Copepod Crustaceans of Great Smoky Mountains National Park:
First Collections, Preliminary Impressions, and Results

Janet W. Reid
Research Associate, Department of Invertebrate Zoology
National Museum of Natural History, Smithsonian Institution
Washington DC 20560-0163
phone (202) 357-4674 fax (202) 357-3043

1. Collections and conditions:

My husband Willis and I made 2, 10-day collecting excursions to the Park, in October 1998 and late May-early June 1999. During each visit we made day trips to about 4 general areas and sorted the samples live between trips. In October we stayed in the house at the Natural Resources Center at Twin Creeks uphill from Gatlinburg, and in May/June at the ranger house at Twentymile Creek, then at the house at Cosby dedicated to the ATBI collectors, in the southwest and northeast corners of the Park respectively.

Our needs for sorting consist basically of a sturdy table, electricity, and a sink, and we could set up quite comfortably in the barn at Twin Creeks, on the kitchen table at the small Twentymile house, and more spaciously at the second house at Cosby which has been slightly adapted as a sorting facility. We took and pre-sorted some 90 samples (including multiple samples of different substrates- mud, moss, creek gravel, "plankton", crayfish holes, etc.- at each general site) during each visit.

Keith Langdon, Chuck Parker, Becky Nichols, Jody Flemming, and all the park personnel were extremely helpful. Keith and Chuck have an almost encyclopedic knowledge of likely collecting spots, geology, water types, use history, etc., and with their help and excellent maps we got to a nice array of habitats. Not their fault that in October/98 some waterbodies were dry or nearly so after extreme drought conditions in summer/98; things were a little wetter this May.

We reached most collecting areas via car, and did only a little hiking. The roads were all in good shape, but of course winding, and travel time was significant. Those periods are of course popular times for visitors, so we avoided collecting on weekends. Any hikers we met readily accepted our explanation that we were "doing water sampling" or "surveying small aquatic animals for the park". There were no incidents with other flora or fauna, including the bear in Cades Cove who was more interested in a passing deer than in us.

To sum up, on both visits we were comfortably housed and efficiently provided with all possible practical support, for which we thank all the people at the Park, plus the NPS and Discover Life which provided the housing. We can also recommend some good restaurants ...

During 1999, Will Reeves of Clemson University made several visits to the Park, collecting invertebrates in small springs and caves. Will sorted and sent me 4 samples of copepods, including one (unfortunately an unidentifiable juvenile) from Rich Mountain Blowhole Cave. Will continues to collect, and we anticipate interesting finds.

Prof. David K. Smith of the University of Tennessee, Knoxville has kindly made identifications of moss samples from which copepods were collected. Eventually, knowledge of moss species preferred by particular species of copepods should yield useful information on the range of environmental conditions preferred.

2. The Smokies free-living copepods - impressions:

During the October, 1998 trip, we collected 40 species, and, with determination of the May/June/99 samples only about 20% complete, have added 22 more. Of these, 17 taxa are new or probably new (i.e., members of groups in need of revision).

The first published paper including information on Smokies copepods came out this year (Strayer & Reid, 1999). One sample from Little River was included in the data for that paper. I have now completed a review of the historical literature, including contacting several colleagues, and biologists at the Tennessee Valley Authority in regard to Fontana and other bordering reservoirs. So far, I have found no historical literature, and I have located no one who has collected copepods from the park. Therefore we begin basically at ground zero for this group.

At the 1997 ATBI planning workshop I made a wild estimate of 100 copepod species to be expected in the park. This would be considered a rather speciose fauna; to compare some other, relatively well-collected temperate regions, there are 54 native free-living species recorded from the District of Columbia, 66 species from North Carolina, 70 from the Laurentian Great Lakes, and 80 from Belgium.

We found only 3 calanoid species, all in the TVA reservoirs bordering the Park. This low number was expected, since calanoids are mainly planktonic in lakes and ponds, of which there are only a few in the Park. A few additional species may show up in the larger ephemeral pools such as Gum Swamp; in fact, we did collect a single juvenile calanoid in Gum Swamp, but were unable to identify it before it was eaten by a cyclopoid copepod in the same sample.

However, two of the calanoid species are interesting. Fontana Lake is in the extreme northern part of the range of Epischura fluviatilis, a southeastern US member of the family Temoridae. Skistodiaptomus carolinensis (family Diaptomidae) was described in 1986 from Lake Ravenel, Highlands, NC. Because of its extremely restricted distribution, S. carolinensis was listed as a species of "special concern" by Clamp et al. (1999) and as "threatened/vulnerable" in the 1996 IUCN Red Book, and I am pleased to find it in another location.

As expected, the remaining species are about evenly divided between the orders Cyclopoida and Harpacticoida. Most of the known species are common and widely distributed. However, Stolonicyclops heggiensis, first described in 1998 from seeps on Heggie's Rock, a granite monadnock in Georgia, has shown up; as has Rheocyclops indiana, known previously only from one location, a streambed, in Indiana of course. These are somewhat surprising range extensions for species that are both known only from substrate-bound, groundwater-related situations.

The most spectacular range extension, however, was the find of "Diacyclops" yezoensis, described from Japan and, in North America, previously found only in southeastern Alaska. A single female occurred in the spring behind the Natural Resources Center. With a Japanese colleague I am proposing a new genus for this taxon (Reid and Ishida, in review).

Certain primarily northern species such as the harpacticoid Attheyella obatogamensis are common in the Park, though rather rare in lowland piedmont areas. The Smokies collection data from October/98 allowed me to re-evaluate the status of A. obatogamensis and a few other species in North Carolina for the report by Clamp et al. (1999). However, we know so little about the distribution and habitat preferences of non-planktonic copepods that it is difficult to make generalizations. All the copepod species found so far in the Smokies appear to be North American natives.

At the 1997 workshop, I predicted that there would be a high degree of endemism, mainly of the more groundwater-associated species. The 17 new taxa of 62 is certainly a "high" proportion; in comparison, 5 of the 54 native species from DC are known only from DC or nearby. The number of "endemic" species will doubtless rise as we begin to get samples from less accessible groundwater habitats such as the streambeds and the caves. Keith Langdon informed me that old USGS wells are being located and re-opened, which will help access some of the deep fauna.

This type of sampling will of course be much more labor-intensive and will require special drilling and pumping equipment. "Streambeds" in the Smokies are thick with rocks and boulders, making access to the hyporheic fauna via the usual hand-driven Bou-Rouch pump extremely difficult or impossible. David Strayer told me that the Little River gravel bar site that he sampled was his "least favorite" in the entire Appalachians, because of the rock factor. Moreover, sorting tiny interstitial copepods from the mass of organic floc in the stream sediments is a long, long job. However, most of the more "interesting" (new, rare, or restricted to a single drainage basin) species would be expected to appear in the hyporheic and other groundwaters.

There are perhaps 20 additional widely distributed but not often collected species known from the eastern US, which might be expected to show up in the park. Taking all this into account, I still predict that the copepod list for the park will eventually include 80-100 species.

I prepared an abstract for a poster presentation at the VII International Conference on Copepoda in Curitiba, Brazil, 25-30 July 1999. This may be referenced on the Conference website, although the numbers in it are outdated (Reid, 1999).

I participated in the workshop "Copepod Diversity in Neotropis: Present Knowledge and New Directions for Research", São Sebastião, São Paulo, Brazil, 21-23 July 1999. Data from the Smokies contributed to the species list used to compare the structure and composition of the copepod communities of tropical Rio de Janeiro and temperate North Carolina. An article for the volume of workshop proceedings is in review.

I am preparing a manuscript describing the 3 new species of Elaphoidella, and a related species from DC.


Clamp, J.C. (compiler), W.F. Adams, J.W. Reid, A.Y. Taylor, J.E. Cooper, C. McGrath, D.J. Williams, D.J. DeMont, W.O. McLarney, G.
Mottesi and J. Alderman. 1999. A report on the conservation status of North Carolina's freshwater and terrestrial crustacean fauna. Scientific Council on Freshwater and Terrestrial Crustaceans, Report to North Carolina Wildlife Resources Commission, Raleigh. 92pp.

Reid, J.W. 1999. The all-taxa biodiversity inventory (ATBI) of the Great Smoky Mountains National Park, U.S.A.; initial collections of free-living copepods. Poster, VII International Conference on Copepoda, Curitiba, Brazil, 25-30 July 1999. Abstract on conference website:

Reid, J.W. & T. Ishida. Itocyclops, a new genus proposed for Diacyclops yezoensis (Copepoda: Cyclopoida). Journal of Crustacean Biology, in review.

J. W. Reid and C. E. F. Rocha. Are tropical copepods more diverse? Comparison of the faunas of the states of Rio de Janeiro, Brazil, and North Carolina, U.S.A. Proceedings of the workshop "Copepod Diversity in Neotropis: Present Knowledge and New Directions for Research," São Sebastião, Brazil, 21-23 July 1999. In review.
Strayer, D.L. & J.W. Reid. 1999. Distribution of hyporheic cyclopoids (Crustacea: Copepoda) in the eastern United States. Archiv für Hydrobiologie 145(1): 79-92.

3. Accomplishments and milestones:

4. Acknowledgments

Willis and I are extremely grateful to the NPS and to Discover Life in America for their support in terms of providing housing and research facilities in the Park, and for making it possible for me to participate in the planning workshop in December 1998. The grant from DLIA made to me this year will go directly toward publication costs for two articles in peer-reviewed journals, one of which is now in review and the other in preparation.

Our thanks to Keith Langdon, Chuck Parker, Becky Nichols, Jody Flemming, and many other people at the Park; Will Reeves for collections; and D.K. Smith for moss identifications.

List of Copepod Crustacean Species:

Order Calanoida
Family Centropagidae
Osphranticum labronectum S. A. Forbes, 1882

Family Diaptomidae
Skistodiaptomus carolinensis Yeatman, 1986

Family Temoridae
Epischura fluviatilis Herrick, 1883

Order Cyclopoida
Family Cyclopidae
Subfamily Eucyclopinae
Ectocyclops phaleratus (Koch, 1838)
Eucyclops agilis (Koch, 1838)
Eucyclops conrowae Reid, 1992
Eucyclops elegans (Herrick, 1884)
Eucyclops cf. prionophorus Kiefer, 1931
Eucyclops A
Eucyclops B
Eucyclops C
Macrocyclops albidus (Jurine, 1820)
Megacyclops A
Paracyclops chiltoni (Thomson, 1882)
Paracyclops fimbriatus-group A (Fischer, 1853)
Paracyclops poppei (Rehberg, 1880)
Tropocyclops extensus Kiefer, 1931
Tropocyclops prasinus mexicanus Kiefer, 1938
Tropocyclops A

Subfamily Cyclopinae
Acanthocyclops brevispinosus (Herrick, 1884)
Acanthocyclops exilis Coker, 1934
Acanthocyclops robustus (Fischer, 1820; "warm-water strain" of Dodson, 1994)
Acanthocyclops robustus-group A
Acanthocyclops venustoides Coker, 1934
Acanthocyclops vernalis (Fischer, 1853)
Acanthocyclops A
Acanthocyclops B
Acanthocyclops C
Diacyclops crassicaudis var. brachycercus (Kiefer, 1929)
Diacyclops haueri Kiefer, 1931
Diacyclops nearcticus Kiefer, 1934
Diacyclops thomasi (S. A. Forbes, 1882)
Diacyclops A
Itocyclops yezoensis (Ito, 1954)
Mesocyclops edax (S. A. Forbes, 1891)
Microcyclops rubellus (Lilljeborg, 1901)
Orthocyclops modestus (Herrick, 1883)
Rheocyclops indiana Reid, 1999
Stolonicyclops heggiensis Reid & Spooner, 1998
Order Harpacticoida

Family Canthocamptidae
Attheyella americana (Herrick, 1884)
Attheyella carolinensis Chappuis, 1932
Attheyella illinoisensis (S.A. Forbes, 1882)
Attheyella obatogamensis (Willey, 1925)
Bryocamptus hiemalis (Pearse 1905)
Bryocamptus hutchinsoni Kiefer, 1929
Bryocamptus newyorkensis (Chappuis, 1927)
Bryocamptus nivalis (Willey, 1925)
Bryocamptus zschokkei (Schmeil, 1893)
Bryocamptus A
Bryocamptus B
Bryocamptus C
Bryocamptus D
Canthocamptus assimilis Kiefer, 1931
Elaphoidella bidens (Schmeil, 1894)
Elaphoidella A
Elaphoidella B
Elaphoidella C
Epactophanes richardi Mrazek, 1893
Moraria cristata Chappuis, 1929
Moraria virginiana Carter, 1944

Family Parastenocarididae
Parastenocaris A

Family Phyllognathopodidae
Phyllognathopus viguieri (Maupas, 1892)

Total 62 species (17 new)