Illinois Natural History Survey - University of Illinois

THE AQUATIC ANNELIDA (CLITELLATA) 
OF THE COLORADO RIVER AND ITS TRIBUTARIES, 
GRAND CANYON NATIONAL PARK, ARIZONA


This web site established and maintained by

Mark J. Wetzel, Research Scientist
- Curator, Collections Manager, and Database Manager, INHS Annelida Collection
Illinois Natural History Survey (INHS), University of Illinois Institute of Natural Resource Sustainability,
Champaign, Illinois 61820 USA

Introduction.  In 1990, the Glen Canyon Environmental Studies Program in cooperation with the Department of Biology, Northern Arizona University, Flagstaff (NAU), began a long-term study of the benthic dynamics of the Colorado River and its tributaries located in the Grand Canyon National Park (GCNP), Coconino and Mohave counties, Arizona. For several years, this project was directed by Drs. Dean W. Blinn and Joseph P. Shannon (NAU), with assistance (through 1993) from Dr. Larry E. Stevens (now with Grand Canyon Monitoring and Research Center); more recently (after Dr. Blinn retired from the faculty of NAU), the project has been directed by Dr. Shannon.


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Photos (© M.J. Wetzel, INHS) taken during our 20-30 June 2001 trip on the Colorado River in Grand Canyon National Park, Arizona [Colorado River (COR) river mile (RM) noted].


 

Numerous taxonomists and systematists with expertise in one or more groups of plants and animals are participating in this project, and are being assisted by ecologists, data specialists, aquatic and terrestrial biologists, teachers, students, and regional volunteers.My primary focus for this project is to identify freshwater oligochaetes, leeches, and any other aquatic or semi-aquatic annelids that I have collected from the Colorado River, its direct tributaries, and other unique habitats within the GCNP. I especially welcome any specimens that you may have collected from aquatic habitats within the GCNP, and your published records of aquatic and terrestrial Annelida from this region. Your findings (with credit) will be incorporated into this website, and your publications will be included in the literature cited section of this document if you forward a reprint to me. Your suggestions for and comments on this project are welcome.

Permits. All research associated with this project in the GCNP complies with and has been / is being conducted under the following Scientific Research and Collecting Permits issued by the USDI-NPS: GRCA1993AHRR (issued to Dr. Dean Blinn, Northern Arizona University, Flagstaff); and GRCA1999-20-11, GRCA-1997-SCI-071, and GRCA-2000-0071 (all issued to Dr. Joseph Shannon, Northern Arizona University, Flagstaff). Information pertinent to the NPS-issued collecting permit covering surveys by MJW on the 1991 expedition is not available at this time, although I was assured that my research and that of the expedition were covered at that time by a current collecting permit. All of the trips were affiliated directly with a continuing project -- monitoring of the effects of interim flows on the aquatic food base [Cladophora, and amphipods, chironomids, oligochaetes, and several other dominant invertebrate taxa] in the Colorado River downstream of the Glen Canyon Dam, Arizona.Please note that this is a working document. If you have additions or corrections to the information presented below, please forward them to me via E-mail, at: * mjwetzel[AT]illinois.edu

Acknowledgements. Funding for this research, in part, has been provided by the U.S. Department of the Interior through the Bureau of Reclamation, Glen Canyon Environmental Studies Program, and the Grand Canyon Monitoring and Research Center--U.S. Geological Survey, in cooperation with the Grand Canyon National Park, Glen Canyon National Recreation Area, and Northern Arizona University.  I also thank the People of the Havasupai Nation - the Havsuw Baaja, or "People of the Blue-Green Waters," and also the People of the Hualapai Nation - "People of the Tall Pines" and "Keepers of the Canyon." Finally, I acknowledge the continuing support of my institution, the Illinois Natural History Survey. I am particularly grateful to Drs. Dean Blinn and Joe Shannon, directors of this project, for their support and encouragement, and for the opportunity to participate in this outstanding project.  Effective collection of annelid specimens during the 1991, 1993, 2001, and 2006 river expeditions would not have been possible without the professional field assistance of numerous colleagues and friends –

1991 and 1993 river trips:
expeditions would not have been possible without the professional field assistance of Joseph P. Shannon, Michael L. Shaver, Jeanette Macauley, Clay Runck, Alla Sineeva, Candace Stewart, Tom Martin, Molly & Peter Absolon, Marcy Demillion, Teresa Yates, Dennis Silva, Mary Moran, OC Davis, Bob Grusy, Jano Kempster, Monte Becker, Lew Steiger, Stephanie Trimmer, Jim Thomas, Larry E. Stevens, Michelle L. Biyal, Susan J. Hueftle, Fredric R. 'Frederheic' Govedich, Emma L. Benenati, Bonnie Ellis, Connie L. Graham, Jack Stanford, G. Richard Marzoff, Jeff Pflueger, and several other volunteers.  Lab assistant M.L. Biyal was responsible for the sorting and processing of macroinvertebrate samples collected in 1991 and 1993, the slide mounting of oligochaete specimens, and database entry of pertinent project information. 

2001 river trip (20-30 June):
Emma L. Benenati, Nelleka DeVane, Allen Haden, Pat Haden, Carolyn Link, Chris Niebuhr, Joe Shannon, Kristin Straka, Susie Vogel, and Kevin Wilson, and to GCMRC personnel Carol 'Fritz' Fritzinger, Park Steffensen, and Jake Tiegs. Peggy Morgan (Florida Department of Environmental Protection), who was assisting me with the collection of oligochaetes during this trip, was instrumental in our ability to concentrate on the diversity and uniqueness of habitats (especially the spring, seep, and COR tributaries), particularly when our time was limited at sites because of time-dependent aspects of the research projects supporting this trip, and also because of schedules associated with river flow, hydroelectric discharge from the Glen Canyon dam, and other characteristics of the COR and its tributaries.  

2006 river trip (10-18 July):
Nick Ballew, Emma L. Benenati, Chris Blattel, Steve Marlatt, Peggy Morgan, Bob Murphy, John Rihs, Jeff Scoggins, and Joe Shannon.I thank Tim Grooms and other personnel of Yellow Springs Instrument Company, Inc. for the loan of the model YSI 556 multiparameter handheld meter (dissolved oxygen, DO % saturation, water temperature, conductivity, hydrogen ion concentration [as pH], ORP, resistivity, and barometric pressure); this meter performed flawlessly, affording a convenient and time-saving approach to accurate measurement of several physical / chemical water quality parameters amidst the relatively extreme conditions present in the Grand Canyon at this time of year. [I have used numerous meters in the YSI product line for over 30 years, and they have been extremely reliable; although I cannot provide specific endorsements of commercially available products, the Model 556 meter did serve flawlessly in an 'emergency field situation' after a comparable meter (manufactured by a competing company) failed to 'meet expectations'.]   


Collecting Sites.

All collecting sites sampled during the river trips of 1991, 1993, 2001, and 2006 are located in either Coconino or Mohave counties in Arizona. Colorado River river mile (COR RM, or RM) coordinates are taken from Stephens (1983). All other locality information, including Universal Transverse Mercator System (UTM) coordinates, has been taken from U.S. Geological Survey 7.5 minute series topographic quadrangle maps (USGS tqm); the North American Datum (NAD) of 1927 applies to all maps.  The edition (year of publication, photorevision, and photoinspection, when applicable) is noted in each site description. Township, Range, and Section information, when available, is governed by the Gila & Salt River Meridian (G&SRM).  UTM coordinates recorded for the 2001 collections were obtained using a Garman GPS III Plus. 


1991 Collections. 

In November and December 1991, 21 aquatic sampling sites were surveyed; these included six sites on the main stem of the Colorado River (including backwater bays); Warm Springs / Lava Well and two unnamed backwater marsh areas that drain into the Colorado River; and 12 direct tributary streams of the Colorado River (Vasey's Paradise, Saddle Canyon Creek, Little Colorado River, Lava [Chuar] Canyon Creek, Bright Angel Creek, Deer Creek, Matkatamiba Canyon Creek, an unnamed spring/seep/creek at the "Ledges", National Canyon Creek, Spring Canyon Creek, Granite Park Canyon spring/seep, and Diamond Creek. Aquatic oligochaetes were collected from 17 of these 21 sites.

1991 sites located in Coconino County:

Colorado River (RM 0.0), at Lees Ferry, north side of river, in small inlet, ~50m upstream boat landing area, in Grand Canyon National Park. G&SRM: Township 40 North, Range 8 East, SW/4, NW/4, NW/4, SW/4, Section 18. U.T.M. Zone 12: 447910m East, 4079950, North. Elevation: 3,107ft. Lees Ferry, Arizona (1985 prov. ed.) USGS tqm. 20 November 1991. Collectors: M.J. Wetzel (#1991-70) (INHS), J. Macauley (NAU).

Colorado River (RM 0.0), at Lees Ferry, several areas near the project's transects T2 and T4 in the immediate vicinity of the gauging station cable, just downstream of the boat launching / parking lot areas, in Grand Canyon National Park. G&SRM: Township 40 North, Range 7 East, SE/4, NE/4, NE/4, SE/4, Section 13. U.T.M. Zone 12: 447640m East, 4079900m North. Elevation: 3,107ft. Lees Ferry, Arizona (1985 prov. ed.) USGS tqm. 20 November 1991. Collectors: M.J. Wetzel (#1991-71) (INHS), J. Macauley (NAU).

Colorado River (RM 0.0), on north side of river, just downstream Lees Ferry boat landing area, 10m downstream gauging station cable, in Grand Canyon National Park. G&SRM: Township 40 North, Range 8 East, SW/4, NE/4, NE/4, SE/4, Section 13. U.T.M. Zone 12: 447605m East, 4079895m North. Elevation: 3,107ft. Lees Ferry, Arizona (1985 prov. ed.) USGS tqm. 20 November 1991. Collectors: M.J. Wetzel (#1991-72) (INHS), J. Macauley (NAU).

Colorado River (RM 31.5R), in small inlet (r.b.f.d.), near "confluence" of South Canyon Creek [when flowng], 100m upstream South Canyon Rapids, in Grand Canyon National Park. U.T.M. Zone 12: 423320m East, 4039980m North. Elevation: 2,875'ft. North Canyon Point, Arizona (1985 prov. ed.) USGS tqm. 21 November 1991. Collectors: M.J. Wetzel (#1991-73) (INHS), Alla Sineeva (Russia).

Vasey's Paradise (hanging spring/waterfall/cascade), on rock face near lower part waterfall/cascade, and in pool at base of cascade, just up (in elevation) from its confluence with the Colorado River (RM 31.8), in Grand Canyon National Park. U.T.M. Zone 12: 423270m East, 4039320m North. Elevation: 2,880ft. Tatahatso Point, Arizona (1988 prov. ed.) USGS tqm. 21 November 1991. Collectors: M.J. Wetzel (#1991-74) (INHS), J. Macauley, T. Martin (NAU).

Small water pot, in boulder, at river's edge, Colorado River (RM 43.2R), 0.6 mi upstream President Harding Rapids, just east of Anasazi Foot Bridge, N of Point Hansbrough, in Grand Canyon National Park. U.T.M. Zone 12: 423340m East, 4028070m North. Elevation: 2,835ft. Tatahatso Point, Arizona (1988 prov. ed.) USGS tqm. 21 November 1991. Collectors: M.J. Wetzel (#1991-75) (INHS), Michael Shaver (NAU).

Saddle Canyon Creek, in pools and riffles below small (10') spillway/falls, near spring, approx. 1.8km upstream (SW) confluence with Colorado River (RM 47R), in Marble Canyon of Grand Canyon National Park. U.T.M. Zone 12: 418850m East, 4024040m North. Elevation: 3,590ft. Point Imperial, Arizona (1988 prov. ed.) USGS tqm. 22 November 1991. Collector: M.J. Wetzel (#1991-76) (INHS).

Colorado River (RM 52.7), in Nankoweap Bay (rbfd), just downstream Nankoweap rapids (second set), in Grand Canyon National Park. U.T.M. Zone 12: 423160m East, 4017380m North. Elevation: 2,785ft. Nankoweap Mesa, Arizona (1988 prov. ed.) USGS tqm. 22 November 1991. Collectors: M.J. Wetzel (#1991-77) (INHS), Tom Martin et al. (NAU).

Backwater marsh, draining into north side of Nankoweap Bay of Colorado River (RM 52.7), just west of the lower set of Nankoweap Rapids; in Grand Canyon National Park. U.T.M. Zone 12: 423080m East, 4017430m North. Elevation: 2,887ft. Nankoweap Mesa, Arizona (1988 prov. ed.) USGS tqm. 22 November 1991. Collector: M.J. Wetzel (#1991-78) (INHS).

Little Colorado River (upstream 'arm', at island split before confluence), on north side (r.b.f.d.) of river (LCR), 200m upstream confluence with main channel Colorado River (RM 61.4), in Grand Canyon National Park. U.T.M. Zone 12: 428400m East, 4005380m North. Elevation: 2,710ft. Cape Solitude, Arizona (1988 prov. ed.) USGS tqm. 23 November 1991. Colectors: M.J. Wetzel (#1991-79) (INHS); with Jeanette Macauley, Michael Shaver (NAU), and Candace Stewart, Jim Thomas (volunteers).

Colorado River (RM 61.4), on upstream side (north edge) of cobble island located in the middle of the main channel of the river at confluence point with the Little Colorado River, in Grand Canyon National Park. U.T.M. Zone 12: 428040m East, 4005500m North. Elevation: 2,710ft. Cape Solitude, Arizona (1988 prov. ed.) USGS tqm. 23 November 1991. Collector: M.J. Wetzel (#1991-80) (INHS).

Lava [Chuar] Creek, 100m upstream confluence with Colorado River (RM 65.5R), at Lava Canyon Rapids, in Grand Canyon National Park. U.T.M. Zone 12: 426460m East, 3999480m North. Elevation: 2,690ft. Cape Solitude, Arizona (1988 prov. ed.) USGS tqm. 23 November 1991. Collectors: M.J. Wetzel (#1991-81) (INHS), Joe Shannon, Dennis Silva, Mary Moran (NAU).

Bright Angel Creek, 0.25 mi (0.4km) S Phantom Ranch, 5m downstream campground footbridge; upstream confluence with Colorado River (RM 87.8R), in Grand Canyon National Park. U.T.M. Zone 12: 401425m East, 3995690m North. Elevation: 2,435ft. Phantom Ranch, Arizona (1988 prov. ed.) USGS tqm. 25 November 1991. Collector: M.J. Wetzel (#1991-82) (INHS).

Deer Creek, from pools and riffles 20m downstream of Deer Creek Falls, just upstream of its confluence with Colorado River (RM 136.1R), in Granite Narrows, Grand Canyon National Park. U.T.M. Zone 12: 364770m East, 4027950m North. Elevation: 1,940ft. Fishtail Mesa, Arizona (1988 prov. ed.) USGS tqm. 27 November 1991. Collector: M.J. Wetzel (#1991-83) (INHS).

Matkatamiba Canyon Creek, 100m upstream confluence w/ Colorado River (RM 147.9L), in Grand Canyon National Park. U.T.M. Zone 12: 350020m East, 4023110m North. Elevation: 1,840ft. Havasu Falls, Arizona (1988 prov. ed.) USGS tqm. 28 November 1991. Collector: M.J. Wetzel (#1991-84) (INHS).

National Canyon Creek, 1.6 km upstream (ESE) "confluence" with Colorado River (RM 166.4L) [stream was intermittent/hyporheic], on Hualapai Indian Reservation, adjacent to Grand Canyon National Park. U.T.M. Zone 12: 331500m East, 4013560m North. Elevation: 1,900ft. Fern Glen Canyon, (1988 prov. ed.) USGS tqm. 29 November 1991. Collector: M. J. Wetzel (#1991-86).

Lava Well / Warm Springs, in pool along bluff of Colorado River (RM 179.4L), 150m SE Lava Falls Rapid, on Hualapai Indian Reservation, adjacent to Grand Canyon National Park. U.T.M. Zone 12: 312880m East, 4007520 m North. Elevation: 1,665ft. Vulcans Throne, Arizona (1967 ed.) USGS tqm. 29 November 1991. Collectors: M. J. Wetzel (#1991-87) (INHS), Michael Shaver (NAU).

Granite Park Canyon spring/seep (Granite Park Canyon Creek), 0.9 km upstream "confluence" with Colorado River (RM 208.8L), in Grand Canyon National Park. U.T.M. Zone 12: 291720 m East, 3982230m North. Elevation: 1,575ft. Granite Park, Arizona (1967 ed.) USGS tqm. 1 December 1991. M.J. Wetzel (#1991-89) (INHS), Michael Shaver (NAU).

1991 sites located in Mohave County:

Spring / seep / creek, at "the Ledges", Colorado River (RM 151.6R); collections taken from pools and on slab rocks located above (in elevation) the river, in Grand Canyon National Park. U.T.M. Zone 12: 345180m East, 4023460m North. Elevation: 1,845ft. Havasu Falls, Arizona (1988 prov. ed.) USGS tqm. 29 November 1991. Collector: M.J. Wetzel (#1991-85) (INHS).

Spring Canyon Creek, 0.4 km upstream "confluence" with Colorado River (RM 204.4R), in Grand Canyon National Park. U.T.M. Zone 12: 287850m East, 3988340m North. Elevation: 1,510ft. Whitmore Point SE, Arizona (1967 ed.) USGS tqm. 30 November 1991. Collector: M.J. Wetzel (#1991-88) (INHS).

Diamond Creek, 30m upstream confluence with Colorado River (RM 225.7L), just S of boat landing area, in Lower Granite Gorge, Lake Mead National Recreation Area, in Grand Canyon National Park. U.T.M. Zone 12: 285540m East, 3960310m North. Elevation: 1,345ft. Diamond Peak, (1967 ed.) USGS tqm. 3 December 1991. Collector: M.J. Wetzel (#1991-90) (INHS).


1993 Collections. In September 1993, 23 aquatic sampling sites were surveyed. These included five sites on the main stem of the Colorado River (including backwater bays); Warm Springs / Lava Well, Pumpkin Springs, and five unnamed backwater marsh areas that drain into the Colorado River; and 16 direct tributary streams of the Colorado River (Nankoweap Creek, Bright Angel Creek, Hermit Creek, Crystal Creek, Shinumu Creek, Royal Arch Creek, 128 Mile Creek, Stone Creek, Tapeats Creek, Deer Creek, Kanab Creek, Matkatamiba Canyon Creek, Havasu Creek, Spring Canyon Creek, Three Springs Canyon Creek and seep, and Diamond Creek. Aquatic oligochaetes were collected from 13 of these 23 sites. 1993 sites located in Coconino County:Colorado River (COR RM 31.65R), in moist sand just above water line, along north edge of South Canyon Bay, 110 m downstream South Canyon Creek Rapids (280m NE Stanton's Cave), upstream (around bend from) Vasey's Paradise. U.T.M. Zone 12: 423250m East, 4039820m North. Elevation: 2,870ft. North Canyon Point, ARIZ. (1985 prov. ed.) USGS tqm. 15 September 1993. Collectors: M.J. Wetzel (#1993-41) (INHS); L.E. Stevens, J. Stanford, J. Pflueger.Colorado River (COR RM 31.65R), along north edge of river, in South Canyon Bay, 110 m downstream South Canyon Creek Rapids (280m NE Stanton's Cave), upstream (around bend from) Vasey's Paradise. U.T.M. Zone 12: 423255 m East, 4039815m North. Elevation: 2,870 ft. North Canyon Point, ARIZ. (1985 prov. ed.) USGS tqm. 15 September 1993. Collectors: M.J. Wetzel (#1993-42 a,b) (INHS), with L.E. Stevens, J. Pflueger.Backwater marsh #1, in drainage 'channel' just 'upstream' (SSE) of inlet / bay created by sand spit along east side Colorado River (COR RM 43.5L), 150m upstream of President Harding Rapids, just ENE Point Hansbrough, in Marble Canyon, Grand Canyon National Park. U.T.M. Zone 12: 423970m East, 4027900m North. Elevation: 2,835ft. Tatahatso Point, Arizona (1988 prov. ed.) USGS tqm. 2 September 1993. Collectors: M.J. Wetzel (#1993-43), M.L. Biyal (INHS).Backwater marsh #2up, at upstream end of drainage into bay created by sand spit, 50-70m ESE bay outlet to Colorado River (COR RM 50.0L), in Marble Canyon, Grand Canyon National Park. U.T.M. Zone 12: 422970m East, 4021400m North. Elevation: 2,840ft. Nankoweap Mesa, Arizona (1988 prov. ed.) USGS tqm. 2 September 1993. Collectors: M.J. Wetzel (#1993-44), M.L. Biyal (INHS).Backwater marsh #2dn, downstream site, near mouth of bay created by sand spit, 5-15m ESE bay outlet to Coloradao River (COR RM 50.0L), in Marble Canyon, Grand Canyon National Park. U.T.M. Zone 12: 422920m East, 4021430m North. Elevation: 2,840ft. Nankoweap Mesa, Arizona (1988 prov. ed.) USGS tqm. 2 September 1993. Collectors: M.J. Wetzel (#1993-45), M.L. Biyal (INHS).Nankoweap Creek, 200m upstream (SW) confluence with Colorado River (COR RM 52.2L) at Nankoweap Rapids, Grand Canyon National Park. U.T.M. Zone 12: 422900m East, 4018620m North. Elevation: 2,830ft. Nankoweap Mesa, Arizona (1988 prov. ed.) USGS tqm. 2 September 1993. Collectors: M.J. Wetzel (#1993-46), M.L. Biyal (INHS); C.L. Graham (NAU volunteer).Nankoweap Bay marsh, on north side of Nankoweap Bay, just west of the lower set of Nankoweap Rapids; marsh drains into north side of this bay of Colorado River (COR RM 52.7L), Grand Canyon National Park. U.T.M. Zone 12: 423080m East, 4017430m North. Elevation: 2,787ft. Nankoweap Mesa, Arizona (1988 prov. ed.) USGS tqm. 2 September 1993. Collector: M.J. Wetzel (#1993-47) (INHS).Backwater slough / marsh, Colorado River (COR RM 55.5R), 0.5 mi (0.8 km) upstream (NW) Kwagunt Creek / rapids, Colorado River (COR RM 55.5R); Grand Canyon National Park. U.T.M. Zone 12: 425150m East, 4013930m North. Elevation: 2,795ft. Nankoweap Mesa, Arizona (1988 prov. ed.) USGS tqm. 3 September 1993. Collectors: M.J. Wetzel (#1993-48), M.L. Biyal (INHS).Bright Angel Creek, 550m upstream confluence with Colorado River (COR RM 87.8R), adjacent to North Kaibab Trail and Phantom Ranch campground, 80m downstream footbridge across creek; Grand Canyon National Park. U.T.M. Zone 12: 401420m East, 3995590m North. Elevation: 2,485ft. Desert View, ARIZ. (1988 prov. ed.) USGS tqm. 5 September 1993. Collectors: M.J. Wetzel (#1993-49), M.L. Biyal (INHS).Hermit Creek, 90m upstream confluence with Colorado River (COR RM 95.0L); Grand Canyon National Park. U.T.M. Zone 12: 391180m East, 3995400m North. Elevation: 2,360ft. Grand Canyon, ARIZ. (1988 prov. ed.) USGS tqm. 6 September 1993. Collectors: M.J. Wetzel (#1993-50), M.L. Biyal (INHS).Crystal Creek, 30-50m upstream confluence with Colorado River (COR RM 98.15R); in Granite Gorge, Grand Canyon National Park. U.T.M. Zone 12: 388140m East, 3999470m North. Elevation: 2,350ft. Shiva Temple ARIZ. (1988 prov. ed.) USGS tqm. 6 September 1993. Collectors: M.J. Wetzel (#1993-51), M.L. Biyal (INHS).Shinumo Creek, 70-120m upstream (N) of confluence with Colorado River (COR RM 108.6R); downstream of waterfall/pool; in Granite Gorge, Grand Canyon National Park. U.T.M. Zone 12: 378820m East, 4011000m North. Elevation: 2,202ft. Havasu Point, Arizona (1988 prov. ed.) USGS tqm. 6 September 1993. Collectors: M.J. Wetzel (#1993-52), M.L. Biyal (INHS).Royal Arch Creek and pools, 120m upstream (S) confluence with Colorado River (COR RM 116.6L); in Granite Gorge, Grand Canyon National Park. U.T.M. Zone 12: 369640m East, 4006515m North. Elevation: 2,200ft. Explorers Monument, Arizona (1988 prov. ed.) USGS tqm. 6 September 1993. Collectors: M.J. Wetzel (#1993-53), M.L. Biyal (INHS).128 Mile Creek, 100m upstream (SE) confluence with Colorado River (COR RM 128.5R); in Middle Granite Gorge, Grand Canyon National Park. U.T.M. Zone 12: 366710m East, 4019030m North. Elevation: 2,075ft. Powell Plateau, Arizona (1988 prov. ed.) USGS tqm. 7 September 1993. Collector: M.J. Wetzel (#1993-54) (INHS).Stone Creek, from 150 to 250m upstream confluence with Colorado River (COR RM 131.9R); Grand Canyon National Park. U.T.M. Zone 12: 369750m East, 4023260m North. Elevation: 2,070ft. Powell Plateau, Arizona (1988 prov. ed.) USGS tqm. 7 September 1993. Collectors: M.J. Wetzel (#1993-55), M.L. Biyal (INHS).Tapeats Creek, from 75-15-m upstream confluence with Colorado River (COR RM 133.7R); Grand Canyon National Park. U.T.M. Zone 12: 368330m East, 4025970m North. Elevation: 2,000ft. Powell Plateau, Arizona (1988 prov. ed.) USGS tqm. 7 September 1993. Collectors: M.J. Wetzel (#1993-56), M.L. Biyal (INHS).Deer Creek, 20-40m upstream (NNW) confluence with Colorado River (COR RM 136.2R); in Granite Narrows, Grand Canyon National Park. U.T.M. Zone 12: 364770m East, 4027950m North. Elevation: 1,940ft. Fishtail Mesa, Arizona (1988 prov. ed.) USGS tqm. 7 September 1993. Collectors: M.J. Wetzel (#1993-57), M.L. Biyal (INHS).Matkatamiba Canyon Creek, in Matkatamiba Canyon, from 100-750m upstream (S, SE) confluence of creek with Colorado River (COR RM 147.9L), just upstream Matkatamiba Rapids; Grand Canyon National Park. U.T.M. Zone 12: 350080m East, 4022960m North. Elevation: 1,930ft. Havasu Falls, Arizona (1988 prov. ed.) USGS tqm. 8 September 1993. Collectors: M.J. Wetzel (#1993-59), M.L. Biyal (INHS).Havasu Creek, 140m upstream confluence with Colorado River (COR RM 156.8L), where foot trail first crosses creek (from SW to NE side of creek); Grand Canyon National Park. U.T.M. Zone 12: 341880m East, 4019240m North. Elevation: 1,810ft. SB Point, Arizona (1988 prov. ed.) USGS tqm. 8 September 1993. Collectors: M.J. Wetzel (#1993-60), M.L. Biyal (INHS).Lava Well / Warm Springs, in pool along bluff, 60m SE Lava Falls rapid (Colorado River (COR RM 179.4L); on Hualapai Indian Reservation adjacent to Grand Canyon National Park. U.T.M. Zone 12: 312880m East, 4007520m North. Elevation: 1,665ft. Vulcan's Throne, ARIZ. (1967 ed.) USGS tqm. 9 September 1993. Collectors: M.J. Wetzel (#1993-61) (INHS), M. Shaver (NAU).Pumpkin Springs, from several sites around perimeter of pool at top of "pumpkin", Colorado River (COR RM 212.8L), at northern edge of bay / campsite; Grand Canyon National Park. U.T.M. Zone 12: 289445m East, 3976960m North. Elevation: 1,440ft. Granite Park, ARIZ. (1967 ed.) USGS tqm. 11 September 1993. Collectors: M.J. Wetzel (#1993-64), M.L. Biyal (INHS).Seep, north side Three Springs Canyon Creek, along trail east into canyon, 220m E of confluence of creek with Colorado River (COR RM 215.6L); in Lake Mead National Recreation Area. U.T.M. Zone 12: 291880m East, 3973510m North. Elevation: 1,460ft. Granite Park, ARIZ. (1967 ed.) USGS tqm. 11 September 1993. Collectors: M.J. Wetzel (#1993-65), M.L. Biyal (INHS).Three Springs Canyon Creek, 330m upstream (E) of confluence of creek with Colorado River (COR RM 215.6L), where trail drops down (SE) to and crosses creek, in heavy vegetation and pool below creek/boulder slides; in Lake Mead National Recreation Area. U.T.M. Zone 12: 291880m East, 3973500m North. Elevation: 1,440ft. Granite Park, ARIZ. (1967 ed.) USGS tqm. 11 September 1993. Collectors: M.J. Wetzel (#1993-66), M.L. Biyal (INHS).Colorado River (COR RM 220.15C), SE corner of cobble island (Gorilla Island?), in middle of Colorado River; in Lower Granite Gorge, Lake Mead National Recreation Area. U.T.M. Zone 12: 289930m East, 3967260m North. Elevation: 1,378ft. Diamond Peak, ARIZ. (1967 ed.) USGS tqm. 11 September 1993. Collectors: M.J. Wetzel (#1993-67) (INHS); M. Shaver (NAU), J. Stanford, B. Ellis (Univ. Montana-Flathead Biol. Sta.).1993 sites located in Mohave County:Kanab Creek, from 20-120m upstream (N, NW) confluence with Colorado River (COR RM 143.3R) at Kanab Rapids; Grand Canyon National Park. U.T.M. Zone 12: 353890m East, 4028550m North. Elevation: 1,890ft. Kanab Point, Arizona (1988 prov. ed.) USGS tqm. 8 September 1993. Collectors: M.J. Wetzel (#1993-58), M.L. Biyal (INHS).Backwater bay / marsh outflow channel, Colorado River (COR RM 204.0R), in Grand Canyon National Park. U.T.M. Zone 12: 288220m East, 3988880m North. Elevation: 1,500 ft. Whitmore Point SE, ARIZ. (7.5' series, 1967 edition; NAD1927) quad map. 10 September 1993. Collectors: M.J. Wetzel (#1993-62) (INHS), with G. Richard "Dick" Marzoff (USGS), and T. Brinton (volunteer).Spring Canyon Creek, 150-250m upstream 'confluence' with Colorado River (COR RM 204.3R); Grand Canyon National Park. U.T.M. Zone 12: 288060m East, 3988380m North. Elevation: 1,500ft. Whitmore Point SE, ARIZ. (1967 ed.) USGS tqm. 10 September 1993. Collector: M.J. Wetzel (#1993-63) (INHS).Diamond Creek, 10-50m upstream confluence with Colorado River (COR RM 225.6L), at takeout point of COR raft expedition, near terminus of road (S) to Peach Springs; in Lower Granite Gorge, Lake Mead National Recreation Area. U.T.M. Zone 12: 285540m East, 3960310m North. Elevation: 1,345ft. Diamond Peak, Arizona (1967 ed.) USGS tqm. 12 September 1993. Collector: M.J. Wetzel (#1993-68) (INHS).


2001 Collections. Between 19 and 29 June 2001, 18 aquatic sampling sites were surveyed.Seep and spring habitats:-13 Mile Spring/seep (COR RM -13L, ~1mi downstream of the Glen Canyon Dam, 13 mi upstream Lee's Ferry);an unnamed spring at Fence's Fault (COR RM 30.3L);Vasey's Paradise (hanging spring garden, COR RM 31.6R);Whale's Armpit (four seep drip pots, COR RM 34.7R);seeps at Lava Well / Warm Springs, along edge of Colorado River at Lava Rapids (COR RM 179.4L);Beecher Spring (COR RM 183.5L).Additional sites included:small backwater slough of the Colorado River at Tanner Flats (COR RM 68.5R);andeleven small to large tributary streams of the Colorado River:Paria River (COR RM 0.8R);Little Colorado River (COR RM 61.3L);Lava Chuar Creek (COR RM 65.5R);Clear Creek (COR RM 84.1R);Bright Angel Creek (COR RM 87.7R);Crystal Creek (COR RM 98.1R);127 Mile Creek (COR RM 126.8R);Tapeats Creek (COR RM 133.7R);140 Mile Creek (COR RM 139.8L);Kanab Creek (COR RM 143.4R);Spring Canyon Creek (COR RM 204.4R).More specific locality and collecting information for each site will be provided here soon. Digital photographs of the sites also will be included.


2006 Collections. Between 10 and 18 July 2006, 19 stream and spring habitats were visited; of these, extensive stream assessments following the AZ Department of Environmental Quality stream assessment protocols [AZDEQ] were completed at 10 of the sites. Water quality samples [WQ] were collected from seven additional sites. Surveys for aquatic annelids [AqAnnel] were conducted at 15 of these sites.Vasey's Paradise (hanging spring garden, COR RM 31.6R);10 July: AqAnnel, WQ.Whale's Armpit (four seep drip pots, COR RM 34.7R); 11 July: AqAnnel.Nankoweap Creek (COR RM 52.2R); 11 July: AZDEQ, AqAnnel, WQ.Lava Chuar Creek (COR RM 65.5R); 11 July: WQ only.Clear Creek (COR RM 84.1R); 12 July: recently flashed, thus no assessment; AqAnnel.Bright Angel Creek (COR RM 87.7R); 12 July: recently flashed, thus no assessment or collection.Monument Creek (COR RM 93.5L); 13 July: AZDEQ, AqAnnel, WQ.Hermit Creek (COR RM 95.0L); 13 July: AZDEQ, AqAnnel, WQ.Crystal Creek (COR RM 98.1R); 14 July: AZDEQ, AqAnnel, WQ.Shinumo Creek (COR RM 108.6R); 14 July: AZDEQ, AqAnnel, WQ.Royal Arch Creek and pools (COR RM 116.6L); 15 July: AZDEQ, AqAnnel, WQ.Tapeats Creek (COR RM 133.7R); 15 July: AZDEQ, AqAnnel, WQ.Deer Creek - pool below falls, and outflow stream (COR RM 136.2R); 15 July: AZDEQ, AqAnnel, WQ.Kanab Creek (COR RM 143.4R); 16 July: AZDEQ, AqAnnel, WQ.Matkatamiba Creek (COR RM 179.9L); 16 July: AqAnnel only.the Ledges seep (COR RM ~151.55R); 17 July: WQ only.National Canyon Creek (COR RM 166.4L); 16 July: AZDEQ, AqAnnel, WQ.Spring Canyon Creek (COR RM 204.4R); 17 July: AZDEQ, AqAnnel, WQ.Pumpkin Spring (COR RM 204.4R); 17 July: WQ only.Diamond Creek (COR RM 225.65L) [>1.5 km upstream (south) confluence with COR]; 18 July: AqAnnel, WQ.More specific locality and collecting information for each site will be provided here soon. Digital photographs of the sites also will be included.


Field sampling and laboratory methodologies.Physical and chemical parameters were measured at each site immediately prior to habitat characterizations and biological collections.  Parameters measured ambient and water temperatures (in °Celsius, using red liquid thermometer); dissolved oxygen (in mg/l), percent DO saturation, conductivity, hydrogen ion concentration (as pH), ORP, and barometric pressure were all measured simultaneously using the Yellow Springs Instruments Model 556 MPS multimeter.Aquatic oligochaetes (and other macroinvertebrates) were collected qualitatively using standard D-ring dip nets and small, fine-meshed aquarium dip nets.  Dip net collections for specimens were supplemented by the hand-picking of various natural substrates (gravel, cobble, boulders, sticks, branches, logs, leaf packets, and root mats).  Riffle, run, pool, backwater, and cascade habitats were agitated by hand and foot, with dislodged material drifting into dip nets.  Additionally, aquatic vegetation, detritus, and other substrates (gravel, cobble, detritus) were rinsed in buckets of habitat water to dislodge oligochaetes, followed by the straining of rinse water through fine-meshed Nytexcloth or an aquarium net.  Most samples containing oligochaete specimens were relaxed immediately after collection in a weak ethanol solution, then fixed with buffered formalin.  Aquatic insects, when separated from other material after capture in the nets or buckets, were preserved in 70% ethanol.  Mollusks, when observed, were preserved in 70% ethanol; ancylids, sphaeriids, and a few gastropod species were collected from several  sites, but neither live specimens nor shell material of the pelecypod families Unionidae, Corbiculidae, and Dreissenidae was observed a any of the sites visited in 1991, 1993, or 2001.  A priority during our surveys was to minimize physical disturbance to streams and other sites from which aquatic macroinvertebrates were collected.  In particular, extreme care was taken during our abbreviated collecting efforts for aquatic macroinvertebrates in the finite habitats associated with the small streams and rock pools.  Every effort was made to cull live vertebrate specimens (fishes and amphibians) from netted material, immediately returning them to the specific habitat from which they were obtained.After field samples were brought to the INHS laboratory, and initial processing of samples was completed, all samples were labeled and stored in 70% ethanol in  vials or jars appropriate for the specimen.  Oligochaetes were dehydrated through an ethanol series, cleared for 5-10 min in a 1:1 ethanol-xylene solution, mounted under coverslips on glass slides in Canada Balsam or Permount, and identified using an Olympus BH-2 or BX-50 compound microscope equipped with Nomarski DIC.  Nomenclature and taxonomic interpretations follow Reynolds and Cook (1976, 1981, 1989, 1993), Kathman and Brinkhurst (1998), Reynolds and Wetzel (2006, in prep), Wetzel et al. (2006) and those published most recently in the literature; the most current classification and list of species known to occur in North America is posted HERE.  All specimens obtained during this study are deposited in the Illinois Natural History Survey Annelida Collection, Champaign.<![if !vml]><![endif]>- Preliminary Checklist -
Aquatic Annelida occurring in the Colorado River and its tributaries
Grand Canyon National Park, Arizona (GCNP)
Introduction to the List. There are no historical records in the published literature documenting the occurrence of aquatic Annelida in the Colorado River and its tributaries within the Grand Canyon National Park. A survey of the invertebrate faunas of three stream caves in the Grand Canyon by Peck (1980) yielded neither aquatic annelids nor other aquatic invertebrates. Several recent papers relating to this current research on benthic dynamics of the Colorado River have noted the presence of Oligochaeta in the river and its tributaries (Benenati et al. 2000; Blinn et al. 1995; Blinn et al. 1999; Shannon et al. 1994; Shannon et al. 1996; Shaver et al. 1997; Stevens et al. 1997; Oberlin et al. 1999) but none provided identifications beyond that of Class Oligochaeta or the families Lumbricidae, Lumbriculidae, Naididae, and Tubificidae. Wetzel et al. (1999) published the first records for aquatic oligochaetes of Montezuma Well, Arizona; in that paper, the authors reviewed past records of specimens from elsewhere in Arizona, and recognized 10 species as new for the state. Historical and recent records of aquatic oligochaetes occurring in North America are noted in Brinkhurst and Jamieson (1971), Brinkhurst and Wetzel (1984), Klemm (1985, 1995), Brinkhurst (1986), Brinkhurst and Gelder (1991, 2001), Kathman and Brinkhurst (1998), Wetzel et al. (2006a, 2006b, 2009), Coates and Wetzel (unpublished); INHS Annelida Collection records; and numerous other published papers.The binomial nomenclature of aquatic annelids provided below reflect recent nomenclatural and systematic changes, and correct date-of-publication errors that had been perpetuated in past literature. References noted in the text of this document are included in the 'Literature Cited' section which follows the checklist of species.Please note that this is a working document. Information pertaining to the distribution of aquatic annelids in or adjacent to the GCNP will be added as it becomes available. The list below has been compiled from identifications completed on specimens collected in November and December 1991; specimens collected in September 1993, June 2001, and July 2006 currently are being mounted on microscope slides prior to identification.To date, no representatives of the freshwater annelid groups Aeolosomatida (suction-feeding worms) or Branchiobdellida (crayfish worms) have been collected from aquatic habitats within the GCNP (sites described above).


P H Y L U M - A N N E L I D ACLASS OLIGOCHAETA
Superorder Microdrili
Order Lumbriculida

FAMILY LUMBRICULIDAE

species yet to be determined.

Order Enchytraeida
Suborder Enchytraeina
Superfamily Enchytraeoidea

FAMILY ENCHYTRAEIDAE

species yet to be determined.

Order Tubificida
Suborder Tubificina
Superfamily Tubificoidea

FAMILY NAIDIDAE

Subfamily Naidinae

Genus Chaetogaster von Baer, 1827
     Chaetogaster diaphanus (Gruithuisen, 1828)

Genus Nais Müller, 1773

? Nais barbata Müller, 1773
Nais communis Piguet, 1906

Nais elinguis Müller, 1773

Nais pardalis Piguet, 1906

Nais pseudobtusa Piguet, 1906

Nais variabilis Piguet, 1906

Genus Ophidonais Gervais, 1838

Ophidonais serpentina (Müllerller, 1773)

Subfamily Pristininae

Genus Pristina Ehrenberg, 1828 emended Brinkhurst, 1984

Pristina aequiseta Bourne, 1901

Pristina osborni (Walton, 1906)

Pristina sp.; yet to be determined

Subfamily Tubificinae

Genus Limnodrilus Claparède, 1862

Limnodrilus claparedeianus Ratzel, 1868

Limnodrilus hoffmeisteri Claparède, 1862

Genus Tubifex Lamarck, 1816

Tubifex tubifex (Müller, 1774)

<![if !supportEmptyParas]> <![endif]>>>>

Please Note: the genus Pristinella, established by Brinkhurst (1985), was suppressed by Collado and Schmelz (2000) based upon their description of two new species in the genus Pristina, and their reassessment of the morphology of Pristina and Pristinella.

* Current Status of the Family Naididae.

Order Opisthopora
Suborder Lumbricina
Superfamily Lumbricoidea

FAMILY LUMBRICIDAE

Genus Eiseniella Michaelsen, 1900

Eiseniella tetraedra (Savigny, 1826)

SUBCLASS HIRUDINIDA
Order Arhynchobdellida

FAMILY ERPOBDELLIDAE

Genus Erpobdella de Blainville, 1818

Erpobdella punctata (Leidy, 1870)

 

 

LITERATURE CITED

Benenati, E.P.; Shannon, J.P.; Blinn, D.W.; Wilson, K.P.; Hueftle, S.J.  2000. Reservoir-river linkages: Lake Powell and the Colorado River, Arizona. Journal of the North American Benthological Society 19(4): 742-755.

Blinn, D.W., J.P. Shannon, L.E. Stevens, and J.P. Carder. 1995. Consequences of fluctuating discharge for lotic communities. Journal of the North American Benthological Society 14: 233-248.

Blinn, D.W., J.P. Shannon, K.P. Wilson, C. O'Brien, and P.L. Benenati. 1999. Response of benthos and organic drift to a controlled flood. Pp. 259-272, In: R.H. Webb, J.C. Schmidt, G.R. Marzolf, & R.A. Valdez (eds.). The controlled flood in Grand Canyon. Geophysical Union Monograph 110, American Geophysical Union, Washington, D.C.

Brinkhurst, R O. 1985. The generic and subfamilial classification of Naididae (Annelida: Oligochaeta). Proceedings of the Biological Society of Washington 98(2): 470-475.

Brinkhurst, R.O.  1986.  Guide to the freshwater aquatic microdrile oligochaetes of North America.  Canadian Special Publication in Fisheries and Aquatic Sciences  84.  vi + 259 pp. 

Brinkhurst, R.O., and S.R. Gelder. 1991. Annelida: Oligochaeta and Branchiobdellida. Pp. 401-435, In: J.H. Thorp and A.P. Covich, eds. Ecology and classification of North American freshwater invertebrates. Academic Press - Harcourt Brace Jovanovich, Publishers, San Diego, CA, USA. [ Citation for second edition noted below ].

Brinkhurst, R.O., and S.R. Gelder. 2001. Annelida: Oligochaeta, including Branchiobdellidae. Pp. 431-463, In: J.H. Thorp and A.P. Covich, eds. Ecology and classification of North American freshwater invertebrates, Second Edition. Academic Press, San Diego, CA, USA. [To order, contact Academic Press, Book Marketing Dept. #10520, 1250 Sixth Avenue, San Diego, California, 92101-9665 USA. Tel.: 1-800-321-5068.

Brinkhurst, R.O., and B.G M. Jamieson. 1971: Aquatic Oligochaeta of the world. University of Toronto Press, Toronto, ON, and Buffalo, NY. xi + 860 pp. [out of print].

Brinkhurst, R.O., and M.J. Wetzel. 1984. Aquatic Oligochaeta of the world: Supplement. A catalogue of new freshwater species, descriptions, and revisions. Canadian Tech. Rpt. Hydrography Ocean Sci. No. 44. v + 101 pp.

Coates, K.A., and M.J. Wetzel, editors. Manuscript in prep. Common and scientific names of aquatic invertebrates from the United States and Canada: Clitellate annelids, including Aphanoneura, Branchiobdellae, Hirudinea, and Oligochaeta.

Collado, R., and R.M. Schmelz. 2000. Pristina silvicola and Pristina terrena spp. nov., two new soil-dwelling species of Naididae (Oligochaeta, Annelida) from the tropical rain forest near Manaus, Brazil, with comments on the genus Pristinella. Journal of Zoology, London 252: 509-516.

Erséus, C., and L. Gustavsson. 2002. A proposal to regard the former family Naididae as a subfamily within Tubificidae (Annelida, Clitellata). Hydrobiologia 485: 253-256.

Erséus, C., L. Gustavsson, and R.O. Brinkhurst. 2005. Case 3305. Tubificidae Vejdovsky, 1876 (Annelida, Clitellata): proposed precedence over Naididae Ehrenberg, 1828. Bulletin of Zoological Nomenclature 62(4):226-231.

Erséus, C., and C.M. Källersjö. 2004. 18S rDNA phylogeny of Clitellata (Annelida). Zoologica Scripta 33(2): 187-196.

Erséus, C.M. Källersjö, M. Ekman, and R. Hovmöller. 2002. 18S rDNA phylogeny of the Tubificidae (Clitellata) and its constituent taxa: dismissal of the Naididae. Molecular Phylogenetics and Evolution 22(3): 414-422.

Erséus, C., M.J. Wetzel, and L. Gustavsson. 2008. ICZN rules--a farewell to Tubificidae (Annelida, Clitellata). Zootaxa 1744: 66-68.

Kathman, R.D., and R.O. Brinkhurst. 1998. Guide to the freshwater oligochaetes of North America. Aquatic Resources Center, College Grove, TN 37046 USA. iv + 264 pp. USD $55.00 (post paid). A link to ordering information for this guide is posted HERE.

Klemm, D. J., ed. 1985. A guide to the freshwater Annelida (Polychaeta, naidid and tubificid Oligochaeta, and Hirudinea) of North America. Kendall/Hunt Publishing Company, Dubuque, IA. xiii + 198 pp. [now out of print, but copies may be available through used book sellers]

Klemm, D. J. 1995. Identification guide to the freshwater leeches (Annelida: Hirudinea) of Florida and other southern states. Florida Department of Environmental Protection (FDEP), Division of Water Facilities. Bureau of Surface Water Management, Twin Towers Office Building, 2600 Blair Stone Road, Tallahassee, FL 32399-2400 USA. [This guide is available, without charge, by visiting the following website: FDEP Bureau of Laboratories List of Keys and Guides, and then scrolling down this page to 'Leeches.pdf' to download a pdf version of this key, or to the next entry on this page - 'Leechx.exe' to receive a self-extracting .zip to html file of this same document.  

Oberlin, G.E., J.P. Shannon, and D.W. Blinn. 1999. Watershed influence on the macroinvertebrate fauna of ten major tributaries of the Colorado River through Grand Canyon, Arizona. Southwestern Naturalist 44(1): 17-30.

Peck, S.B. 1980. Climatic change and the evolution of cave invertebrates in the Grand Canyon, Arizona. NSS Bulletin 42: 53-60.

Reynolds, J. W., and D. G. Cook. 1976. Nomenclatura oligochaetologica. A catalogue of names, descriptions and type specimens of the Oligochaeta. Univ. New Brunswick, Fredericton, New Brunswick. x + 217 pp. [This catalog, and the first supplement, cited below, are out of print].

Reynolds, J. W., and D. G. Cook. 1981. Nomenclatura oligochaetologica. Supplementum primum. A catalogue of names, descriptions and type specimens of the Oligochaeta. [published by the University of New Brunswick] Fredericton, New Brunswick. v + 39 pp. [out of print].

Reynolds, J. W., and D. G. Cook. 1989. Nomenclatura oligochaetologica. Supplementum secundum. A catalogue of names, descriptions and type specimens of the Oligochaeta. New Brunswick Mus. Monogr. Ser. (Nat. Sci.) No. 8. v + 37 pp. [this second supplement is available, at no charge, from the New Brunswick Museum, 277 Douglas Avenue, Saint John, NB, Canada E2K 1E5.].

Reynolds, J. W., and D. G. Cook. 1993. Nomenclatura oligochaetologica. Supplementum tertium. A catalogue of names, descriptions and type specimens of the Oligochaeta. New Brunswick Mus. Monogr. Ser. (Nat. Sci.) No. 9. vi + 33 pp. [this third supplement is available, at no charge, from the New Brunswick Museum, 277 Douglas Avenue, Saint John, NB, Canada E2K 1E5.].

Reynolds, J.W. and M.J. Wetzel. 2004. Terrestrial Oligochaeta (Annelida: Clitellata) in North America north of Mexico. Megadrilogica 9(11): 71-98.

Reynolds, J.W. and M.J. Wetzel. 2006. Nomenclatura oligochaetologica. Supplementum quartum. A catalogue of names, descriptions and type specimens of the Oligochaeta. In Prep.

Shannon, J.P., D.W. Blinn, and L.E. Stevens. 1994. Trophic interactions and benthic animal comunity structure in the Colorado River, Arizona. Freshwater Biology 31: 213-220.

Shannon, J.P., D.W. Blinn, P.L. Benenati, and K.P. Wilson. 1996. Organic drift in a regulated desert River. Canadian Journal of Fisheries and Aquatic Sciences 53: 1360-1363.

Shaver, M.L., J.P. Shannon, K.P. Wilson, P.L. Benenati, and D.W. Blinn. 1997. Effects of suspended sediment and desiccation on the benthic tailwater community in the Colorado River. Hydrobiologia 357: 63-72.

Stephens, L. 1983. "The Colorado River Grand Canyon - A comprehensive guide to its natural and human history." Third Edition. Red Lake Books, P.O. Box 1315, Flagstaff, AZ 86002. 110 pp.

Stevens, L.E., J.P. Shannon, andd D.W. Blinn. 1997. Colorado River benthic ecology in Grand Canyon, Arizona, USA: dam, tributary and geomorphic influences. Regulated Rivers: Research & Management 13: 129-149.

Wetzel, M.J., G.E. Oberlin, and D.W. Blinn. 1999. The aquatic Oligochaeta (Annelida: Clitellata) of Montezuma Well, Arizona: a near thermally constant limnocrene. Southwestern Naturalist 44(4): 514-518.

Wetzel, M.J., S.V. Fend, K.A. Coates, R.D. Kathman, and S.R. Gelder. 2006a,b. Taxonomy, systematics, and ecology of the aquatic Oligochaeta and Branchiobdellidae (Annelida, Clitellata) of North America, with emphasis on the fauna occurring in Florida. A workbook prepared for two workshops: a) underwritten by / covened at the Florida Department of Environmental Protection, Tallahassee, 21-23 February 2006, and b) underwritten by the Florida Association of Benthologists, convened at the University of Florida Department of Entomology and Nematology, 9-11 May 2006. [vi + 269 pp. + 4 color plates].

Wetzel, M.J., S.V. Fend, K.A. Coates, R.D. Kathman, and S.R. Gelder. 2009. Taxonomy, systematics, and ecology of the freshwater oligochaetes and branchiobdellidans (Annelida, Clitellata) of North America. A workbook. 3 March. vi + 280 pp. + 4 color plates. Copyright (c) 2009. {The overall scope of this workbook is the North American continent; this serves as a compliment to the most recent guide published by R.D. Kathman and R. Brinkhurst in 1998.}
To purchase a copy of this workbook, first send an *E-mail to {mjwetzel'{AT}'illinois.edu} expressing your interest in this workbook / and to ascertain its availability; note that payment must be in form of personal check or money order -- credit card payments not accepted).

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Copyright 2000-2009, by Mark J. Wetzel (Illinois Natural History Survey, Champaign) unless otherwise noted; All Rights Reserved. This website contains original, copyrighted material; it is being provided here as a professional courtesy, exclusively for your private, non-commercial use. Reference to or redistribution of any part of the information contained herein - whether it be through oral, printed, electronic, or other tangible medium of expression - shall acknowledge the Illinois Natural History Survey (INHS) and this website as its source, and shall acknowledge the U.S. Department of the Interior Bureau of Reclamation, Glen Canyon Environmental Studies Program, the Grand Canyon Monitoring and Research Center� U.S. Geological Survey, Grand Canyon National Park, and Glen Canyon National Recreation Area, and Northern Arizona University for their long-term support of this research. Should you have any questions whatsoever regarding the warranty, liability, or proprietary rights of, or credits for information contained within this website, please refer to the INHS License Agreement. [To the best of my knowledge - and with the exception of the INHS logo - all icons, line breaks, dots, arrows, and globes are not copyrighted.]


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Suggested citation for this electronic web page:

Wetzel, M. J. 2009. The Aquatic Annelida of the Colorado River and its tributaries, Grand Canyon National Park, Arizona. World Wide Web URL:
[ http://www.inhs.uiuc.edu:80/~mjwetzel/AqAnnel.GCNP.html ]. 15 April 2009.

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