae and archaeocytes are transformed into

adults. The larval stage is the shortest stage of the life cycle; lasting only a few

hours or days. Larvae undergo metamorphosis to become adults. The adult stage

which represents the vegetative phase is responsible for producing the next genera-

tion of sponges. The degeneration of the adult and gemmulation phase occurs as

environmental conditions unfavorable to the adult are encountered, which in Ten-

nessee is during periods of drought or with the onset of falling water temperatures

as winter approaches. Freshwater sponges exist as gemmules during hibernation or

estivation. The time spent in hibernation or estivation represents the quiescence

phase of the life cycle. When environmental condition become favorable for the

survival of the adult sponge gemmules germinate (release Thesocytes) giving rise to

adults (Figure 7).

Figure 7. Annual life-cycle of freshwater sponge.

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Ecological Role:

Sponges are filter feed-

ers. Think of a sponge as

acting as a water filter.

Sponges have water chan-

nels which have an opening

on each end of the channel;

an ostium (pl. ostia) by

which water enters the

sponge, and an osculum (pl.

oscula) through which water

exits the sponge (figure 8).

Food is captures as water

passes through chambers

lined with choanocytes.

Each choanocyte has a col-

lar of microvilli which cap-

tures and filters food and a

flagellum. Flagella through their movements facilitate water movement within the

sponge. Foods include bacteria, phytoplankton, zooplankton, and organic detrital

particulates. Their symbiotic relationship with algae is important in habitats short

on nutrients. Algae provide sponges with photosynthetic produced sugars and oxy-

gen. Algae benefit from sponge produced carbon dioxide and nutrients such as ni-

trogen.

Freshwater sponges serve as a link between benthic and pelagic food webs.

Crayfish of the genus Orconectes, ring-neck duck (Aythya collaris), and few fish

species have been reported to consume freshwater sponges.

Relationship to Other Organisms

Animals and choanoflagellates are considered to have a common eukaryotic

ancestor. The choanoflagellates are a group of free-living unicellular and colonial

flagellate eukaryotes. Choanoflagellates are of particular interest to evolutionary

biologists studying the origins of multicellularity in animals.

The anatomy of a sponge choanocyte is similar to that of choanoflagellates.

Both have a collar of microvilli for capturing food and a flagellum. It has been

thought that choanoflagellates gave rise to sponges. However, molecular studies

have confirmed the two groups are monophyletic, so neither is a descendant of the

other. Sponge choanocytes can differentiate to become sperm.

Historically, sponges have been considered the sister group to all other animals.

Recently some biologists have provided evidence which supports their belief that

jelly combs should occupy this position. However, the traditional view that spong-

es are the sister group is accepted by most biologists.

Figure 8. Surface of freshwater sponge. A, osculum; B,

ostium.

7

Importance of Freshwater Sponges

Freshwater sponges perform several important functions within the ecosystems

they occupy. Some of the functions carried out by freshwater sponges are: 1) as

filter feeders sponges actively and efficiently pump and clean large amounts of wa-

ter, 2) their pumping activity helps circulate the water surrounding them, especially

in lentic waters, 3) their pumping activity traps particulate and dissolved organic

matter thus they play an important role in the re-cycling of organic material 4) pro-

vide a living refuge for a wide array of organisms such as bacteria, algae, protists,

and larvae of caddisflies, spongillaflies and other organisms, 5) can have a symbi-

otic relationship with autotrophic microorganisms, which contributes to primary

production, 6) contribute to the formation of sediments by releasing spicules upon

their death and 7) serve as a link in the transfer of energy between, benthic, pelagic,

and terrestrial food webs. In addition, sponge spicules are used in human skin care

products, and in the production of ceramic products. Sponges and their microbial

symbionts are being investigated as sources for antimicrobial compounds.

Finding Freshwater Sponges:

Finding freshwater sponges is easy and fun. They are found in both lotic

(streams and rivers) and lentic (ponds and lakes) waters. Sponges can be found by

wading, swimming, snorkeling, scuba diving, or dredging. Look for freshwater

sponges on hard substrates (rocks, logs, bridge pilings, shells of mollusks, man-

made objects). The undersides of objects which can be lifted should be viewed.

Rarely are they found on soft substrates such as sand or mud. Size can vary from a

millimeter to over a meter in diameter.

Are Freshwater Sponges Endangered or Threatened?

No freshwater sponges are currently listed as endangered or threatened by the

U. S. Fish and Wildlife Service (USFWS). This may be due to a lack of data. Un-

fortunately, for some species very little is known concerning their abundance and

distribution. Some such as Corvomeyenia carolinensis appears to have a very lim-

ited distribution, currently known from two ponds, the type locality in South Caroli-

na and from one in Connecticut. Others such as Corvospongilla becki are known

from a few widely separated locations. In Tennessee C. becki is known from a sin-

gle location. The lack of information needed for listing by the USFWS could con-

tribute to the extinction of some species.

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Checklist and Distribution Notes of Tennessee Freshwater Sponges

Any checklist should be considered a work in progress that represents the most

reliable information available at the time. Much remains to be learned concerning

Tennessee’s sponges. It is very likely additional species will be discovered. Distri-

bution information is lacking. Most of what we know concerning the distribution

of the freshwater sponges in Tennessee is the result of a single state-wide survey.

.

Potamolepidae:

Cherokeesia armata Copeland, Pronzato, Manconi, 2015: Currently known

only from Tennessee. Apparently locally common in Nolichucky and Hiwassee

Rivers. Also, collected from Buffalo, Red and Sequatchie Rivers

Spongillidae:

Corvospongilla becki Poirrier, 1978: Presently known from a single site on the

Duck River, within Henry Horton State Park, Marshall County.

Ephydatia fluviatilis (Linnaeus, 1759): Collected from the Holston River, Haw-

kins County, and Indian Creek (a tributary of the Powell River) in Claiborne

County.

Ephydatia mulleri (Lieberkuhn, 1855): Collected from Clear Creek of the

Emory River drainage in Morgan County and from the Elk River in Lincoln

County

Eunapius fragilis (Leidy, 1851): A very common and widely distributed species

in Tennessee. Collected from the Buffalo, Calfkiller, Clinch, Duck, East Fork

Obey, Elk, Harpeth, Holston, Nolichucky, Powell, Red, Sequatchie, and South

Fork Holston Rivers, and from Clear Creek and Daddy’s Creek of the Emery

River drainage.

Heteromeyenia latitenta (Potts, 1881): Collected only from the Pigeon River in

Cocke County, where it appears to be common. Until the finding in Tennessee

it had been reported only from the northeastern United States.

Heteromeyenia tubisperma (Potts, 1881): Collected from the Elk River, Giles

County; Richland Creek, Davidson County; Roaring River, Overton County;

and Wolf River, Pickett County.

Racekiela ryderi (Potts, 1882): Common in the Emory River drainage. Collect-

ed from Clear Creek Morgan County, Daddy’s Creek, Cumberland and Morgan

Counties; Obed River, Morgan County. Also collected from Hiwassee River,

Polk County.

9

Radiospongilla cerebellata (Bowerbank, 1863): This sponge in the United

States is an enigma. R. cerebellata is an oriental species but has reported from

Texas, Alabama, and now Tennessee. Manconi and Pronzato (2016) doubt that

this North American sponge is R. cerebellata. This sponge appears phenotypi-

cally to be R. cerebellata but could represent a new species. DNA analysis is

needed to clarify this situation. This sponge has been collected from eastern

Tennessee waters. Collected from French Broad River, Cocke County; Nol-

ichucky River, Green and Washington Counties; and Pigeon River, Cocke

County.

Radiospongilla crateriformis (Potts, 1882): This sponge is broadly distributed

across Tennessee. Collected from Abrams Creek, Blount County; Elk River,

Lincoln County; Mill Creek, Davidson County; Nolichucky River, Greene

County; and Sequatchie River, Marion County.

Spongilla lacustris (Linnaeus, 1759): A common and broadly distributed

sponge in Tennessee. Collected from Buffalo River, Lewis County; Cane

Creek, Van Buren County; Clear Creek, Morgan County; Cumberland River,

Davidson County; Duck River, Maury County; Little Sequatchie River, Marion

County; Mill Creek, Davidson County; New River, Scott County; Nolichucky

River Greene and Washington Counties; Piney Creek, Van Burren County;

Obed River, Cumberland County; Red River, Robertson County; Richland

Creek, Davidson County; and South Fork Holston River, Sullivan County.

Trochospongilla horrida (Weltner, 1893): A common and widely distributed

sponge in Tennessee. Collected from Abrams Creek, Blount County; Buffalo

River, Lewis County; Calfkiller River, White County; Clear Creek, Morgan

County; Clinch River, Hancock County; Collins River, Warren County; Cona-

suga River, Polk County; Cumberland River, Davidson River; Duck River Mar-

shall and Maury Counties; Elk River, Giles County; Harpeth River, Cheatham

County; Little Sequatchie River, Marion County; New River, Scott County;

Nolichucky River, Washington County; and Obed River, Cumberland County.

Trochospongilla leidii (Bowerbank, 1863): Current known distribution is lim-

ited to the Nashville Basin. Collected from Cumberland and Harpeth Rivers,

Davidson and Cheatham Counties.

10

References:

Copeland, J., S. Kunigelis, J. Tussing, T. Jett, and C. Rich. 2019. Freshwater spong-

es (Porifera: Spongillidae) of Tennessee. The American Midland Naturalist,

181:310-326.

Copeland, J. E., J. A. Tussing, T. M. Jett, and S. Kunigelis. 2015a. Freshwater Po-

rifera of eastern Tennessee. J. TN Acad. of Sci., 90:23.

Copeland, J., R. Pronzato, and R. Manconi. 2015b. Discovery of living Potamolepi-

dae (Porifera: Spongillina) from Neartic freshwater with description of a new

genus. Zootaxa, 3957:37- 48.

Hoff, C. C. 1943. Some records of sponges, branchiobdellids, and molluscs from

the Reelfoot Lake region. J. TN. Acad. Sci., 18:223-227.

Kunigelis, S. C. and J. E. Copeland. 2014. Identification of isolated and in situ

freshwater sponge spicules of eastern Tennessee. Microsc. Microanal., 20

(Suppl. 3):1294-1295.

Manconi, R. and R. Pronzato. 2002. Suborder: Spongillina subord. Nov.: Freshwa-

ter Sponges, InHooper, J.N.A. & Van Soest, R.W.M. (Eds), Systema Porifera:

A Guide to the classification of Sponges. Kluwer Academic/Plenum Publishers,

New York, pp.921-1019.

Manconi, R. and R. Pronzato. 2015. Phylum Porifera, p. 133-157. In: Thorp, J. and

D. C. Rogers (eds.) Ecology and General Biology: Thorp and Covich’s Fresh-

water Invertebrates. 4th Edition, Vol. 1. Academic Press, London, UK.

Manconi, R. and R. Pronzato. 2016a. How to survive and persist in temporary

freshwater? Adaptive traits of sponges (Porifera: Spongillida): A review. Hy-

drobiologia, 782:11-22.

Manconi, R. and R. Pronzato. 2016b. Phylum Porifera, p. 39-83. In: Thorp, J. and

D. C. Rogers (eds.): Keys to Nearctic Fauna: Thorp and Covich’s Freshwater

Invertebrates. 4th Edition, Vol. 2. Academic Press, London, UK.

Matsuoka, T. and Y. Masuda. 2000. A new potamolepid freshwater sponge

(Demospongiae) from the Miocene Nakamura Formation central Japan, Paleon-

tol. Res., 4:131-137.

Mobley, A. S. 2010. The bacterial community of a freshwater sponge, Radiospon-

gilla cerebellata: a comparison of terminal restriction fragment length poly-

morphisms (T-RFLP) and 16S RRNA clone library methods. M. S. Thesis, The

University of Alabama at Birmingham. p. 63.

Parchment, J. G. 1966. Notes on the ecology of sponges. J. TN Acad. Sci., 41:65.

Penney, J. T. 1960. Distribution and bibliography (1892-1957) of the fresh water

sponges. University of South Carolina Publications, Series 3, 3:1-97.

Penney, J. T. and A. A. Racek. 1968. Comprehensive Revision of a Worldwide Col-

lection of Freshwater Sponges (Porifera: Spongillidae). U. S. Nat. Mus. Bull.,

272.

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Pisera, A., P. A. Siver, and A. P. Wolf. 2013. A first account of freshwater potamo-

lepid sponge (Demospongiae, Spongillina, Potamolepiidae) from the middle

Eocene: biogeographic and paleoclimatic implications. J. Paleontol., 87:373-

378.

Poirrier, M. A. 1972. Additional records of Texas freshwater sponges

(Spongillidae) with the First record of Radiospongilla cerebellata (Bowerbank,

1863) from the Western Hemisphere. The Southwest. Nat., 16:434-435.

Poirrier, M. A. 1978. Corvospongilla becki n. sp., a new fresh-water sponge

from Louisiana. Trans. Amer. Micros. Soc., 97:240-243.

Pronzato, R. and R. Manconi. 1994. Adaptive strategies of sponges in inland wa-

ters. Bollettino di Zoologica, 61, 395-401.

Reiswig, H. M., T. M. Frost, and A. Ricciardi. 2010. Chapter 4 Porifera, p. 91-123.

In: Thorp, J. H. and A. P. Covich, (eds.). Ecology and Classification of North

American Freshwater Invertebrates. Academic Press, London, UK.

Ricciardi, A. and H. M. Reiswig. 1993. Freshwater sponges (Porifera, Spongillidae)

of Eastern Canada: taxonomy, distribution, and ecology. Can J. Zool., 71:665-

682.

Van Soest, R. W. M., N. Boury-Esnault, J. N. A. Hooper, K. Rützler, N. J. de

Voogd, B. Alvarez, E. Hajdu, A. B. Pisera, R. Manconi, C. Schӧnberg, M.

Klautau, B. Picton, M. Kelly, J. Vacelet, M. Dohrmann, M.-C. Diaz, P. Cárde-

nas, J. L. Carballo, P. Rios, and R. Downey. 2018.World Porifera Database.

Accessed at http://www.marinespecies.org/porifera/ on 2018-6-1.

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About the Cumberland Mountain Research Center

Named for the Cumberland Mountains in which Lincoln Memorial University

(LMU) was founded, the Cumberland Mountain Research Center (CMRC) was es-

tablished in 1990 on the LMU campus to foster a collaborative research environ-

ment among faculty and students as part of the larger science education mission of

the University, and to provide a broad range of research facilities, opportunities,

and resources to LMU faculty and student researchers. The original objective was

to take advantage of the unique ecology of southern Appalachia that surrounds the

Cumberland Gap region. Working with a variety of institutions and organizations,

including state and federal agencies, these efforts quickly expanded to include many

areas across the Southeastern United States, as well as westward into Arkansas.

Shortly after this time, the CMRC also began developing international opportunities

and it continues to provide and facilitate regional, national, and international re-

search and educational opportunities for LMU faculty and students to this day.

With a particular interest in conservation biology and ecology, the CMRC is

committed to enhancing research support to LMU faculty and students by facilitat-

ing the scholarship of discovery through high-quality research, creative activities,

and outreach initiatives. Central to this mission, the CMRC seeks research opportu-

nities, partnerships, and collaborations to foster strong research and outreach ef-

forts, as well as seek funding and resources necessary to facilitate these endeavors.

As a point of contact for all it’s stakeholders, partners, and collaborators, the

CMRC facilitates, as needed, project and research development, management, com-

munication, and dissemination of results while adhering to the highest ethical stand-

ards and professional best practices.

For additional information about the CMRC and its activities, please contact

the CMRC Director, LaRoy Brandt, via email at laroy.brandt@LMUnet.edu, the

LMU School of Math and Sciences, or the LMU Department of Biology. You can

also follow the Cumberland Mountain Research Center on Facebook.

13

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