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Western North American Naturalist 62(4), © 2002, pp. 385–404
STONEFLIES (PLECOPTERA) OF MOUNT
RAINIER NATIONAL PARK, WASHINGTON
B.C. Kondratieff1 and Richard A. Lechleitner2
ABSTRACT.—Mount Rainier National Park, with an area of 95,356 ha, is approximately one-third as large as the state
of Rhode Island. The lowest point is 490 m in elevation in the southeastern corner near where the Ohanapecosh River
crosses the southern boundary. Columbia Crest is the highest point at 4392 m. The entire park is a rugged landscape
marked by the major topographical feature, Mount Rainier, comprising over 25,899 ha, almost one-third of the park. The
park lies entirely west of the crest line of the Cascade Range. Most streams in the park originate on Mount Rainier;
however, several large rivers meander through the park near its boundaries. One of the first attempts to summarize the
stoneflies of Washington, including Mount Rainier National Park, was Hoppe’s 1938 work that reported ca 8 species.
Jewett (1959) reviewed the stoneflies of the Pacific Northwest and listed 7 species that had type localities in the park:
Megaleuctra kincaidi Frison, Doddsia occidentalis (Banks), Soliperla fenderi ( Jewett), Frisonia picticeps (Hanson), Isoperla
rainiera Jewett, Megarcys irregularis (Banks), and M. subtruncata (Hanson). Subsequently, Kathroperla takhoma Stark
and Surdick (1987) was described from the park. Samples of adult stoneflies from 1994 to 2001 indicate the presence of
at least 82 species, with 64% of these typical Pacific Northwest species, and 30 species, or 36%, widespread western
North American species. Seventeen new Washington state records are listed, including a substantial range extension for
Lednia tumana (Ricker). One undescribed species in the Sweltsa borealis complex was also discovered. We also present
illustrations of male terminalia for Despaxia augusta (Banks) and Moselia infuscata (Claassen) to aid in the identification
of these species.
Key words: stoneflies, Plecoptera, Mount Rainier National Park, Washington.
Mount Rainier National Park is in Lewis
and Pierce counties on the western slope of
the Cascade Range of Washington State. Mount
Rainier is located approximately 65–110 km
from the Seattle-Tacoma metropolitan area
known as Puget Sound (Fig. 1). The park area
is 95,356 ha, and elevations range from 490 m
above sea level to 4392 m at the summit.
Mount Rainier, one of the highest and topographically most impressive of the world’s
volcanoes, owes its scenic beauty to many features. The broad cone rises about 2300 m
above its 2100-m foundation and stands in
solitary splendor as the highest peak in the
Cascade Range. Its rocky ice/snow-mantled
slopes above timberline contrast with the dense
green forests, giving Mount Rainier the appearance of an arctic island in a temperate sea.
There are 26 major glaciers on Mount Rainier
and numerous unnamed snow or ice patches.
Mount Rainier was born during the Pleistocene, and gradually the high main cone was
built up. As eruptions diminished, Mount
Rainier began to deteriorate by explosion,
collapse, and erosion. It has been estimated
that the cone was once about 600 m higher than
it is now. Even before Mount Rainier reached
its greatest height, rivers and glaciers were
cutting deep valleys and huge, bowl-shaped
cirques into its sides. During the last major
glaciation, which ended about 10,000 years
ago, the valley glaciers grew to as much as 60
km long, and smaller glaciers nestled in cirques
above 1400 m. The peak has essentially remained the same in appearance since the last
major glaciations. The last major eruption of
the volcano occurred about 2000 years ago,
although the last eruption was approximately
150 years ago (Mullineaux 1974).
Mount Rainier is situated within a temperate, maritime climate. Several climatic zones
exist elevationally and geographically around
the park. However, the east-southeast side of
the park is generally the driest, and the northwest side is the wettest sector (especially during spring and summer months). Annual precipitation is heavy, ranging from about 1.5 m at
lowest elevations to over 2.5 m in the subalpine
1Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, CO 80523.
2Mount Rainier National Park, Tahoma Woods, Star Route, Ashford, WA 98304.
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WESTERN NORTH AMERICAN NATURALIST
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Fig. 1. Map of western Washington, indicating Mount Rainier National Park, Washington.
regions. Because southwesterly winds bear
much of the moisture, a rain shadow occurs on
the northeast side of the park. Over 90% of the
precipitation occurs between November and
April. Much of the winter precipitation is snow
that accumulates into depths of 6 to 8 m at
higher elevations. At Paradise the average
annual snowfall is about 17 m, with over 26 m
of snow during the winter of 1998–99. Winter
temperatures are relatively warm (mean January
temperatures of about –4° to –1°C). Summers
tend to be cool (mean July temperatures of 10°
to 20°C), and extended periods of cloudiness
are not uncommon. July and August are usually comparatively dry. Fog and high winds may
be expected any day of the year.
The topography of the park is rugged and
precipitous, consisting mainly of peaks and
valleys. Nine major rivers and their tributaries
drain the flanks of the mountain, with all but 2
flowing into Puget Sound near Tacoma, Washington. The Muddy Fork and Ohanapecosh
rivers are the exception, flowing into the Cowlitz River, outside the park, and then draining
into the Columbia River and on to the Pacific
Ocean. Each major river occupies a deep canyon, its floor 300–1000 m below the adjacent
divides. Valley floor gradients are steep and increase markedly upstream, especially in Tahoma
Creek, North and South Puyallup and Mowich
rivers. The park includes 470 mapped rivers
and streams, some 400 mapped lakes and ponds,
and over 1200 ha of other wetland types, including numerous thermal and mineral springs.
About 800 species of plants are known from
the park (Franklin et al. 1988), reflecting the
varied climatic and environmental conditions
encountered across the 3800-m elevation gradient. Approximately 58% of the park is covered
by forests, mostly old-growth stands ranging
from 200 to 1000 years old. The subalpine
parkland covers approximately 23% of the
park; vegetation in this zone is a mosaic of tree
clumps and herbaceous meadows extending
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MOUNT RAINIER NATIONAL PARK STONEFLIES
from 1500 m to about 2200 m elevation. The
alpine zone extends from treeline to the
mountain’s summit and covers approximately
19% of the park. Permanent snow and ice cover
half of the alpine zone, with fragile alpine vegetation growing on rock and soil outcrops.
According to historic records, coho (silver)
salmon, chinook, and steelhead were once present in many of the park’s major rivers before
the construction of dams outside the park.
Chinook, steelhead, and coho are now transported around some dams in an effort to restore these anadromous fish runs, thereby
allowing access to the headwaters in the park.
These fish species are thought to occur in the
White and Carbon rivers inside the park. Bull
trout are present throughout the park. In addition to native fish species, nonnative and
hatchery fish have been introduced to lakes
and streams in the park. Nonnative and hatchery fish stocks have probably altered natural
lake and stream ecosystems.
One of the first attempts to summarize the
stoneflies for the state of Washington, including
Mount Rainier National Park, was the work of
Hoppe (1938). She reported at least 8 species
from the park: Yoraperla brevis (Banks) (as Peltoperia [sic] brevis), Megarcys irregularis (Banks)
(as Perlodes irregularis), Claassenia sabulosa
(Banks) (as Perla languida Needham and Claassen), Isoperla sordida Banks, I. fulva Claassen
(as I. cascadensis Hoppe), Ostrocerca dimicki
(Claassen) (as Nemoura dimicki), Zapada cinctipes (Banks) (as Nemoura cinctipes), and
Doddsia occidentalis (Banks) (as Taeniopteryx
occidentalis). Stark and Nelson (1994) did not
report Y. brevis from Washington, and Hoppe’s
records probably refer to Y. nigrisoma (Banks)
or Y. mariana (Ricker). Hoppe’s record may be
the one originally cited by Needham and
Claassen (1925). Ostrocerca dimicki was not
collected from the park during the present
study, and the reported specimens were not
located. Her record may refer to O. foersteri
(Ricker); however, both species are known
from Washington. These small nemourids are
often difficult to specifically identify (Young et
al. 1989). Additionally, we have been unable to
confirm the presence of the widespread and
common I. fulva from the park. Hoppe (1938)
listed it as I. cascadensis from the Nisqually
River, a stream draining the park. Szczytko and
Stewart (1979) examined the female specimen.
387
Jewett (1959) in his review of the stoneflies
of the Pacific Northwest listed 7 species that
had type localities in the park: Megaleuctra
kincaidi Frison, D. occidentalis, Soliperla fenderi ( Jewett), Frisonia picticeps, (Hanson), I.
rainiera Jewett, M. irregularis, and M. subtruncata (Hanson). Szczytko and Stewart (1979)
included a paratype female for I. bifurcata
from Longmire; and Stark and Surdick (1987)
described Kathroperla takhoma from Falls
Creek. Table 1 summarizes the primary type
localities in the park. Paradise Valley was relatively easy to access in the early history of the
park, and a resort was established there.
Our study had 2 main objectives: to document the general distribution of stoneflies of
Mount Rainier National Park, and to provide
information on endemic, rare, or potentially
threatened species. Stoneflies are known to be
major indicators of water quality (Baumann
1979) and are often dominant food-web components of most temperate lotic ecosystems
(Stewart and Stark 1988). Knowledge of the
occurrence and distribution of these insects
will help serve as a basis for future biomonitoring programs as dramatic land-use changes
occur in adjacent areas.
METHODS
We sampled over 110 sites between 1994
and 2001 (Fig. 2). These sites were primarily
TABLE 1. Primary types originally described from Mount
Rainier National Park, Washington.
LEUCTRIDAE
Megaleuctra kincaidi Frison 1942. Type locality: Fryingpan Creek
TAENIOPTERYGIDAE
Doddsia occidentalis (Banks) 1900. Type locality: Mount
Rainier National Park
PELTOPERLIDAE
Soliperla fenderi ( Jewett) 1955. Type locality: St.
Andrews Creek
CHLOROPERLIDAE
Kathroperla takhoma Stark and Surdick 1987. Type
locality: Falls Creek
PERLODIDAE
Frisonia picticeps (Hanson) 1942. Type locality: Paradise
River
Isoperla rainiera Jewett 1954. Type locality: Mount
Megarcys irregularis (Banks) 1900. Type locality: Mount
Megarcys subtruncata (Hanson) 1942. Type locality:
Paradise Valley
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Fig. 2. Map of Mount Rainier National Park, Washington, indicating major streams and roads. Major collecting sites of
stoneflies during this study are indicated by ▲.
chosen to cover all relatively accessible major
drainages. Large areas of the park (Fig. 2) are
roadless or difficult to access due to extreme
topography. All adult and larval specimens
were preserved in 80% ethanol. We employed
standard collection techniques, including beating sheets, aerial nets, and collection by hand.
Specimens collected during the study were
deposited in the C.P. Gillette Museum of
Arthropod Diversity, Colorado State University. We also present illustrations of the male
terminalia for 2 common leuctrid species, Despaxia augusta (Banks) (Figs. 3, 4) and Moselia
infuscata (Claassen) (Figs. 5, 6) to aid in their
identification.
RESULTS AND DISCUSSION
We collected over 3600 adult specimens
representing at least 82 species of stoneflies in
the park during the study; they are listed in
Table 2. Sixty-four percent of these taxa are
typical Pacific Northwest species (Ricker 1943,
Jewett 1959, Ricker and Scudder 1975), and
30 species, or 36%, are widespread western
North American species (Table 3; Baumann
et al. 1977). One undescribed species in the
Sweltsa borealis complex was also discovered
and will be described later.
Seventeen new stonefly state records for
Washington were collected during the study
(Table 2; see Nelson and Baumann 1989, Stark
1998). Sixteen of these species are known from
adjacent states or British Columbia (Ricker and
Scudder 1975, Stark 1998) and were expected
records. However, Lednia tumana (Ricker) represents a major range extension. The remarkable
Nearctic nemourid genus Lednia is currently
known only in the literature from Glacier
National Park, Montana (Stewart and Stark
1988). R.W. Baumann (personal communication) indicates that specimens of this genus are
also known from North Cascades National
Park, Washington, and a site in the California
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Figs. 3–4. Despaxia augusta (Banks), male terminalia: 4, lateral; 5, dorsal.
Sierra Nevada. There may be additional relict
populations of Lednia along the Coast and
Cascade mountains of the Pacific Northwest.
Jewett (1955) described S. fenderi from a
single male collected from St. Andrews Creek
in the park. Stark (1983) reported this species
only from Mount Rainier, mostly from the St.
Andrews Creek area (B.P. Stark, personal communication, has specimens from Mount Adams,
Shamama Co.). This species can be collected
from spring seeps and rheocrenes throughout
the park (Appendix).
Isoperla rainiera was originally described
by Jewett (1954) from the park from a single
male. The female was described later from Mt.
Hood, Oregon ( Jewett 1962). Szczytko and
Stewart (1979) did not report any additional
specimens from the park. Recently, I. rainiera
was collected in Montana (S.W. Szczytko personal communication). Another rare species of
stonefly known from the park and only a few
outside localities is M. yosemite (Needham and
Claassen). This species was originally described
from Mt. Lyell, Yosemite National Park, California, and was collected during this study from
Fryingpan Creek (Van Wieren et al. 2001).
Additionally, M. irregularis, known only from
Washington and British Columbia, occurs
abundantly in the park. Ricker (1943) once
described a flight of adults at Fryingpan Creek
as “a small canyon filled with flying adults to a
depth of possibly 100 feet, resembling the
swarms of large termites. . . .”
The nymphs of Setvena tibialis (Banks) are
the dominant stonefly predators of most rheocrenes and small streams of the park. Stewart
and Stanger (1985) previously reported this
species from the park. Jewett (1959) considered
S. tibialis a rare species of the Pacific Northwest,
and collections from the park represent substantial new material for study.
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Figs. 5–6. Moselia infuscata (Claassen), male terminalia: 6, lateral; 7, dorsal.
Despaxia augusta was found to be an abundant autumn- and early winter-emerging species in the park, occurring in small streams
and rheocrenes. This species ranges from
Alaska to Montana, southwest to northern California. Ricker (1954) clarified the taxonomy of
this species. Baumann et al. (1977) indicated
adults of D. augusta are uncommonly collected,
probably because of their late emergence. Figures 3 and 4 present illustrations of the male
terminalia.
The other abundant leuctrid in the park is
Moselia infuscata. However, very little biological information is available for this species
(Stewart and Stark 1988). Nymphs are common
in most small streams of the park. Adult emergence is generally from April to July. Illustrations of the distinctive male terminalia are
presented in Figures 5 and 6.
No major distributional patterns for the 82
stonefly species were discernible for the park.
Of the species with an adequate number of
records, a few species such as Mesocapnia
oenone, Malenka californica, Doroneuria baumanni, and Kogotus nonus appear restricted to
drainages on the west side of the park (Appendix). Most other species are generally distributed throughout the park.
Despite 7 years of relatively intensive collecting, at least 11 species are represented only
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391
TABLE 2. Stonefly species recorded from Mount Rainier National Park, Washington. An asterisk (*) indicates a new
state record for Washington (see Stark [1998] and Nelson and Baumann [1989]). Zapada cinctipes (Banks) is not indicated from Washington in Stark (1998); however, the type locality of this species is Olympia, Washington.
EUHOLOGNATHA
CAPNIIDAE
Bolshecapnia sasquatchi (Ricker)*
Capnia confusa Claassen
C. elongata Claassen
C. excavata Claassen
C. gracilaria Claassen
C. licina Jewett
C. melia Frison
C. nana Claassen
C. sextuberculata Jewett
Eucapnopsis brevicauda Claassen*
Isocapnia agassizi Ricker
I. grandis (Banks)*
I. spenceri Ricker*
I. vedderensis (Ricker)*
Mesocapnia oenone (Neave)
Paracapnia oswegaptera ( Jewett)
LEUCTRIDAE
Leuctrinae
Despaxia augusta (Banks)
Moselia infuscata (Claassen)
Paraleuctra forcipata (Frison)*
P. occidentalis (Banks)*
P. projecta (Frison)
P. vershina Gaufin & Ricker
Perlomyia collaris Banks*
P. utahensis Needham & Claassen*
Megaleuctrinae
Megaleuctra kincaidi Frison
NEMOURIDAE
Amphinemurinae
Malenka californica (Claassen)*
M. cornuta (Claassen)
Nemourinae
Lednia tumana (Ricker)*
Ostrocerca foersteri (Ricker)
Podmosta decepta (Frison)*
P. delicatula (Claassen)*
Prostoia besametsa (Ricker)*
Soyedina interrupta (Claassen)
S. producta (Claassen)
Visoka cataractae (Neave)
Zapada cinctipes (Banks)
Z. columbiana (Claassen)
Z. cordillera (Baumann & Gaufin)
Z. frigida (Claassen)
Z. haysi (Ricker)
Z. oregonensis (Claassen)*
TAENIOPTERYGIDAE
Brachypteryinae
Doddsia occidentalis (Banks)
Taenionema kincaidi (Hoppe)
T. pallidum (Banks)
SYSTELLOGNATHA
CHLOROPERLIDAE
Chloroperlinae
Alloperla fraterna Frison
A. serrata Needham & Claassen
Plumiperla diversa (Frison)
Suwallia dubia (Frison)
S. forcipata (Neave)
S. pallidula (Banks)
Sweltsa borealis (Banks)
S. exquisita (Frison)
S. occidens (Frison)
S. revelstoka ( Jewett)
S. n. sp.
Paraperlinae
Kathroperla perdita (Banks)
K. takhoma Stark & Surdick
Paraperla frontalis (Banks)
P. wilsoni Ricker
PELTOPERLIDAE
Soliperla fenderi ( Jewett)
Yoraperla mariana (Ricker)
Y. nigrisoma (Banks)
Y. siletz Stark & Nelson
PERLIDAE
Doroneuria baumanni Stark & Gaufin
Hesperoperla pacifica (Banks)
Claassenia sabulosa (Banks)
PERLODIDAE
Isoperlinae
Isoperla bifurcata Szczytko & Stewart
I. fusca Needham & Claassen
I. gravitans (Needham & Claassen)
I. petersoni Needham & Christenson*
I. rainiera Jewett
I. sobria (Hagen)
I. sordida Banks
I. tilasqua Szczytko & Stewart*
Perlodinae
Arcynopterygini
Frisonia picticeps (Hanson)
Megarcys irregularis (Banks)
M. subtruncata Hanson
M. yosemite (Needham & Claassen)
Setvena tibialis (Banks)
Skwala americana (Klapalek)
Diploperlini
Kogotus nonus (Needham & Claassen)
PTERONARCYIDAE
Pteronarcys princeps Banks
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Fig. 7. Seasonal distribution of adult stoneflies occurring at Mount Rainier National Park, Washington.
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2002]
393
TABLE 3. Stonefly species occurring at Mount Rainier National Park that are considered widespread western North
American species.
Capnia confusa Claassen
C. gracilaria Claassen
C. nana Claassen
Eucapnopsis brevicauda Claassen
Isocapnia grandis (Banks)
Paraleuctra occidentalis (Banks)
P. projecta (Frison)
P. vershina Gaufin & Ricker
Perlomyia utahensis Needham & Claassen
Malenka californica (Claassen)
Podmosta decepta (Frison)
P. delicatula (Claassen)
Prostoia besametsa (Ricker)
Taenionema pallidum (Banks)
Zapada cinctipes (Banks
by single collections (Appendix). Included are
Capnia excavata, Paracapnia oswegaptera, Perlomyia utahensis, L. tumana, Hesperoperla pacifica, Claassenia sabulosa, I. bifurcata, I. fusca,
I. petersoni, I. sobria, and Frisonia picticeps.
Other populations of these species surely occur
in the park, yet to be discovered.
The emergence phenology of comparable
species of stoneflies of Mount Rainier National
Park is very similar to what Jewett (1959),
Ricker (1943), and Kerst and Anderson (1974)
reported from streams of southwestern British
Columbia and Oregon (Fig. 7). For many
species, adults can be collected for long periods. Generally, there are winter-emerging
species, including the capniids and 2 species
of Zapada, Z. cinctipes and Z. columbiana
(Fig. 7). Adults of numerous species can be
collected beginning in March and April, with
some of these species, for example Taenionema
kincaidi, still present in August. The largest
group of species begins to emerge throughout
May and June (Fig. 7). There is also a rather
distinct late summer- and fall-emerging group
of species, typified by M. oenone, D. augusta,
Alloperla serrata, and K. nonus.
ACKNOWLEDGMENTS
We thank Drs. Richard W. Baumann, Brigham Young University, Stanley W. Szczytko,
University of Wisconsin, and Bill P. Stark,
Mississippi College, for verifying species
identifications and providing records. Drs.
Baumann and Stark also provided prepublication reviews of the manuscript. We also thank
Z. columbiana (Claassen)
Z. frigida (Claassen)
Z. haysi (Ricker)
Z. oregonensis (Claassen)
Doddsia occidentalis (Banks)
Sweltsa borealis (Banks)
S. revelstoka ( Jewett)
Suwallia pallidula (Banks)
Kathroperla perdita Banks
Paraperla frontalis (Banks)
Isoperla petersoni Needham & Christenson
I. sobria (Hagen)
Skwala americana (Klapalek)
Hesperoperla pacifica (Banks)
Claassenia sabulosa (Banks)
Dr. Edward A. Lisowski, Yakima, Washington,
and Robert Dopiriak, Mount Rainier National
Park, for making valuable collections of stoneflies from Mount Rainier available for study.
LITERATURE CITED
BAUMANN, R.W. 1979. Nearctic stonefly genera as indicators of ecological parameters (Plecoptera: Insecta).
Great Basin Naturalist 39:241–244.
BAUMANN, R.W., A.R. GAUFIN, AND R.F. SURDICK. 1977.
The stoneflies (Plecoptera) of the Rocky Mountains.
Memoirs of the American Entomological Society 31.
208 pp.
FRANKLIN, J.F., W.H. MOIR, M.A. HEMSTROM, S.E. GREENE,
AND B.G. SMITH. 1988. The forest communities of
Mount Rainier National Park. Scientific Monograph
Series 19. U.S. Department of Interior, Washington,
DC. 194 pp.
HOPPE, G.N. 1938. Plecoptera of Washington. University
of Washington Publication in Biology 4:139–174.
JEWETT, S.G. 1954. New stoneflies (Plecoptera) from western North America. Journal of the Fisheries Research
Board of Canada 11:543–549.
______. 1955. Notes and descriptions concerning western
North American stoneflies (Plecoptera). Wasmann
Journal of Biology 13:145–153.
______. 1959. The stoneflies (Plecoptera) of the Pacific
Northwest. Oregon State Monographs, Studies in
Entomology 3:1–95.
______. 1962. New stoneflies and records from the Pacific
coast of the United States. Pan-Pacific Entomologist
38:15–20.
KERST, C.D., AND N.H. ANDERSON. 1974. Emergence patterns of Plecoptera in a stream in Oregon, USA.
Freshwater Biology 4:205–212.
MULLINEAUX, D.R. 1974. Pumice and other pyroclastic
deposits in Mount Rainier National Park, Washington. U.S. Geological Survey Bulletin 1326. 83 pp.
NEEDHAM, J.G., AND P.W. CLAASSEN. 1925. A monograph
of the Plecoptera or stoneflies of America north of
Mexico. Thomas Say Foundation, Entomological
Society of America 2. 397 pp.
394
NELSON, C.R., AND R.W. BAUMANN. 1989. Systematics and
distribution of the winter stonefly genus Capnia (Plecoptera: Capniidae) in North America. Great Basin
Naturalist 49:289–363.
RICKER, W.E. 1943. Stoneflies of southwestern British
Columbia. Indiana University Publications in Science,
Series 12. 145 pp.
______. 1954. Nomenclatorial notes on Plecoptera. Proceedings of the Entomological Society of British
Columbia 51:37–39.
RICKER, W.E., AND G.G.E. SCUDDER. 1975. An annotated
checklist of the Plecoptera (Insecta) of British Columbia. Syesis 8:333–348.
STARK, B.P. 1983. A review of the genus Soliperla (Plecoptera: Peltoperlidae). Great Basin Naturalist 43:
30–44.
______. 1998. North American stonefly list. www.mc.edu/
~stark/stonefly.html
STARK, B.P., AND C.R. NELSON. 1994. Systematics, phylogeny and zoogeography of genus Yoraperla (Plecoptera: Peltoperlidae). Entomologica Scandinavica
25:241–273.
STARK, B.P., AND R.F. SURDICK. 1987. A new Kathroperla
species from western North America (Plecoptera:
Chloroperlidae). Proceedings of the Entomological
Society of Washington 89:527–531.
STEWART, K.W., AND J.A. STANGER. 1985. The nymphs, and
a new species, of North American Setvena Illies
(Plecoptera: Perlodidae). Pan-Pacific Entomologist
61:237–244.
STEWART, K.W., AND B.P. STARK. 1988. Nymphs of North
American stonefly genera (Plecoptera). Thomas Say
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12:1–460.
SZCZYTKO, S.W., AND K.W. STEWART. 1979. The genus
Isoperla (Plecoptera) of western North America:
holomorphology and systematics, and a new stonefly
genus Cascadoperla. Memoirs of the American
Entomological Society 32. 120 pp.
VANWIEREN, B.J., B.C. KONDRATIEFF, AND B.P. STARK. 2001.
A review of the North American species of Megarcys
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YOUNG, D.C., B.C. KONDRATIEFF, AND R.F. KIRCHNER. 1989.
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Received 1 May 2001
Accepted 15 September 2001
[Volume 62
APPENDIX. Localities for collections records of
Plecoptera of Mount Rainier National Park, Washington.
Order Plecoptera
Suborder Arctoperlaria
Group Euholognatha
Family Capniidae
Genus Capnia
Bolshecapnia sasquatchi
Ohanapecosh River, Hwy 143, 17-March-70, R.A. Hite &
D.S. Potter, 2 males (BYU); Ohanapecosh River, Hwy 143,
16-March-73, D.S. Potter & L.M. Preble, 1 male (BYU).
Capnia confusa
0.6 mile S of Ipsut Campground, 04-Apr-97, 1 female;
White River Highway 410, 29-May-97, 2 males, 1 female;
Tahoma Creek, 30-May-97, 1 male; Tahoma Creek at
Nisqually to Paradise Road, 30-May-97, 2 males; stream
by South Mowich River Camp, 04-May-98, 2 males, 1
female; Carbon River entrance, 15-Aug-99, 1 female; Fryingpan Creek at Sunrise Road Bridge, 16-Aug-99, 3
females; Tahoma Creek at Fish Creek, 08-Jul-00, 1 female;
Cataract Creek at Wonderland Trail, 09-Jul-00, 1 male, 9
females; St. Andrews Creek at Westside Road, 14-Jul-00,
1 male, 2 females; Fryingpan Creek at Sunrise Road, 16Jul-00, 1 female; Tahoma Creek, 13-May-01, 1 female;
seeps into White River on Highway 410, 14-May-01, 3
males, 4 females; Longmire, 15-May-01, 1 female;
Ohanapecosh Campground, 16-May-01, 1 female; White
River Highway 410, 0.5 mile S of park boundary, 16-May01, 3 males, 4 females.
Capnia elongata
0.7 mile SE of Ipsut Campground, 25-Mar-95, 1 male;
Paradise River in Paradise Valley, 02-Aug-00, 2 females;
Ohanapecosh River at the Grove of the Patriarchs, 16-May01, 1 male; White River Highway 410, 0.5 mile S of park
boundary, 16-May-01, 2 males.
Capnia excavata
Boggy stream tributary to Tahoma Creek, 07-May-01, 1
male, 1 female.
Capnia gracilaria
Paradise River at Fourth Crossing, 21-Apr-96, 1 male;
south of Chenuis Falls parking lot, 23-Feb-97, 1 male;
Westside Road at Paradise Road, 29-Mar-97, 1 male, 1
female; Kautz Creek at Nisqually to Paradise Road, 04Apr-01, 18 males, 6 females.
Capnia licina
Nisqually River at creek from Tato Falls, 14-Apr-94, 2 males,
1 female; 2.2 miles SE Ipsut Campground, 25-Mar-95, 2
males; Nahunta Falls, 16-Apr-95, 3 males; Glacier View
Bridge, 16-Apr-95, 2 males, 1 female; 0.1 mile NE Glacier
View Bridge, 14-Apr-96, 5 males; Snow Lake inflow, 15Jun-97, 7 males, 2 females; Shaw Creek at White River
Road, 07-Jul-99, 2 males; Paradise River in Paradise Valley, 10-Jul-00, 4 females; Paradise River in Paradise Valley,
02-Jul-01, 4 males, 3 females.
2002]
Capnia melia
Ipsut Falls, 23-Feb-95, 54 males, 19 females; 2.2 miles SE
of Ipsut Campground, 25-Mar-95, 1 male; 0.7 mile SE of
Ipsut Campground, 25-Mar-95, 1 male, 4 females; Nahunta
Falls, 16-Apr-95, 3 males; Nisqually River at creek from
Tato Falls, 18-Feb-96, 2 males; Pinnacle Peak Trail stream,
14-Apr-96, 5 males, 3 females; Nisqually River at creek
from Tato Falls, 14-Apr-96, 2 males; south of Chenuis Falls
parking lot, 23-Feb-97, 4 males, 1 female; 0.7 mile SE of
Ipsut Campground, 23-Feb-97, 4 males, 1 female; 0.6 mile
S of Ipsut Campground, 23-Feb-97, 6 males; Westside
Road 1.3 miles N of Paradise Road, 29-Mar-97, 2 males, 6
females; Tahoma Creek at Nisqually to Paradise Road, 29Mar-97, 1 male; 0.6 mile S of Ipsut Campground, 04-Apr97, 2 males, 2 females; 0.7 mile SE of Ipsut Campground,
04-Apr-97, 5 males, 9 females; Ipsut Falls on Ipsut Creek,
04-May-97, 9 males, 10 females; Nickel Creek at Stevens
Canyon Road, 28-May-97, 1 male; Crystal Creek at Highway 410, 29-May-97, 6 males, 5 females; unnamed stream
N of Crystal Creek, Highway 410, 29-May-97, 2 males, 11
females; seeps into White River on Highway 410, 29May-97, 2 males; Glacier View Bridge, 30-May-97, 4
females; Eagle Peak Creek at Nisqually River, 30-May-97,
3 males, 3 females; Eagle Peak Creek 4360 ft, 14-Jun-97,
3 males; Snow Lake inflow, 15-Jun-97, 1 male; Olallie
Creek, 28-Jun-97, 2 males; Fryingpan Creek at Sunrise
Road Bridge, 01-Apr-98, 1 male, 1 female; White River
Campground, 01-Apr-98, 1 male, 1 female; St. Andrews
Creek at Westside Road, 29-Apr-98, 2 males, 6 females;
Mountain Meadow Stream at Paul Peak Trail, 07-Jul-98, 1
male; Falls Creek at Stevens Canyon Road, 05-Jun-99, 4
males, 1 female; Falls Creek at Stevens Canyon Road, 11Jun-99, 3 males; Fryingpan Creek at Sunrise Road Bridge,
06-Jul-99, 4 males; South Puyallup River at Westside
Road, 24-Sep-00, 1 female; Eagle Peak Creek at Nisqually
River, 13-May-01, 5 males, 9 females; Ipsut Falls at Ipsut
Creek, 14-May-01, 1 male; Falls Creek at Stevens Canyon
Road, 07-Jun-01, 1 female; White River at Sunrise Road,
15-Jun-01, 1 male; seeps into Fryingpan Creek 0.5 mile
upstream from Sunrise Road, 15-Jun-01, 1 male, 2 females.
395
Canyon Road, 05-Jun-99, 3 females; Falls Creek at
Stevens Canyon Road, 11-Jun-99, 2 males, 1 female; tributary to Tahoma Creek at Westside Road, 08-Jul-99, 1
female; Paradise River in Paradise Valley, 16-Aug-99, 1
male, 1 female; Paradise River above Narada Falls, 16Aug-99, 1 female; Falls Creek at Stevens Canyon Road,
25-Jun-00, 4 females; Ohanapecosh River at the Grove of
the Patriarchs, 16-May-01, 2 females; White River Highway 410, 0.5 mile S of park boundary, 16-May-01, 4 males,
3 females; Falls Creek at Stevens Canyon Road, 07-Jun01, 1 male, 2 females; Panther Creek at Highway 123, 09Jun-01, 1 male, 1 female; Deer Creek at Eastside Trail,
09-Jun-01, 2 males, 1 female; Ohanapecosh River 0.5 mile
S of Shriner Peak Trailhead, 09-Jun-01, 1 male, 2 females;
Paradise River above Narada Falls, 02-Jul-01, 1 male, 1
female.
Genus Isocapnia
Isocapnia abbreviata
Seeps into White River on Highway 410, 14-May-01, 1
male, 3 females; White River Highway 410, 0.5 mile S of
park boundary, 16-May-01, 2 males, 6 females.
Isocapnia agassizi
White River on Highway 410, 29-May-97, 1 male; Tahoma
Creek at Nisqually to Paradise Road, 30-May-97, 1
female; seeps into White River on Highway 410, 14-May01, 2 males, 1 female; White River Highway 410, 0.5 mile
S of park boundary, 16-May-01, 1 male, 3 females.
Isocapnia grandis
White River on Highway 410, 28-May-97, 1 female; Kautz
Creek at Nisqually to Paradise Road, 04-Apr-01, 1 male;
males, 2 females; White River Highway 410, 0.5 mile S of
park boundary, 16-May-01, 2 males, 1 female.
Isocapnia spenceri
Capnia nana
Ohanapecosh Campground, 28-May-97, 1 female; seeps
into White River on Highway 410, 14-May-01, 1 male, 1
female.
Tahoma Creek at Nisqually to Paradise Road, 29-Mar-97,
1 female; Glacier View Bridge, 30-May-97, 3 females;
Longmire, 15-May-01, 1 female.
Mesocapnia oenone
Capnia sextuberculata
Deer Creek at Eastside Trail, 09-Jun-01, 1 female; Paradise River above Narada Falls, 02-Jul-01, 1 female.
Genus Eucapnopsis
Eucapnopsis brevicauda
Laughingwater Creek, 27-Jun-95, 1 female; Nisqually
River at Longmire, 28-May-97, 2 males; Ohanapecosh
Hot Springs, 28-May-97, 1 male; Ohanapecosh Campground, 28-May-97, 6 males, 5 females; Nickel Creek at
Stevens Canyon Road, 28-May-97, 19 males, 28 females;
Falls Creek at Stevens Canyon Road, 28-May-97, 11
males, 10 females; Ohanapecosh River at the Grove of the
Patriarchs, 28-May-97, 15 males, 4 females; Ohanapecosh
River at Sheep Creek, 29-May-97, 13 males, 7 females;
Panther Creek at Highway 123, 29-May-97, 22 males, 17
females; Olallie Creek, 28-Jun-97, 2 females; Huckleberry
Creek Bridge, 28-Apr-98, 1 male; Falls Creek at Stevens
Genus Mesocapnia
Carbon River Entrance, 20-Sep-97, 8 males, 3 females;
Carbon River Entrance, 15-Aug-99, 1 male; west fork of
White River, 13-Sep-99, 1 female.
Genus Paracapnia
Paracapnia oswegaptera
Seeps into Eagle Peak Creek, 28-May-97, 1 male.
Family Leuctridae
Genus Despaxia
Despaxia augusta
Westside Road, 0.2 mile NE of Paradise Road, 18-Dec-94,
1 male; Pinnacle Peak trail near road (Reflection Lakes),
28-Oct-95, 1 female; Paradise Valley Road 0.7 mile S of
Paradise, 28-Oct-95, 3 males, 6 females; 0.1 mile S of
Cayuse Pass, 24-Nov-95, 1 female; Fish Creek near
Tahoma Creek, 19-Dec-95, 1 female; 0.1 mile N of Fryingpan Creek Bridge, 03-Nov-96, 1 male, 3 females; 0.2 mile
396
NE of Fryingpan Creek bridge, 03-Nov-96, 1 female; 0.1
mile E of Ranger Creek Bridge, 17-Nov-96, 1 female; Kautz
Creek Trail 0.7 mile N of Nisqually Road, 08-Dec-96, 2
females; Ipsut Creek at Carbon River Road Bridge, 08Sep-97, 2 males; Carbon River Entrance, 20-Sep-97, 8
males, 3 females; small tributary to Shaw Creek, 07-Jul99, 6 males, 6 females; tributary to Tahoma Creek at Paradise Road, 08-Jul-99, 1 female; Carbon River Entrance,
15-Aug-99, 1 female; Carbon River at Alice Falls, 15-Aug99, 1 male; stream by Lake James, 13-Sep-99, 1 male, 1
female; west fork of White River, 13-Sep-99, 1 male; Van
Horn Creek at Waterfall, 13-Sep-99, 2 males; Fryingpan
Creek at Wonderland Trail Bridge, 29-Jun-00, 1 male, 1
female; seeps 0.9 mile S of St. Andrews Creek, 16-Sep-00,
6 males, 2 females; seeps 0.8 mile S of St. Andrews Creek,
16-Sep-00, 2 males, 2 females; stream 0.4 mile S of St.
Andrews Creek, 16-Sep-00, 15 males, 10 females; stream
1.75 miles S of St. Andrews Creek, 16-Sep-00, 9 males; St.
Andrews Creek at Westside Road, 16-Sep-00, 18 males, 2
females; stream 1.5 miles S of St. Andrews Creek, 16-Sep00, 2 males, 3 females; stream 1.15 miles S of St. Andrews
Creek, 16-Sep-00, 4 males, 1 female; Dick Creek at Wonderland Trail, 23-Sep-00, 1 female; St. Andrews Creek at
Westside Road, 17-Oct-00, 2 females; boggy stream tributary to Tahoma Creek, 07-May-01, 1 male.
Genus Moselia
Moselia infuscata
Carbon River Entrance, Rainforest Loop Trail, 08-May95, 2 females; Reflection Lake, stream near Pinnacle Peak
Trail, 14-Apr-96, 1 male; Tahoma Creek at Nisqually to
Paradise Road, 29-Mar-97, 1 male; Carbon River Entrance,
Rainforest Loop Trail, 27-May-97, 3 males, 2 females;
Carbon River Entrance, 27-May-97, 4 males, 5 females;
Ohanapecosh River at the Grove of the Patriarchs, 28May-97, 1 female; Ohanapecosh Campground, 28-May97, 1 female; Falls Creek at Stevens Canyon Road, 28May-97, 8 males, 5 females; seeps into Eagle Creek, 28May-97, 3 males, 6 females; Ohanapecosh River at Sheep
Creek, 29-May-97, 1 male, 1 female; unnamed stream N
of Crystal Creek, Highway 410, 29-May-97, 14 males, 11
females; unnamed stream 3.4 miles S of park boundary
Highway 410, 29-May-97, 9 males, 5 females; seeps into
White River on Highway 410, 29-May-97, 12 males, 7
females; Panther Creek at Highway 123, 29-May-97, 3
females; Nisqually River at Glacier View Bridge, 07-Jul99, 7 males, 6 females; Eagle Peak Creek at Nisqually
River, 30-May-97, 10 males, 6 females; Ipsut Creek at
Carbon River Road Bridge, 06-Jun-97, 2 males; Eagle
Peak Creek 4360 ft, 14-Jun-97, 5 males, 1 female; St.
Andrews Creek at Westside Road, 19-Jun-97, 6 males, 15
females; Olallie Creek, 28-Jun-97, 1 female; White River
Campground, 23-Jul-97, 4 males, 7 females; White River
at Sunrise Road Bridge, 02-Jul-98, 1 male, 2 females;
Mountain Meadow Stream at Paul Peak Trail, 07-Jul-98, 2
males; Falls Creek at Stevens Canyon Road, 05-Jun-99, 4
males; Falls Creek at Stevens Canyon Road, 11-Jun-99, 3
males, 4 females; Falls Creek at Stevens Canyon Road, 06Jul-99, 4 males, 8 females; Fryingpan Creek at Sunrise
Road Bridge, 06-Jul-99, 36 males, 38 females; tributary to
Ohanapecosh River, 07-Jul-99, 1 male, 4 females; stream
into White River at Highway 410 N of White River
Entrance, 07-Jul-99, 1 male; Nisqually River at Glacier
View Bridge, 30-May-97, 38 males, 11 females; tributary
to Tahoma Creek at Westside Road, 08-Jul-99, 7 males, 10
[Volume 62
females; Seep into Fryingpan Creek, 08-Jul-99, 3 males,
10 females; small tributary on Westside Road 2 miles N of
Paradise Road, 08-Jul-99, 1 female; tributary to Tahoma
Creek at Paradise Road, 08-Jul-99, 1 female; high-gradient tributary at Nisqually River at Longmire, 09-Jul-99, 1
male, 1 female; Paradise River in Paradise Valley, 16-Aug99, 1 female; Paradise River at Fourth Crossing, 16-Aug99, 1 female; creek behind Mystic Lake Ranger Station,
04-Sep-99, 1 male, 2 females; Falls Creek at Stevens
Canyon Road, 25-Jun-00, 5 males, 6 females; stream on
the Westside Road 1.5 miles from the Paradise Road, 26Jun-00, 1 male, 3 females; Paradise River in Paradise Valley, 10-Jul-00, 2 males, 4 females; St. Andrews Creek at
Westside Road, 14-Jul-00, 6 males, 3 females; Paradise
River at Fourth Crossing, 23-Jul-00, 1 male; Paradise River
in Paradise Valley, 02-Aug-00, 2 males; seeps into Eagle
Peak Creek, 07-May-01, 10 males, 2 females; boggy
stream tributary to Tahoma Creek 07-May-01, 17 males,
15 females; seeps by Nisqually River Bridge in Longmire,
13-May-01, 2 males, 2 females; Eagle Peak Creek at
Nisqually River, 13-May-01, 13 males, 3 females; Ipsut
Falls at Ipsut Creek, 14-May-01, 3 males, 4 females;
Huckleberry Creek Bridge, 14-May-01, 7 males; Eagle
Peak Creek at Nisqually River, 15-May-01, 1 male; small
stream near Ohanapecosh Visitor Center, 16-May-01, 1
male, 3 females; stream into White River at Highway 410
0.5 mile S of park boundary, 16-May-01, 2 males, 2
females; Panther Creek at Highway 123, 09-Jun-01, 2
females; Deer Creek at Eastside Trail, 09-Jun-01, 2 males,
1 female; St. Andrews Creek at Westside Road, 16-Jun-01,
2 males, 1 female; boggy stream tributary to Tahoma
Creek, 17-Jun-01, 7 males, 6 females; small stream near
Narada Falls, 23-Jul-01, 1 male.
Genus Paraleuctra
Paraleuctra forcipata
Westside Road 2.7 miles NNE of Nisqually Entrance, 29Mar-97, 1 male; Tahoma Creek, 1.8 miles NE of Nisqually
Entrance, 29-Mar-97, 1 male, 2 females; Carbon River at
Alice Falls, 06-Jun-97, 5 males; Huckleberry Creek
Bridge, 28-Apr-98, 2 males; Mountain Meadow Stream at
Paul Peak Trail, 07-May-98, 2 males; Falls Creek at
Stevens Canyon Road, 05-Jun-99, 1 male, 1 female; Falls
Creek at Stevens Canyon Road, 11-Jun-99, 2 males; Fryingpan Creek at Sunrise Road Bridge, 06-Jul-99, 2 males;
stream into White River at Highway 410 N of White
River Entrance, 07-Jul-99, 3 males; small tributary to
Shaw Creek, 07-Jul-99, 3 males; small tributary on Westside Road 2 miles N of Paradise Road, 08-Jul-99, 1 male;
Falls Creek at Stevens Canyon Road, 25-Jun-00, 2 males,
4 females; Tahoma Creek at Fish Creek, 08-Jul-00, 1 male;
St. Andrews Creek at Westside Road, 14-Jul-00, 2 males,
1 female; Fryingpan Creek at Sunrise Road Bridge, 16Jul-00, 1 male.
Paraleuctra occidentalis
White River Road 0.5 mile N of ranger station, 21-Apr-96,
1 male; Westside Road 1.3 miles N of Paradise Road, 29Mar-97, 3 males; Tahoma Creek 1.8 miles NE of Paradise
Road, 29-Mar-97, 3 males, 4 females; Ohanapecosh
Campground, 28-May-97, 1 male; Eagle Peak Creek at
Nisqually River, 30-May-97, 4 females; Huckleberry Creek
Bridge, 21-Apr-98, 1 male; Huckleberry Creek Bridge,
28-Apr-98, 2 males; St. Andrews Creek at Westside Road,
29-Apr-98, 1 male, 2 females; stream by South Mowich
2002]
397
River Camp, 07-May-98, 1 male; Fryingpan Creek at Sunrise Road Bridge, 06-Jul-99, 1 female; tributary to
Ohanapecosh River, 07-Jul-99, 4 males; tributary to
Tahoma Creek at Westside Road, 08-Jul-99, 1 male, 1
female; Falls Creek at Stevens Canyon Road, 08-Jul-99, 1
male; Paradise River in Paradise Valley, 16-Aug-99, 1
male, 1 female; Falls Creek at Stevens Canyon Road, 25Jun-00, 2 males, 4 females; Paradise River in Paradise Valley, 10-Jul-00, 3 females; Falls Creek at Stevens Canyon
Road, 16-Jul-00, 1 female; Fryingpan Creek 0.25 mile
upstream of Sunrise Road, 16-Jul-00 1 male; Ipsut Falls at
Ipsut Creek, 14-May-01, 1 male, 2 females; Ohanapecosh
River at the Grove of the Patriarchs, 16-May-01, 1 male, 1
female; White River Highway 410, 0.5 mile S of park
boundary, 16-May-01, 2 males, 2 females; Paradise River
in Paradise Valley, 02-Jul-01, 1 male.
Megaleuctra kincaidi
Paraleuctra projecta
Malenka californica
Boggy stream tributary to Tahoma Creek, 07-May-01, 3
males; seeps by Nisqually River Bridge in Longmire, 07May-01, 8 males; Eagle Peak Creek at Nisqually River,
13-May-01, 35 males, 18 females; seeps by Nisqually
River Bridge in Longmire, 13-May-01, 1 male; Ipsut Falls
at Ipsut Creek, 14-May-01, 1 male; Eagle Peak Creek at
Nisqually River, 15-May-01, 2 males, 7 females; Nisqually
River, 2 miles W of Longmire, 15-May-01, 1 male.
Carbon River Entrance, 27-May-97, 1 male; Carbon River
Entrance, 21-Sep-97, 1 male; Carbon River Entrance, 15Aug-99, 2 females; Carbon River Entrance, 17-Aug-99, 6
males.
Paraleuctra vershina
Rainforest Loop Trail at Carbon River Entrance, 08-May95, 7 males, 1 female; Carbon River Entrance, 24-May-95,
1 male; Carbon River Entrance, Rainforest Loop Trail, 27May-97, 5 males, 3 females; Nisqually River at Longmire,
28-May-97, 1 male, 2 females; Ohanapecosh Campground,
28-May-97, 1 male; Ohanapecosh River at the Grove of
the Patriarchs, 28-May-97, 1 male, 1 female; Ohanapecosh
River at Sheep Creek, 29-May-97, 1 male; seeps into White
River on Highway 410, 29-May-97, 4 males, 3 females;
Panther Creek at Highway 123, 29-May-97, 2 males; stream
into White River at Highway 410 N of White River
entrance, 07-Jul-99, 2 males; Fryingpan Creek at Sunrise
Road Bridge, 29-Jun-00, 1 female; Fish Creek at Westside
Road, 14-Jul-00, 1 female; Fish Creek at Dry Creek, 07May-01, 1 male, 1 female; seeps into White River on
Highway 410, 14-May-01, 3 males, 2 females; White River
Highway 410, 0.5 mile S of park boundary, 16-May-01, 1
male, 2 females; Ohanapecosh Campground, 16-May-01,
2 males, 1 female; White River at Sunrise Road, 15-Jun01, 7 males, 4 females; Paradise River in Paradise Valley,
02-Jul-01, 1 male; Fryingpan Creek at Sunrise Road Bridge,
02-Jul-01, 2 males, 3 females; seeps into Fryingpan Creek
0.5 mile upstream from Sunrise Road, 15-Jun-01, 6 males,
1 female.
female; Huckleberry Creek Bridge, 14-May-01, 2 males, 1
female.
Perlomyia utahensis
Eagle Peak Creek at Bridge, 30-May-97, 1 male.
Genus Megaleuctra
Ohanapecosh Campground, 09-May-96, 1 female; Fryingpan Creek at Sunrise Road Bridge, 06-Jul-99, 1 male;
seep into Fryingpan Creek, 08-Jul-99, 1 male, 2 females;
boggy stream tributary to Tahoma Creek, 07-May-01, 1
male.
Family Nemouridae
Genus Malenka
Malenka cornuta
Seeps into Eagle Creek, 28-May-97, 3 females; unnamed
stream 3.4 miles S of park boundary on Highway 410, 29May-97, 1 male; White River on Highway 410, 29-May-97,
1 male, 3 females; seeps into White River on Highway 410,
29-May-97, 1 female; Ipsut Creek at Campground, 08Sep-97, 1 female; Falls Creek at Stevens Canyon Road,
06-Jul-99, 1 male; Nisqually River at Glacier View Bridge,
07-Jul-99, 1 female; small tributary on Westside Road 2
miles N of Paradise Road, 08-Jul-99, 1 male; tributary to
Tahoma Creek at Paradise Road, 08-Jul-99, 10 males, 6
females; Carbon River Entrance, 15-Aug-99, 2 females;
small tributary on Westside Road 2 miles N of Paradise
Road, 17-Aug-99, 3 females; West fork of White River, 13Sep-99, 1 female; Falls Creek at Stevens Canyon Road,
25-Jun-00, 1 male, 1 female; stream on Westside Road 1.5
miles from Paradise Road, 26-Jun-00, 1 male; Fish Creek
at Westside Road, 14-Jul-00, 1 female; Falls Creek at
Stevens Canyon Road, 16-Jul-00, 1 male; seep 0.8 mile S
of St. Andrews Creek, 16-Sep-00, 1 female; White River
at Sunrise Road, 15-Jun-01, 1 male; boggy stream tributary
to Tahoma Creek, 17-Jun-01, 4 males, 7 females; boggy
stream tributary to Tahoma Creek, 09-Jul-01, 1 male.
Genus Ledina
Ledina tumana
Fryingpan Creek at Sunrise Road Bridge, 16-Jul-00, 1 male.
Genus Ostrocerca
Genus Perlomyia
Perlomyia collaris
Huckleberry Creek Bridge, 28-Apr-98, 1 male; stream by
South Mowich River Camp, 04-May-98, 1 male; North
Mowich River at Wonderland Trail, 07-May-98, 1 male;
Mountain Meadow Stream at Paul Peak Trail, 07-May-98,
1 male; seeps by Nisqually River Bridge in Longmire, 07May-01, 10 males, 1 female; seeps by Nisqually River
Bridge in Longmire, 13-May-01, 2 males, 1 female; Eagle
Peak Creek at Nisqually River, 13-May-01, 17 males, 4
females; seeps into White River on Highway 410, 14May-01, 1 male; Ipsut Falls at Ipsut Creek, 14-May-01, 1
Ostrocerca foersteri
Ipsut Creek at campground, 18-May-95, 3 males; Falls
Creek on Carbon River Road, 27-May-97, 1 male;
Nisqually River at Longmire, 28-May-97, 3 males; Falls
Creek at Stevens Canyon Road, 06-Jul-99, 1 male; small
stream near Longmire, 09-Jul-99, 1 male.
Genus Podmosta
Podmosta decepta
Small stream into Ohanapecosh River near Highway 123,
07-Jul-99, 1 female; Paradise River in Paradise Valley, 16-
398
Aug-99, 3 males, 5 females; Paradise River above Narada
Falls, 16-Aug-99, 1 female; Paradise River in Paradise Valley, 07-Sep-99, 2 females.
Podmosta delicatula
Paradise River above Narada Falls, 16-Aug-99, 1 male;
Paradise River in Paradise Valley, 16-Aug-99, 9 males, 7
females; Paradise River in Paradise Valley, 02-Aug-00, 1
male; Eagle Peak Creek at Nisqually River, 13-May-01, 1
female.
Genus Prostoia
[Volume 62
Bridge, 06-Jul-99, 8 males, 4 females; Paradise River in
Paradise Valley, 16-Aug-99, 13 males, 3 females; Paradise
River above Narada Falls, 16-Aug-99, 3 males, 4 females;
Paradise River in Paradise Valley, 02-Aug-00, 1 male;
boggy stream tributary to Tahoma Creek, 07-May-01, 3
males, 2 females; seeps into White River on Highway 410,
14-May-01, 2 males; stream into White River at Highway
410, 0.5 mile S of park boundary, 16-May-01, 5 males, 4
females; St. Andrews Creek at Westside Road, 16-Jun-01,
1 male; boggy stream tributary to Tahoma Creek, 17-Jun01, 5 males; Paradise River above Narada Falls, 02-Jul-01,
5 males, 4 females.
Prostoia besametsa
Genus Visoka
Ohanapecosh River at the Grove of the Patriarchs, 28May-97, 1 male, 1 female; Ohanapecosh Campground, 28May-97, 4 males, 5 females; Panther Creek at Highway
123, 29-May-97, 2 males, 5 females; Fryingpan Creek at
Sunrise Road Bridge, 06-Jul-99, 1 male; Fryingpan Creek
at Sunrise Road Bridge, 16-Jul-00, 1 male, 1 female; seeps
into White River on Highway 410, 14-May-01, 7 males, 6
females; Longmire, 15-May-01, 1 male, 1 female; Ohanapecosh Campground, 16-May-01, 2 males, 5 females;
White River on Highway 410, 0.5 mile S of park boundary, 16-May-01, 17 males, 8 females; Ohanapecosh River
at the Grove of the Patriarchs, 16-May-01, 2 males, 1
female; Deer Creek at Eastside Trail, 09-Jun-01, 1 female.
Genus Soyedina
Soyedina interrupta
Summerland, 23-Jul-95, 1 male; 0.5 mile W of White
River Ranger Station, 21-Apr-96, 1 male; seeps into Eagle
Creek, 28-May-97, 2 males, 1 female; unnamed stream 3.4
miles S of park boundary on Highway 410, 29-May-97, 1
male; seeps into White River on Highway 410, 29-May97, 4 males, 1 female; Nisqually River at Glacier View
Bridge, 30-May-97, 11 males, 3 females; St. Andrews
Creek at Westside Road, 19-Jun-97, 1 male, 2 females;
White River across from the White River Campground,
02-Jul-98, 1 male; Falls Creek at Stevens Canyon Road,
05-Jun-99, 1 male, 2 females; Falls Creek at Stevens
Canyon Road, 11-Jun-99, 1 male, 5 females; Falls Creek at
Stevens Canyon Road, 06-Jul-99, 4 males; Fryingpan
Creek at Sunrise Road Bridge, 06-Jul-99, 4 males, 5
females; Nisqually River at Glacier View Bridge, 07-Jul99, 3 males, 5 females; seep into Fryingpan Creek, 08-Jul99, 3 males, 1 female; stream on Westside Road 1.5 miles
from Paradise Road, 26-Jun-00, 1 male, 1 female; Fryingpan Creek at Wonderland Trail, 29-Jun-00, 1 male; South
Puyallup River at Westside Road, 14-Jul-00, 1 male, 1
female; Longmire, 13-May-01, 1 male; Eagle Peak Creek
at Nisqually River, 13-May-01, 1 male; Ipsut Falls at Ipsut
Creek, 14-May-01, 1 male, 1 female; Eagle Peak Creek at
Nisqually River, 15-May-01, 1 male; White River Highway 410, 0.5 mile S of park boundary, 16-May-01, 1 male.
Soyedina producta
Unnamed stream 3.4 miles S of park boundary on Highway 410, 29-May-97, 3 males; seeps into White River on
Highway 410, 29-May-97, 1 male; Nisqually River at
Glacier View Bridge, 30-May-97, 24 males, 3 females;
Carbon River at Alice Falls, 06-Jun-97, 3 males, 4 females;
Ipsut Creek at Carbon River Road Bridge, 06-Jun-97, 1
male, 1 female; North Mowich River at Wonderland Trail,
07-May-98, 2 males; Fryingpan Creek at Sunrise Road
Visoka cataractae
0.6 mile S of Ipsut Campground, 04-Apr-97, 1 male; Ipsut
Falls on Ipsut Creek, 04-May-97, 17 males, 1 female.
Genus Zapada
Zapada cinctipes
0.7 mile SE of Ipsut Campground, 25-Mar-95, 4 males, 2
females; Tahoma Creek at Westside Road, 02-Mar-96, 1
male, 3 females; Tahoma Creek 0.1 mile N of Dry Creek,
02-Mar-96, 1 male, 3 females; Reflection Lake, stream
near Pinnacle Peak Trail, 14-Apr-96, 2 males, 2 females;
White River Road 0.5 mile N of ranger station, 21-Apr-96,
1 male, 4 females; Ohanapecosh Campground, 12-May96, 1 female; Ipsut Falls at Ipsut Creek, 23-Feb-97, 1
female; south of Chenuis Falls parking lot, 23-Feb-97, 2
males, 13 females; 0.7 mile SE of Ipsut Campground, 23Mar-97, 1 female; Westside Road 1.3 miles N of Paradise
Road, 29-Mar-97, 4 females; Tahoma Creek, 1.8 miles NE
of Nisqually Entrance, 29-Mar-97, 4 males, 8 females;
Westside Road, 2.6 miles NNE of Nisqually Entrance, 29Mar-97, 5 males, 6 females; 0.6 mile S of Ipsut Campground, 04-Apr-97, 1 male, 1 female; Shaw Creek at
White River Road, 21-Apr-97, 1 male; White River Campground, 01-Apr-98, 1 female; tributary to Tahoma Creek at
Westside Road, 08-Jul-99, 1 female; Paradise River in Paradise Valley, 16-Aug-99, 9 males, 8 females; Paradise River
above Narada Falls, 07-Sep-99, 1 male, 2 females; Paradise River at Fourth Crossing, 16-Jul-00, 2 males, 1
female; Paradise River at Fourth Crossing, 23-Jul-00, 1
male; Paradise River in Paradise Valley, 02-Aug-00, 2
males; boggy stream tributary to Tahoma Creek, 07-May01, 3 females; Longmire, 15-May-01, 1 male.
Zapada columbiana
2.2 miles SE of Ipsut Campground, 25-Mar-95, 2 males, 2
females; Nahunta Falls, 16-Apr-95, 12 males; Nisqually
River at Longmire, 16-Apr-95, 1 male; 0.15 mile NE of
Glacier View Bridge, 16-Apr-95, 2 males; Squaw Lake, 07May-95, 1 male; Nisqually River at creek from Tato Falls,
14-Apr-96, 5 males, 5 females; Reflection Lake, stream
near Pinnacle Peak Trail, 14-Apr-96, 11 males, 5 females;
0.1 mile NE of Glacier View Bridge, 14-Apr-96, 1 male;
Paradise River at Fourth Crossing, 21-Apr-96, 22 males, 9
females; Edith Creek at Fourth Crossing, 05-May-96, 2
males, 1 female; Edith Creek at Fourth Crossing, 16-May96, 15 males, 6 females; Ipsut Falls, 23-Feb-97, 10 males,
3 females; Ipsut Creek, 0.6 mile S of Ipsut Campground,
23-Feb-97, 1 male; Tahoma Creek, 02-Mar-97, 1 male, 1
female; Westside Road, 2.6 miles NNE of Nisqually
Entrance, 29-Mar-97, 4 males, 3 females; Westside Road
2002]
1.3 miles N of Paradise Road, 29-Mar-97, 2 males; 0.6
mile S of Ipsut Campground, 04-Apr-97, 1 male; 0.7 mile
SE of Ipsut Campground, 04-Apr-97, 10 males, 9 females;
Ipsut Falls on Ipsut Creek, 04-May-97, 1 female; Crystal
Creek at Highway 410, 29-May-97, 1 male, 2 females; Eagle
Peak Creek at 4360 ft, 14-Jun-97, 2 males, 2 females;
White River Campground, 01-Apr-98, 15 males, 12 females;
Fryingpan Creek at Sunrise Road Bridge, 01-Apr-98, 5
males, 4 females; Paradise River in Paradise Valley, 10-Jul00, 3 males, 11 females; Paradise River in Paradise Valley,
16-Jul-00, 2 males, 3 females; Eagle Peak Creek at Nisqually River, 13-May-01, 1 male, 1 female; Deer Creek at
Eastside Trail, 09-Jun-01, 1 male; Paradise River in Paradise Valley, 02-Jul-01, 1 male.
Zapada cordillera
Westside Road 1.3 miles N of Paradise Road, 29-Mar-97, 1
male, 1 female; Tahoma Creek, 1.8 miles NE of Nisqually
Entrance, 29-Mar-97, 1 female; boggy stream tributary to
Tahoma Creek, 07-May-01, 1 female; seeps into Eagle
Peak Creek, 07-May-01, 1 male.
Zapada frigida
0.6 mile WSW of Shaw Creek Bridge, 03-Nov-96, 1 female;
Carbon River Entrance, Rainforest Loop Trail, 08-May95, 1 male, 1 female; Carbon River Entrance, Rainforest
Loop Trail, 27-May-97, 1 male; Ohanapecosh Campground,
28-May-97, 2 males; Falls Creek at Stevens Canyon Road,
28-May-97, 1 female; Carbon River at Alice Falls, 06-Jun97, 4 males, 2 females; Ipsut Creek at Carbon River Road
Bridge, 06-Jun-97, 1 male; St. Andrews Creek at Westside
Road, 19-Jun-97, 5 males; White River across from the
White River Campground, 02-Jul-98, 3 males, 1 female;
Fryingpan Creek at Sunrise Road Bridge, 06-Jul-99, 11
males, 5 females; Falls Creek at Stevens Canyon Road, 06Jul-99, 4 males; seep into Fryingpan Creek, 08-Jul-99, 3
males; tributary to Tahoma Creek at Westside Road, 08Jul-99, 2 males, 1 female; Falls Creek at Stevens Canyon
Road, 08-Jul-99, 2 males; high-gradient tributary at
Nisqually River Longmire, 09-Jul-99, 2 males; Paradise
River in Paradise Valley, 16-Aug-99, 5 males, 4 females;
Falls Creek at Stevens Canyon Road, 25-Jun-00, 1 male, 2
females; Paradise River in Paradise Valley, 10-Jul-00, 2
females; St. Andrews Creek at Westside Road, 14-Jul-00,
1 male; Paradise River in Paradise Valley, 02-Aug-00, 1 male,
1 female; seeps into Fryingpan Creek 0.5 mile upstream
from Sunrise Road, 15-Jun-01, 4 males, 1 female.
Zapada haysi
02-Jul-98, 1 male; White River Highway 410, 0.5 mile S
of park boundary, 16-May-01, 2 males; Ohanapecosh
River at the Grove of the Patriarchs, 16-May-01, 1 male;
boggy stream tributary to Tahoma Creek, 04-Jul-01, 1
male.
Zapada oregonensis
Carbon River Entrance, Rainforest Loop Trail, 27-May97, 4 males, 1 female; unnamed stream N of Crystal Creek,
Highway 410, 29-May-97, 3 males; seeps into White River
on Highway 410, 29-May-97, 1 male; White River Highway 410, 29-May-97, 1 male; Mountain Meadow Stream
at Paul Peak Trail, 07-May-98, 2 males; White River across
from the White River Campground, 02-Jul-98, 2 males, 2
399
females; tributary to Tahoma Creek at Westside Road, 08Jul-99, 3 males, 1 female; Paradise River in Paradise Valley, 07-Sep-99, 1 male; Ohanapecosh River at the Grove
of the Patriarchs, 16-May-01, 1 male; White River Highway 410, 0.5 mile S of park boundary, 16-May-01, 2 males;
boggy stream tributary to Tahoma Creek, 17-Jun-01, 4
males, 1 female.
Family Taeniopterygidae
Genus Doddsia
Doddsia occidentalis
Nisqually River at Longmire, 16-Apr-95, 2 males; Paradise
Ice Caves, 20-Aug-95, 2 females; Westside Road 2.6 miles
NNE of Nisqually Entrance, 29-Mar-97, 6 males, 5 females;
Nisqually River at Longmire, 28-May-97, 5 females; seeps
into White River on Highway 410, 29-May-97, 7 females;
Carbon River at Alice Falls, 06-Jun-97, 2 males; Glacier
Basin, 20-Jul-97, 3 females; Glacier Basin, 21-Jul-97, 3
females; Glacier Basin, 27-Jul-97, 16 females; Huckleberry Creek Bridge, 17-Apr-98, 2 males, 2 females; Longmire, 20-Apr-98, 1 male; Huckleberry Creek Bridge, 21Apr-98, 1 male, 5 females; South Puyallup River at Westside
Road, 21-Apr-98, 1 female; St. Andrews Creek at Westside
Road, 29-Apr-98, 3 males, 1 female; North Mowich River
at Wonderland Trail, 07-May-98, 1 male, 1 female; Glacier
Basin, 06-Jun-98, 24 females; Falls Creek at Stevens Canyon Road, 11-Jun-99, 2 females; Falls Creek at Stevens
Canyon Road, 06-Jul-99, 2 females; small stream into
Ohanapecosh River near Highway 123, 07-Jul-99, 1 female;
River Entrance, 07-Jul-99, 1 female; Pebble Creek at
7200 feet, 20-Jul-99, 6 females; Muir Snowfield at 8000 ft,
23-Jul-99, 1 female; Fryingpan Creek at Wonderland Trail,
29-Jun-00, 1 male, 1 female; Paradise River in Paradise
Valley, 10-Jul-00, 3 females; Fryingpan Creek at Sunrise
Road Bridge, 10-Jul-00, 2 females; Fryingpan Creek at
Sunrise Road Bridge, 16-Jul-00, 19 females; Kautz Creek
at Nisqually to Paradise Road, 04-Apr-01, 1 male, 1 female;
Paradise Ice Caves, 12-May-01, 2 females; Kautz Creek at
Nisqually to Paradise Road, 13-May-01, 1 female; Tahoma
Creek at Nisqually to Paradise Road, 13-May-01, 1 male,
1 female; Ipsut Falls at Ipsut Creek, 14-May-01, 3 females;
males, 7 females; Eagle Peak Creek at Nisqually River, 15May-01, 1 male; Nisqually River, 2 miles W of Longmire,
15-May-01, 8 males, 6 females; Longmire, 15-May-01, 13
males, 4 females; Paradise River at Stevens Canyon Road,
15-May-01, 1 male, 1 female; Ohanapecosh River at the
Grove of the Patriarchs, 15-May-01, 1 male, 1 female;
White River Highway 410, 0.5 mile S of park boundary,
16-May-01, 1 male, 2 females; Ohanapecosh Campground,
16-May-01, 4 females.
Genus Taenionema
Taenionema kincaidi
Kautz Creek Trail 0.7 mile N of Nisqually Road, 08-May95, 1 male, 2 females; South Mowich River Camp, 31-May95, 2 males, 1 female; Tahoma Woods, 29-Apr-96, 1 male;
Ohanapecosh Campground, 28-May-97, 6 males, 5 females;
Ohanapecosh River at the Grove of the Patriarchs, 28May-97, 3 males, 3 females; Nisqually River at Longmire,
28-May-97, 4 males, 1 female; Panther Creek at Highway
123, 29-May-97, 5 males, 2 females; seeps into White River
on Highway 410, 29-May-97, 17 males, 7 females; Deer
400
[Volume 62
Creek on Highway 123, 29-May-97, 5 males, 3 females;
Ohanapecosh River at Sheep Creek, 29-May-97, 5 males,
4 females; Tahoma Creek at Nisqually to Paradise Road,
30-May-97, 6 females; Carbon River at Alice Falls, 06Jun-97, 6 males, 6 females; Ipsut Creek at Carbon River
Road Bridge, 06-Jun-97, 5 males, 4 females; Mountain
Meadow Stream at Paul Peak Trail, 07-May-98, 7 males, 8
females; North Mowich River at Wonderland Trail, 07May-98, 14 males, 4 females; Fryingpan Creek at Sunrise
Road Bridge, 03-Jul-98, 1 female; Fryingpan Creek at Sunrise Road Bridge, 06-Jul-99, 18 males, 7 females; Nisqually
River at Glacier View Bridge, 07-Jul-99, 1 male, 2 females;
River Entrance, 07-Jul-99, 1 female; Shaw Creek at White
River Road, 07-Jul-99, 2 males; tributary to Tahoma Creek
at Westside Road, 08-Jul-99, 2 females; Falls Creek at
Stevens Canyon Road, 08-Jul-99, 1 male; small stream
near Longmire, 09-Jul-99, 1 female; Paradise River at
Fourth Crossing, 16-Aug-99, 1 male; Paradise River above
Narada Falls, 16-Aug-99, 2 males, 27 females; Fryingpan
Creek at Sunrise Road Bridge, 16-Aug-99, 5 females; Falls
Creek at Stevens Canyon Road, 25-Jun-00, 1 male; Fryingpan Creek at Wonderland Trail, 29-Jun-00, 2 females;
Cataract Creek at Wonderland Trail, 09-Jul-00, 2 males, 3
females; Paradise River in Paradise Valley, 10-Jul-00, 1
male, 2 females; Fryingpan Creek at Sunrise Road, 10-Jul00, 1 female; Fish Creek at Westside Road, 14-Jul-00, 5
females; Fryingpan Creek at Sunrise Road, 16-Jul-00, 4
females; Fryingpan Creek 0.25 mile upstream from Sunrise Road, 16-Jul-00, 1 male, 2 females; Falls Creek at
Stevens Canyon Road, 16-Jul-00, 1 female; Paradise River
at Fourth Crossing, 16-Jul-00, 4 males, 1 female; Fish Creek
at Westside Road, 05-Aug-00, 1 female; seeps by Nisqually
River Bridge in Longmire, 07-May-01, 2 males, 1 female;
Fish Creek at Dry Creek, 07-May-01, 8 males; Kautz
Creek at Nisqually to Paradise Road, 13-May-01, 8 males;
Eagle Peak Creek at Nisqually River, 13-May-01, 1 male;
Tahoma Creek at Nisqually to Paradise Road, 13-May-01,
6 males; Ipsut Falls at Ipsut Creek, 14-May-01, 2 females;
males; Huckleberry Creek Bridge, 14-May-01, 1 male;
Paradise River at Stevens Canyon Road, 15-May-01, 1
male; Nisqually River, 2 miles W of Longmire, 15-May01, 4 males, 1 female; White River Highway 410, 0.5 mile
S of park boundary, 16-May-01, 4 males; Ohanapecosh
Campground, 16-May-01, 2 males; Panther Creek at Highway 123, 09-Jun-01, 1 female; Ohanapecosh River 0.5
mile S of Shriner Peak Trailhead, 09-Jun-01, 4 males, 3
females; Deer Creek at Eastside Trail, 09-Jun-01, 1 male;
White River at Sunrise Road, 15-Jun-01, 1 male, 1 female;
Paradise River above Narada Falls, 02-Jul-01, 2 females.
Alloperla serrata
Taenionema pallidum
Suwallia forcipata
North Mowich River at Wonderland Trail, 07-May-98, 2
males, 1 female; Tahoma Creek at Nisqually to Paradise
Road, 13-May-01, 1 male; seeps into White River on
Highway 410, 14-May-01, 1 male.
Nisqually River at Longmire, 04-Apr-96, 1 male, 8 females;
Carbon River Entrance, 20-Sep-97, 10 males, 13 females;
Huckleberry Creek Bridge, 27-Sep-97, 8 females.
Group Systellognatha
Family Chloroperlidae
Genus Alloperla
South Mowich River Camp, 31-May-95, 4 males, 5
females.
Edith Creek at Fourth Crossing, 13-Sep-97, 1 male; Paradise River in Paradise Valley, 16-Aug-99, 3 males; Paradise
River above Narada Falls, 16-Aug-99, 9 males; Paradise
River at Fourth Crossing, 16-Aug-99, 2 males; Paradise
River in Paradise Valley, 07-Sep-99, 1 male, 2 females;
females.
Genus Plumiperla
Plumiperla diversa
Nisqually River at Longmire, 28-May-97, 1 male; seeps
female; Tahoma Creek at Nisqually to Paradise Road, 30May-97, 3 males, 2 females; Longmire, 27-Jul-98, 1 male;
Falls Creek at Stevens Canyon Road, 06-Jul-99, 5 males, 5
females; small stream near Longmire, 07-Jul-99, 2 males;
Nisqually River at Glacier View Bridge, 07-Jul-99, 14
males, 11 females; tributary to Ohanapecosh River, 07Jul-99, 1 male, 6 females; Tahoma Creek, 08-Jul-99, 1
male, 3 females; tributary to Tahoma Creek at Westside
Road, 08-Jul-99, 1 female; Paradise River above Narada
Falls, 16-Aug-99, 2 males; Fryingpan Creek at Sunrise
Road Bridge, 16-Aug-99, 2 males, 2 females; Falls Creek
at Stevens Canyon Road, 25-Jun-00, 2 females; Fryingpan
Creek at Sunrise Road Bridge, 29-Jun-00, 1 male, 2
females; Tahoma Creek at Fish Creek, 08-Jul-00, 5 males,
2 females; Cataract Creek at Wonderland Trail, 09-Jul-00,
1 male, 1 female; South Puyallup River at Westside Road,
14-Jul-00, 20 males, 7 females; Fryingpan Creek at Sunrise Road Bridge, 16-Jul-00, 2 males, 5 females; Fryingpan
Creek 0.25 mile upstream of Sunrise Road, 16-Jul-00, 3
males, 4 females; Tahoma Creek at Nisqually to Paradise
Road, 13-May-01, 1 male; White River at Sunrise Road
Bridge, 15-Jun-01, 3 males, 2 females; boggy stream tributary to Tahoma Creek, 17-Jun-01, 2 males, 1 female;
small stream near Narada Falls, 23-Jul-01, 3 males.
Genus Suwallia
Suwallia dubia
Ipsut Creek at Carbon River Road Bridge, 08-Sep-97, 1
female; Carbon River Entrance, 15-Aug-99, 1 male, 1
female; Carbon River Entrance, 16-Aug-99, 1 male; Falls
Creek at Stevens Canyon Road, 16-Aug-99, 1 male; small
tributary on Westside Road 2 miles N of Paradise Road,
17-Aug-99, 1 male; Van Horn Creek at Waterfall, 13-Sep99, 3 males, 2 females; West fork of White River, 13-Sep99, 1 female; South Puyallup River at Westside Road, 29Sep-00, 16 males, 6 females.
Suwallia pallidula
Genus Sweltsa
Alloperla fraterna
Falls Creek at Stevens Canyon Road, 16-Aug-99, 6 males;
small stream near Ohanapecosh Visitor Center, 16-May01, 1 male.
Sweltsa borealis
Carbon River Entrance, Rainforest Loop Trail, 08-May95, 3 males, 3 females; Wonderland Trail 0.5 mile E of Paul
2002]
401
Peak Trail junction, 01-Jun-95, 1 male, 1 female; Carbon
River Entrance, 27-May-97, 1 male, 2 females; Carbon
River Entrance, Rainforest Loop Trail, 27-May-97, 2 males,
2 females; Nisqually River at Longmire, 28-May-97, 1
female; Falls Creek at Stevens Canyon Road, 28-May-97,
5 males, 3 females; White River Highway 410, 29-May-97,
1 male, 1 female; unnamed stream N of Crystal Creek,
Highway 410, 29-May-97, 10 males, 4 females; seeps into
White River on Highway 410, 29-May-97, 3 females; Panther Creek at Highway 123, 29-May-97, 1 female; Eagle
Peak Creek at Nisqually River, 30-May-97, 1 female; Ipsut
Creek at Carbon River Road Bridge, 06-Jun-97, 1 male, 1
female; St. Andrews Creek at Westside Road, 14-Jun-97, 1
male; Olallie Creek, 28-Jun-97, 2 males; Falls Creek at
Stevens Canyon Road, 06-Jul-99, 4 males, 2 females; stream
into White River at Highway 410 N of White River entrance, 07-Jul-99, 1 female; tributary to Ohanapecosh
River, 07-Jul-99, 7 females; Nisqually River at Glacier
View Bridge, 07-Jul-99, 3 females; small stream into
Ohanapecosh River near Highway 123, 07-Jul-99, 1 male;
tributary to Tahoma Creek at Westside Road, 08-Jul-99, 3
males, 4 females; tributary to Tahoma Creek at Paradise
Road, 08-Jul-99, 3 females; small tributary on Westside
Road 2 miles N of Paradise Road, 08-Jul-99, 1 female;
high-gradient tributary at Nisqually River Longmire, 09Jul-99, 1 male; Paradise River above Narada Falls, 16-Aug99, 1 male, 3 females; Paradise River in Paradise Valley,
16-Aug-99, 5 males, 7 females; Paradise River in Paradise
Valley, 07-Sep-99, 1 male, 1 female; Van Horn Creek at
Waterfall, 13-Sep-99, 1 female; Falls Creek at Stevens
Canyon Road, 25-Jun-00, 4 males, 2 females; Cataract
Creek at Wonderland Trail, 09-Jul-00, 1 female; Paradise
River in Paradise Valley, 10-Jul-00, 3 males; Fish Creek at
Westside Road, 14-Jul-00, 1 male; South Puyallup River
at Westside Road, 14-Jul-00, 1 female; Paradise River in
Paradise Valley, 23-Jul-00, 1 female; Paradise River in Paradise Valley, 02-Aug-00, 1 male, 4 females; Olallie Creek
at Olallie Creek Camp, 03-Aug-00, 1 female; boggy stream
tributary to Tahoma Creek, 07-May-01, 2 males; Ipsut
Falls at Ipsut Creek, 14-May-01, 1 male; Ohanapecosh
boggy stream tributary to Tahoma Creek, 17-Jun-01, 3
males, 4 females; small stream near Narada Falls, 23-Jul01, 2 males, 2 females.
River in Paradise Valley, 16-Aug-99, 8 males, 4 females;
Paradise River in Paradise Valley, 07-Sep-99, 2 males, 2
females; Falls Creek at Stevens Canyon Road, 25-Jun-00,
2 males; stream on Westside Road 1.5 miles from Paradise
Road, 26-Jun-00, 2 males, 4 females; Fryingpan Creek at
Sunrise Road Bridge, 29-Jun-00, 1 male; Cataract Creek
at Wonderland Trail, 09-Jul-00, 10 males, 3 females; Fryingpan Creek at Sunrise Road Bridge, 10-Jul-00, 2 males,
2 females; Fish Creek at Westside Road, 14-Jul-00, 2
males, 11 females; St. Andrews Creek at Westside Road,
14-Jul-00, 10 males, 5 females; Fryingpan Creek at Sunrise Road Bridge, 16-Jul-00, 1 male, 2 females; Fryingpan
Creek 0.25 mile upstream from Sunrise Road, 16-Jul-00, 4
males; Paradise River at Fourth Crossing, 23-Jul-00, 3
males; Paradise River in Paradise Valley, 02-Aug-00, 1
male, 6 females; Fish Creek at Westside Road, 05-Aug-00,
1 male, 2 females; boggy stream tributary to Tahoma Creek,
01-Jun-01, 8 males, 2 females; Falls Creek at Stevens Canyon Road, 07-Jun-01, 1 male, 1 female; Panther Creek at
Highway 123, 09-Jun-01, 1 male; boggy stream tributary
to Tahoma Creek, 17-Jun-01, 6 males; Paradise River
above Narada Falls, 02-Jul-01, 1 male; boggy stream tributary to Tahoma Creek, 09-Jul-01, 2 males, 1 female; small
stream near Narada Falls, 23-Jul-01, 2 males.
Sweltsa exquisita
Sweltsa revelstoka
Carbon River Entrance, Rainforest Loop Trail, 27-May97, 1 male; Carbon River Entrance, 27-May-97, 1 male, 1
female; Carbon River at Chenuis Falls Trailhead, 06-Jun97, 8 males; St. Andrews Creek at Westside Road, 24-Jul97, 3 males, 1 female; Edith Creek at Fourth Crossing, 13Sep-97, 2 males; Falls Creek at Stevens Canyon Road, 06Jul-99, 7 males, 13 females; Fryingpan Creek at Sunrise
Road Bridge, 06-Jul-99, 1 male; tributary to Ohanapecosh
River, 07-Jul-99, 2 males; Nisqually River at Glacier View
Bridge, 07-Jul-99, 2 males; small stream into Ohanapecosh River near Highway 123, 07-Jul-99, 9 males; tributary to Tahoma Creek at Westside Road, 08-Jul-99, 2
males, 3 females; Falls Creek at Stevens Canyon Road, 08Jul-99, 4 males; small stream near Longmire, 09-Jul-99, 2
males, 3 females; High gradient tributary at Nisqually
River at Longmire, 09-Jul-99, 1 male; Paradise River at
Fourth Crossing, 16-Aug-99, 1 male; Paradise River above
Narada Falls, 16-Aug-99, 7 males, 4 females; Falls Creek at
Stevens Canyon Road, 16-Aug-99, 1 female; Paradise
White River Campground, 23-Jul-97, 7 males, 6 females;
inflow upper Palisade Lake, 08-Aug-97, 17 males, 8 females;
White River across from White River Campground, 02Jul-98, 4 males, 1 female; seep into Fryingpan Creek, 08Jul-99, 1 male; Fryingpan Creek at Sunrise Road Bridge,
16-Aug-99, 1 female; Paradise River above Narada Falls,
16-Aug-99, 3 males, 5 females; Paradise River in Paradise
Valley, 16-Aug-99, 5 males, 3 females; Paradise River at
Fourth Crossing, 16-Aug-99, 5 males, 7 females; creek behind Mystic Lake Ranger Station, 04-Sep-99, 1 female;
Cataract Creek at Wonderland Trail, 09-Jul-00, 1 female;
Fryingpan Creek at Sunrise Road, 10-Jul-00, 6 males, 2
females; Fryingpan Creek at Sunrise Road, 16-Jul-00, 1
female; Paradise River at Fourth Crossing, 23-Jul-00, 7
males, 1 female; Paradise River in Paradise Valley, 02-Aug00, 4 males, 4 females; Tamanos Creek at Trail, 03-Aug-00,
2 males; Paradise River in Paradise Valley, 02-Jul-01, 1 male;
Fryingpan Creek at Sunrise Road, 02-Jul-01, 1 female;
Dick Creek at Wonderland Trail, 27-Jul-01, 1 female.
Sweltsa occidens
Laughingwater Creek, 27-Jun-95, 5 males; St. Andrews
Creek at Westside Road, 24-Jul-97, 4 males, 32 females;
high-gradient tributary at Nisqually River Longmire, 09Jul-99, 3 males; Paradise River in Paradise Valley, 16-Aug99, 4 males, 7 females; Paradise River above Narada Falls,
16-Aug-99, 52 males, 15 females; Paradise River at Fourth
Crossing, 16-Aug-99, 1 male; Falls Creek at Stevens Canyon
Road, 25-Jun-00, 1 male; Cataract Creek at Wonderland
Trail, 09-Jul-00, 2 males; St. Andrews Creek at Westside
Road, 14-Jul-00, 4 females; Fryingpan Creek at Sunrise
Road Bridge, 16-Jul-00, 1 male, 2 females; Fryingpan Creek
0.25 mile upstream Sunrise Road, 29-Jun-00, 1 male; Paradise River at Fourth Crossing, 23-Jul-00, 2 males, 1 female;
Tamanos Creek at trail, 03-Aug-00, 10 males, 5 females;
Panther Creek at Highway 123, 09-Jun-01, 1 male; boggy
stream tributary to Tahoma Creek, 17-Jun-01, 1 male; Fryingpan Creek at Sunrise Road Bridge, 02-Jul-01, 1 male;
small stream near Narada Falls, 23-Jul-01, 1 male.
402
Sweltsa new species
Tributary to Tahoma Creek at Westside Road, 08-Jul-99, 1
male.
Genus Kathroperla
Kathroperla perdita
White River Highway 410, 29-May-97, 2 males; unnamed
stream 3.4 miles S of park boundary on Highway 410, 29May-97, 1 female; Crystal Creek at Highway 410, 29May-97, 1 male, 1 female; unnamed stream N of Crystal
Creek, Highway 410, 29-May-97, 2 females; Panther
Creek at Highway 123, 29-May-97, 1 female; Tahoma
Creek at Nisqually to Paradise Road, 30-May-97, 2 males;
Mountain Meadow Stream at Paul Peak Trail, 07-May-98,
1 male; seeps into White River on Highway 410, 14-May01, 1 female; Ipsut Falls at Ipsut Creek, 14-May-01, 1
male, 1 female; White River Highway 410, 0.5 mile S of
park boundary, 16-May-01, 1 male, 2 females; Deer Creek
at Eastside Trail, 09-Jun-01, 1 male; White River at Sunrise Road, 15-Jun-01, 1 female.
Kathroperla takhoma
Falls Creek at Stevens Canyon Road, 06-Jul-99, 1 female;
tributary to Ohanapecosh River, 07-Jul-99, 1 female; Falls
Creek at Stevens Canyon Road, 25-Jun-00, 1 female;
small stream near Ohanapecosh Visitor Center, 16-May01, 1 male.
Genus Paraperla
Paraperla frontalis
Ohanapecosh Ranger Station, 22-Jun-96, 1 female; Carbon River Entrance, 27-May-97, 2 females; Ohanapecosh
White River Highway 410, 29-May-97, 1 male.
Paraperla wilsoni
0.15 mile NE of Glacier View Bridge, 16-Apr-95, 1 female;
0.6 mile S of Ipsut Campground, 04-Apr-97, 1 female;
Nisqually River at Longmire, 28-May-97, 3 females, 1
nymph; Panther Creek at Highway 123, 09-Jun-01, 1 male.
Family Peltoperlidae
Genus Soliperla
Soliperla fenderi
Tributary to Tahoma Creek, 08-Jul-99, 4 males, 1 female;
Road, 08-Jul-99, 1 female; Falls Creek at Stevens Canyon
Road, 08-Jul-99, 1 male, 1 female; small tributary on
Westside Road 2 miles N of Paradise Road, 17-Aug-99, 4
males, 1 female; boggy stream tributary to Tahoma Creek,
17-Aug-99, 1 female; stream on Westside Road 1.5 miles
from Paradise Road, 26-Jun-00, 1 male; Cataract Creek at
Wonderland Trail, 09-Jul-00, 1 male; Dick Creek at Wonderland Trail, 27-Jul-01, 4 males.
Genus Yoraperla
[Volume 62
02-Jul-98, 1 female; boggy stream tributary to Tahoma
Creek, 07-May-01, 1 male, 2 females; White River Highway 410, 0.5 mile S of park boundary, 16-May-01, 1
female; stream into White River at Highway 410, 0.5 mile
S of park boundary, 16-May-01, 1 male; Ohanapecosh
River 0.5 mile S of Shriner Peak Trailhead, 09-Jun-01, 5
females.
Yoraperla nigrisoma
Mowich Road at Grindstone Trail Crossing, 07-Jul-95, 1
female; Ipsut Creek at Carbon River Road Bridge, 06Jun-97, 1 female; Falls Creek at Stevens Canyon Road, 11Jun-99, 2 males; Fryingpan Creek at Sunrise Road Bridge,
06-Jul-99, 1 male, 1 female; small tributary on Westside
Road 2 miles N of Paradise Road, 08-Jul-99, 1 female;
tributary to Tahoma Creek at Westside Road, 08-Jul-99,
25 males, 12 females; tributary to Tahoma Creek at Paradise Road, 08-Jul-99, 2 males, 1 female; Paradise River in
Paradise Valley, 16-Aug-99, 1 male, 1 female; Paradise
River above Narada Falls, 16-Aug-99, 2 females; Falls
Creek at Stevens Canyon Road, 16-Aug-99, 2 females;
Road, 17-Aug-99, 1 male; creek behind Mystic Lake
Ranger Station, 04-Sep-99, 2 males, 3 females; Falls Creek
at Stevens Canyon Road, 25-Jun-00, 9 males, 4 females;
Paradise River in Paradise Valley, 10-Jul-00, 6 males; St.
Andrews Creek at Westside Road, 14-Jul-00, 3 males, 2
females; Falls Creek at Stevens Canyon Road, 16-Jul-00, 1
male, 3 females; Paradise River in Paradise Valley, 02-Aug00, 5 females; Olallie Creek at Olallie Creek Camp, 03Aug-00, 1 female; boggy stream tributary to Tahoma
Creek, 07-May-01, 1 male, 2 females; small stream near
Ohanapecosh Visitor Center, 16-May-01, 1 male; Ohanapecosh River 0.5 mile S of Shriner Peak Trailhead, 09Jun-01, 1 female; boggy stream tributary to Tahoma Creek,
17-Jun-01, 8 males, 2 females; boggy stream tributary to
Tahoma Creek, 04-Jul-01, 1 male; boggy stream tributary
to Tahoma Creek, 09-Jul-01, 2 females.
Yoraperla siletz
Falls Creek at Stevens Canyon Road, 06-Jul-99, 9 males,
11 females; Falls Creek at Stevens Canyon Road, 08-Jul99, 5 males, 5 females; small tributary on Westside Road 2
miles N of Paradise Road, 08-Jul-99, 1 male, 1 female;
Falls Creek at Stevens Canyon Road, 25-Jun-00, 2 males,
2 females.
Family Perlidae
Genus Doroneuria
Doroneuria baumanni
Carbon River Entrance at June Creek, 13-Jul-95, 2 nymphs;
tributary to Tahoma Creek at Paradise Road, 08-Jul-99, 1
female; Carbon River Entrance, 15-Aug-99, 5 males, 1
female.
Genus Hesperoperla
Yoraperla mariana
Hesperoperla pacifica
Ohanapecosh Campground, 28-May-97, 1 male, 4 females;
Panther Creek at Highway 123, 29-May-97, 2 males, 1
female; Ohanapecosh River at Sheep Creek, 29-May-97, 1
male, 1 female; unnamed stream N of Crystal Creek,
Highway 410, 29-May-97, 9 males, 7 females; Mountain
Meadow Stream at Paul Peak Trail, 07-May-98, 1 male;
Ohanapecosh Ranger Station, 17-Jul-97, 1 female.
Genus Claassenia
Claassenia sabulosa
Ohanapecosh Ranger Station, 09-Sep-97, 1 female.
2002]
Family Perlodidae
Genus Isoperla
403
Seeps into White River on Highway 410, 29-May-97, 1
nymph.
Bridge, 06-Jul-99, 1 male, 2 females; Fryingpan Creek at
Sunrise Road Bridge, 16-Aug-99, 1 female; Cataract Creek
at Wonderland Trail, 09-Jul-00, 1 female; Fryingpan Creek
at Sunrise Road Bridge, 10-Jul-00, 2 females; Nisqually
River at Longmire, 13-May-01, 1 male, 1 female; seeps
females; Paradise River above Narada Falls, 02-Jul-01, 1 female; Cataract Creek at Wonderland Trail, 20-Jul-01, 1 female; Puyallup River at Wonderland Trail, 28-Jul-01, 1 male.
Isoperla gravitans
Megarcys subtruncata
Carbon River Entrance, Rainforest Loop Trail, 08-May95, 1 male; Falls Creek at Stevens Canyon Road, 25-Jun00, 1 male.
Deer Creek on Highway 123, 27-May-97, 2 males; Ohanapecosh River at the Grove of the Patriarchs, 28-May-97, 1
male, 1 female; Goat Creek on Highway 410, 29-May-97,
4 males, 1 female; Panther Creek at Highway 123, 29May-97, 1 male; Carbon River at Chenuis Falls Trailhead,
06-Jun-97, 2 females; Ipsut Creek at Carbon River Road
Bridge, 06-Jul-97, 2 females; Glacier Basin, 20-Jul-97, 4
females; Glacier Basin, 21-Jul-97, 4 females; Glacier Basin,
27-Jul-97, 2 females; Fryingpan Creek at Sunrise Road
Bridge, 06-Jul-99,1 male, 1 female; Paradise River above
Narada Falls, 16-Aug-99, 1 female; Paradise River in Paradise Valley, 16-Aug-99, 3 females; Fryingpan Creek at
Sunrise Road Bridge, 16-Aug-99, 1 male, 2 females; Falls
Creek at Stevens Canyon Road, 25-Jun-00, 2 males; Fryingpan Creek at Sunrise Road Bridge, 29-Jun-00, 1 female;
Huckleberry Creek Bridge, 14-May-01, 2 males; Longmire, 15-May-01, 1 male; Ohanapecosh Campground, 16May-01, 1 male; Panther Creek at Highway 123, 09-Jun01, 1 male, 2 females; boggy stream tributary to Tahoma
Creek, 17-Jun-01, 1 male.
Isoperla bifurcata
Carbon River Entrance, 15-Aug-99, 1 female.
Isoperla fusca
Isoperla petersoni
males, 1 female.
Isoperla rainiera
White River Campground, 23-Jul-97, 1 female; Nisqually
River at Glacier Vista Bridge, 07-Jul-99, 1 female; Paradise
River above Narada Falls, 16-Aug-99, 2 males, 1 female;
females; Fryingpan Creek at Wonderland Trail, 31-Oct-00,
1 female.
Isoperla sobria
male, 1 female; White River Highway 410, 0.5 mile S of
park boundary, 16-May-01, 1 male.
Isoperla sordida
Fryingpan Creek at Wonderland Trail, 03-Jul-98, 1 female;
Longmire, 08-Sep-98, 1 female; stream into White River
at Highway 410 N of White River entrance, 07-Jul-99, 3
males; small tributary on Westside Road 2 miles N of Paradise Road, 08-Jul-99, 1 male; boggy stream tributary to
Tahoma Creek, 17-Aug-99, 1 female; Fryingpan Creek at
Sunrise Road Bridge, 05-Sep-99, 1 female; Paradise River
in Paradise Valley, 07-Sep-99, 2 males; Fryingpan Creek at
Sunrise Road Bridge, 13-Sep-99, 1 male; North Puyallup
River at Wonderland Trail, 24-Sep-00, 2 males, 32 females.
Isoperla tilasqua
Stream into White River at Highway 410 N of White
River Entrance, 07-Jul-99, 3 males; tributary to Tahoma
Creek at Paradise Road, 08-Jul-99, 2 males, 3 females;
boggy stream tributary to Tahoma Creek, 17-Aug-99, 1
female.
Megarcys yosemite
Fryingpan Creek at Sunrise Road Bridge, 16-Aug-99, 1
male; Fryingpan Creek at Sunrise Road Bridge, 16-Jul-00,
4 females.
Genus Setvena
Setvena tibialis
Mowich Road at Grindstone Trail Crossing, 07-Jul-95, 1
female; Paradise Ice Caves, 20-Aug-95, 1 male, 1 female;
St. Andrews Creek at Westside Road, 24-Jul-97, 2 females;
Paradise Ice Caves, 13-Sep-97, 1 female; Sunrise Visitor
Center, 13-Aug-98, 1 male; Falls Creek at Stevens Canyon
Road, 06-Jul-99, 3 males, 1 female; Paradise River in Paradise Valley, 16-Aug-99, 2 males; Paradise River at Fourth
Crossing, 16-Aug-99, 1 male; Falls Creek at Stevens Canyon Road, 16-Aug-99, 1 female; small tributary on Westside
Road 2 miles N of Paradise Road, 17-Aug-99, 1 female;
Paradise River in Paradise Valley, 10-Jul-00, 1 male; Paradise River at Fourth Crossing, 23-Jul-00, 1 male.
Genus Skwala
Genus Frisonia
Frisonia picticeps
Skwala americana
Muir Snowfield at 8000 ft, 28-Jul-95, 1 female.
Ohanapecosh Ranger Station, 05-May-96, 1 female; Ohanapecosh Campground, 28-May-97, 2 females; Ohanapecosh
Hot Springs, 28-May-97, 1 female; Ohanapecosh Ranger
Station, 28-May-97, 2 males.
Genus Megarcys
Megarcys irregularis
Nisqually River at Longmire, 28-May-97, 2 males, 1
female; White River Highway 410, 29-May-97, 8 males, 1
female; Tahoma Creek at Nisqually to Paradise Road, 30May-97, 2 females; Fryingpan Creek at Sunrise Road
Genus Kogotus
Kogotus nonus
Ipsut Creek at Carbon River Road Bridge, 08-Sep-97, 1
male; Carbon River Entrance, 15-Aug-99, 1 male, 2 females;
404
female.
Family Pteronarcyidae
Genus Pteronarcys
Pteronarcys princeps
Stevens Canyon Entrance, 15-May-95, 1 female; Panther
Creek at Highway 123, 29-May-97, 1 male; small stream
near Ohanapecosh Visitor Center, 16-May-01, 3 nymphs.
[Volume 62
A 4000-YEAR RECORD OF WOODLAND VEGETATION FROM
WIND RIVER CANYON, CENTRAL WYOMING
Stephen T. Jackson1,3, Mark E. Lyford1, and Julio L. Betancourt2
ABSTRACT.—Plant macrofossil analyses of 16 radiocarbon-dated woodrat middens spanning the past 4000 years from
the Wind River Canyon region in central Wyoming provide information concerning late Holocene development of
juniper woodlands. The study sites are currently dominated by Juniperus osteosperma, with J. scopulorum present
locally. Woodlands in the region were dominated by J. scopulorum from ca 4000 yr BP until at least 2800 yr BP. Juniperus osteosperma invaded and expanded before 2000 yr BP. This expansion fits a regional pattern of J. osteosperma colonization and expansion in north central Wyoming during a relatively dry period between 2800 and 1000 yr BP. At the time
the Wind River Canyon region was colonized by J. osteosperma, the species had populations 50–100 km to both the north
and south. Long-distance seed dispersal was required for establishment in the study area. Genetic studies are necessary
to identify source populations and regions.
Key words: juniper woodlands, vegetation history, woodrat middens, central Wyoming, natural invasions, late Holocene.
Juniper-dominated woodlands are extensive on the flanks of mountain ranges and on
coarse-textured bedrock outcrops in basins of
the central Rocky Mountain region (Knight
1994, West 1999). In Wyoming, juniper woodlands occupy some 5737 km2 (Driese et al.
1997). These woodlands are dominated primarily by Juniperus scopulorum (Rocky Mountain
juniper), which occurs extensively throughout
the Rocky Mountain region, from New Mexico
to Canada. However, scattered juniper woodlands in central Wyoming and adjacent Montana are dominated by J. osteosperma (Utah
juniper; Fig. 1), which occurs primarily in the
Great Basin and Colorado Plateau. The Wyoming and Montana populations represent a
series of isolated outposts separated from each
other by 25–100 km and from the core distribution of the species by 100–500 km. This pattern
contrasts with the distribution of J. osteosperma
in the Colorado Plateau and Great Basin regions,
where the species is abundant and widespread,
and where populations are typically separated
by no more than 10–30 km (Little 1971, West
1999).
Paleoecological records from woodrat middens indicate that J. osteosperma populations
have occurred in the southwestern Great
Basin and southern Colorado Plateau since the
last glacial maximum (Cole 1990, Nowak et al.
1994), and that the species colonized southeastern and northeastern Utah during the early
Holocene (Betancourt 1990, Sharpe 1991).
These observations suggest that the species
migrated from the Utah/Wyoming border north
and east during the Holocene to occupy its
present range. Paleobotanical analyses of an
array of fossil woodrat middens throughout the
modern range of J. osteosperma in Wyoming
and adjacent Utah and Montana are revealing
the timing, rates, and patterns of its Holocene
migration (Lyford et al. 2002a, 2002b).
One of the critical sites in this sampling array
is the Wind River Canyon region of central
Wyoming because of its central location and
the existence of a major north-flowing river
running through the canyon, connecting the
Wind River Basin with the Bighorn Basin (Fig.
1). The canyon may have served as an important migration corridor for J. osteosperma and
other north-moving species during the Holocene. Wind River Canyon is the only canyon
that bisects the Owl Creek Mountains, which
comprise a potential geographic barrier to dispersal between the Bighorn Basin and the Wind
River Basin and other basins to the south (Fig.
1). The Wind River/Bighorn River system comprises one of the few north-flowing rivers in
western North America, and flood flows may
have played an important role in long-distance
1Department of Botany, Aven Nelson Building, University of Wyoming, Laramie, WY 82071.
2U.S. Geological Survey, Desert Laboratory, 1675 W. Anklam Road, Tucson, AZ 85745.
3Corresponding author.
405
406
[Volume 62
Fig. 1. Map of Wyoming and adjacent states showing elevations (white 900–1500 m, light gray 1500–2100 m, medium
gray 2100–2600 m, dark gray ≥ 2600 m), key physiographic features, locations of study sites and other sites mentioned in
text, and distribution of Juniperus osteosperma (black dots) based on records from the Rocky Mountain Herbarium
(http://www.rmh.uwyo.edu).
seed dispersal and Holocene migration of southern species.
We conducted a study of woodrat middens
collected from the Wind River Canyon region
in central Wyoming to better assess the Holocene migration patterns of Juniperus osteosperma, to provide information on the history
of juniper woodlands at the site, and to determine vegetation composition before and after
J. osteosperma colonization (Fig. 1). A 13-midden
series from lower Wind River Canyon spans
the past 4000 years. Three late Holocene middens from Birdseye Creek, 16 km southsoutheast of the Wind River Canyon sites, provide additional information and corroboration.
STUDY SITES
Wind River Canyon is a deep, north-trending canyon, 21 km in length, incised into the
Owl Creek Mountains (Figs. 1, 2). The canyon
cuts through bedrock ranging from Precambrian metavolcanic rocks at the head of the
canyon to limestones of the Permian Phosphoria Formation at the canyon mouth (Maughan
1972, 1987). In between it cuts through a series
of Paleozoic limestones and dolomites, particularly those of the Amsden, Madison, Bighorn,
Gallatin, and Gros Ventre formations. The primary lithologies exposed in the lowermost 4
km of the canyon, where we conducted our
study, are limestones and dolomites of the Park
City, Amsden, and Madison formations.
West- and southwest-facing slopes on the
east side of lower Wind River Canyon are vegetated by juniper woodlands (Fig. 2). Juniperus osteosperma is dominant on these slopes,
although J. scopulorum occurs locally throughout the canyon, especially in mesic sites (draws,
gullies, cliff bases). Rhus trilobata occurs locally
2002]
CENTRAL WYOMING VEGETATION HISTORY
407
Fig. 2. Photo of lower Wind River Canyon near midden-collecting sites. Woodlands in foreground and background are
dominated by Juniperus osteosperma. View is facing upcanyon, WNW. From a color transparency taken May 1998 by
S.T. Jackson.
at cliff bases and in draws. Scattered individuals of Pinus flexilis occur throughout the
canyon but are not abundant on the lower
slopes where we concentrated our collecting
efforts. Other plant species on the slopes include shrubs (Ribes aureus [at cliff bases],
Artemisia frigida, Atriplex canescens, Chrysothamnus spp., Gutierrezia sarothrae), grasses
(Oryzopsis hymenoides, Bromus tectorum, Stipa
comata, Elymus sp.), forbs (Artemisia ludoviciana, Cryptantha spp., Eriogonum spp., Penstemon spp.), and succulents (Opuntia polyacantha).
Birdseye Creek flows south-southwest,
draining uplands of the Owl Creek (Bridger)
Mountains immediately east of Wind River
Canyon. The creek cuts through a highly faulted
series of Paleozoic limestones and dolomites,
locally capped by Eocene sediments of the
Wind River Formation (Thaden 1980). Middens
are in crevices of the Madison Formation. Vegetation in Birdseye Creek Canyon consists of a
mosaic of steppe, dominated by Artemisia tridentata, and open juniper woodlands, domi-
nated by Juniperus osteosperma. Both vegetation types occur within 20 m of our collecting
sites. Subdominants include the same species
as observed in lower Wind River Canyon.
The climate of central Wyoming is semiarid.
Mean annual precipitation at Thermopolis
(1948–2000; 1342 m elevation; 8 km N of the
Wind River Canyon sites) is 290 mm. Mean
annual precipitation at Boysen Dam, at the
head of Wind River Canyon (1948–2000; 1416
m elevation; 7.5 km E of the Birdseye Creek
sites) is 233 mm. Despite its higher elevation,
Boysen Dam receives less precipitation than
Thermopolis owing to its isolation at the
northern edge of the Wind River Basin, where
moisture sources are blocked by the Owl Creek
Mountains. Mean July precipitation at Thermopolis and Boysen Dam is respectively 21
and 13 mm; mean January precipitation is
respectively 10 and 7 mm. Winters are cold
(mean January temperature –5.7°C at Thermopolis and –7.8°C at Boysen Dam) and summers are hot (mean July temperature 22.2°C
at Thermopolis and 23.8°C at Boysen Dam).
408
METHODS
Midden-collecting efforts in Wind River
Canyon were concentrated on west-facing
slopes in the lower portion of the canyon,
where J. osteosperma is most abundant. The
collecting sites range in elevation from 1365 to
1475 m (Table 1); river elevation in this reach
of the canyon is 1340 m and the rim ranges
from 1400 to 1650 m. Using standard field
techniques (Spaulding et al. 1990), we collected
19 midden samples from crevices and bedrock
overhangs. We collected 6 midden samples
from Birdseye Creek Canyon, ranging from
1615 to 1700 m in elevation (Table 1). In all
cases, weathering rinds were removed in the
field, middens were inspected for stratification
and contamination, and a sample from the
core of the midden was collected. In the laboratory, midden samples were reinspected, dispersed in water, oven-dried, and sieved using
1- and 2-mm mesh screens (Spaulding et al.
1990). We examined the 2-mm sieve fraction
thoroughly, separating identifiable plant macrofossils and identifying them using herbariumdocumented reference collections. The 1-mm
sieve fraction was scanned to identify plant
species and morphotypes not represented in
the larger fraction. Each plant taxon in each
midden was assigned a relative abundance
value (1 = single occurrence, 5 = dominant;
Spaulding et al. 1990).
Thirteen middens were selected for radiocarbon-dating from Wind River Canyon, and 3
from Birdseye Creek. We obtained conventional 14C dates on samples of Neotoma fecal
pellets from 8 Wind River Canyon and 2 Birdseye Creek middens. Using accelerator mass
spectrometry (AMS), we dated Juniperus osteosperma foliage from 6 Wind River Canyon middens and 2 Birdseye Creek middens. We also
AMS-dated J. scopulorum foliage from another
Birdseye Creek midden. AMS dating provides
precise dates from small amounts of organic
material and hence yielded direct age-estimates for Juniperus macrofossils from the middens. We used AMS dating to pinpoint temporal occurrences of J. osteosperma at the study
sites and in cases where we suspected possible mixing of plant material of different ages.
For the conventional dates, we submitted
3–10 g of Neotoma fecal pellets to Geochronology Laboratories (Cambridge, MA) for dating. Targets for AMS dating were prepared at
[Volume 62
the USGS Desert Laboratory in Tucson. Samples were pretreated to remove carbonates
and acid- and base-soluble organic matter,
combusted to CO2 on a vacuum line, converted
to graphite, pressed into targets, and then
measured for 14C activity at the University of
Arizona–NSF Accelerator Facility.
All ages reported or discussed in this paper
are calendar-year ages (years Before Present,
with 1950 as the benchmark). Radiocarbon ages
were converted to calendar-year ages using
the Intcal 98 calibration curve, using Method
A (ranges with intercepts) from CALIB 4.3
(Stuiver and Reimer 1993).
RESULTS
The Wind River Canyon middens range in
age from 3876 yr BP to modern (Table 1, Fig.
3). Macrofossil assemblages from the 6 middens dating before 2500 BP show relatively
little variation; all are dominated by Juniperus
scopulorum, with Rhus trilobata and Opuntia
also abundant (Fig. 3, Table 2). Most have
macrofossils of J. osteosperma. The 5 middens
post-dating 2000 yr BP contain both J. osteosperma and J. scopulorum. Rhus and Opuntia
occur as subdominants. Atriplex sp. is absent
from all of these younger middens, and Artemisia nova occurs only in the oldest of these
middens (Fig. 3).
One Birdseye Creek midden is dominated
by J. scopulorum and lacks macrofossils of J.
osteosperma (Fig. 3, Table 2). It contains macrofossils of Artemisia nova and Atriplex sp. This
midden was dated at 3350 yr BP, and foliage of
J. scopulorum was AMS-dated at 2923 yr BP.
Juniperus osteosperma foliage from a 2nd midden, which also contains J. scopulorum macrofossils, was AMS-dated at 2137 yr BP. A 3rd
midden, dominated by J. osteosperma, yielded
modern dates (45 yr BP; 42 yr BP AMS). Both
of these younger middens lack Artemisia nova
and Atriplex sp.
DISCUSSION
During the last glacial period, Juniperus
osteosperma populations were restricted to the
southern Great Basin and southern Colorado
Plateau (Betancourt 1990, Thompson 1990,
Nowak et al. 1994, Lyford et al. 2002b). During the late-glacial (13,000–10,000 yr BP), J.
osteosperma migrated rapidly from northern
Arizona to northeastern Utah (Betancourt 1990,
Lab no.
AA33168
AA38266
AA38265
AA33162
AA38242
AA33156
GX24953
GX24952
GX24951
GX24948
GX24950
GX25842
GX26611
GX24949
GX25480
AA33680
AA38258
AA33673
GX25479
Midden no.
WRC207B
WRC240
WRC208
WRC241B
WRC173
WRC212
WRC242
WRC183
WRC181C
WRC171A
WRC173
WRC182
WRC181A
WRC172
BEC245A
BEC245A
BEC301
BEC174A
BEC174A
J. osteosperma foliage
Neotoma fecal pellets
Material dated
Age (cal-yr BP)
467 (309–514)
505 (341–527)
679 (654–760)
1562 (1422–1708)
1675 (1530–1713)
1657 (1529–1815)
1909 (1723–2118)
2750 (2351–2992)
2781 (2736–2965)
2848 (2747–3058)
3469 (3274–3687)
3655 (3471–3889)
3876 (3698–4086)
3876 (3644–4144)
45 (0–285)
42 (0–277)
2137 (1996–2325)
2923 (2780–3157)
3350 (2952–3633)
Age (14C yr BP)
375 ± 45
430 ± 33
767 ± 43
1680 ± 50
1719 ± 41
1735 ± 50
1970 ± 75
2620 ± 130
2710 ± 80
2750 ± 75
3260 ± 80
3430 ± 80
3590 ± 60
3590 ± 80
80 ± 65
90 ± 45
2155 ± 50
2825 ± 60
3110 ± 130
43°33′28.8″N
43°33′10.1″N
43°33′28.8″N
43°33′10.1″N
43°33′52.8″N
43°33′34.6″N
43°33′10.1″N
43°33′14.9″N
43°33′14.9″N
43°33′44.2″N
43°33′52.8″N
43°33′14.9″N
43°33′14.9″N
43°33′52.8″N
43°23′37.1″N
43°23′37.1″N
43°24′06.7″N
43°23′34.1″N
43°23′34.1″N
Latitude
108°12′20.8″W
108°11′42.6″W
108°12′20.8″W
108°11′42.6″W
108°12′30.3″W
108°12′22.1″W
108°11′42.6″W
108°12′37.6″W
108°12′37.6″W
108°12′42.5″W
108°12′30.3″W
108°12′37.6″W
108°12′37.6″W
108°12′30.3″W
108°05′17.1″W
108°05′17.1″W
108°05′16.2″W
108°05′15.1″W
108°05′15.1″W
Longitude
1455
1421
1455
1421
1431
1482
1421
1416
1416
1367
1431
1416
1416
1431
1645
1645
1657
1639
1639
Elevation (m)
TABLE 1. Radiocarbon age, location, and elevation of middens analyzed from Wind River Canyon (WRC) and Birdseye Creek (BEC). Age estimates in 14C yr BP include 2 standard
errors. Age estimates in calendar-yr BP include median intercept from CALIB 4.2 (Stuiver et al. 1998) and (in parentheses) minimum and maximum ages based on 2 standard errors
from minimum and maximum intercepts.
2002]
409
410
[Volume 62
Fig. 3. Relative abundances of macrofossils of selected taxa in woodrat middens from Wind River Canyon (WRC) and
Birdseye Creek (BEC).
Sharpe 1991). It was established at Dutch John
Mountain, 5 km south of the Utah/Wyoming
border, by 9400 yr BP ( Jackson, Betancourt,
and Lyford, unpublished data). Northeastern
Utah was the most likely portal for invasion of
J. osteosperma and other Great Basin species
into Wyoming. The modern distribution of J.
osteosperma as well as the topographic configuration suggests that migration from the Utah/
Wyoming border northward proceeded from
southwestern Wyoming to the southeastern
foothills of the Wind River Mountains, and
then across the Owl Creek/Bridger Mountains
into the Bighorn Basin (Fig. 1). As the only
drainage cutting across the Owl Creek Mountains, Wind River Canyon would appear to
have been an important migration pathway.
Establishment of J. osteosperma populations
at several sites in the Bighorn Basin predates
the 1st occurrences (1900 yr BP at Wind River
Canyon, 2100 at Birdseye Creek) in the Wind
River Canyon region. The earliest documented
establishments occurred ca 5400 yr BP at
Southern Bighorn Canyon, at the northern
edge of the Bighorn Basin (Lyford et al. 2002a),
and Mahogany Butte, in the southeastern Bighorn Basin (Lyford et al. 2002b; Fig. 1). However, these occurrences predate not only the
1st J. osteosperma occurrences in the Wind
River Canyon region, but also the oldest middens we have in that region (3900 yr BP).
The migration of J. osteosperma in Wyoming
occurred by means of rare, long-distance dispersal events spanning 30–100 km or more.
Such dispersal events were required to cross
geographic barriers (e.g., the Wind River Basin)
and are confirmed by the patchy pattern of
establishment documented from woodrat midden analyses throughout the region (Lyford et
al. 2002b). The Wind River Canyon region may
–
–
–
–
–
–
–
–
–
–
2
3
5
–
2
2
–
–
–
–
2
–
–
–
2
–
–
–
–
–
1.5
Artemisia nova (lf)
Artemisia tridentata (lf)
Atriplex sp. (fr)
Atriplex sp. (lf)
Chrysothamnus sp. (inv)
Cryptantha sp. (lf)
Elymus sp. (fl)
Equisetum sp. (st)
Gutierrezia sarothrae (inv)
Hedeoma sp. (se)
Juniperus osteosperma (fr)
Juniperus osteosperma (se)
Juniperus osteosperma (st)
Juniperus scopulorum (fr)
Juniperus scopulorum (se)
Juniperus scopulorum (st)
Lappula redowskii (se)
Lesquerella sp. (lf)
Lithospermum sp. (se)
Opuntia sp. (se)
Opuntia sp. (st)
Oryzopsis hymenoides (se)
Pinus flexilis (n)
Pinus flexilis (se)
Rhus trilobata (se)
Rosa sp. (lf)
Rosa sp. (se)
Rosa sp. (st)
Stipa sp. (fl)
Stipa comata (se)
Yucca glauca (se)
–
–
–
–
–
–
–
–
–
–
–
3
4
–
2
4
–
–
1
–
2
–
–
–
5
–
–
–
–
–
–
505
–
–
–
–
–
–
–
–
–
–
2
2
5
–
2
2
–
–
–
–
2
–
–
–
1
–
–
–
–
–
–
679
–
–
–
–
–
–
–
–
–
–
2
3
5
–
2
2
–
–
–
2
2
1
–
–
2
–
–
–
–
–
–
1562
–
–
–
–
–
–
–
–
–
–
2
2
2
2
2
5
–
–
–
–
2
2
2
–
2
–
–
–
–
–
–
1657
2
–
–
–
–
–
–
–
2
1
2
2
4
2
3
4
–
–
2
–
2
2
–
–
2
–
–
–
–
–
3
1909
1
–
–
–
–
–
2
–
–
–
–
–
–
–
3
5
–
–
–
–
2
–
–
–
2
–
–
–
–
–
–
2750
2
–
–
–
–
–
–
2
–
–
–
–
–
2
3
5
–
2
–
2
2
1
–
–
2
3
2
2
–
–
–
2781
2
–
3
2
–
–
–
–
2
1
–
–
–
2
4
5
–
2
1
–
2
–
–
–
2
–
–
–
1
–
–
2848
2
–
–
–
–
2
–
–
–
–
–
–
–
2
4
5
1
–
1
2
–
2
4
–
2
–
–
–
–
–
–
3469a
2
–
2
–
–
2
–
1
–
–
–
–
–
2
2
5
–
–
–
–
–
–
–
–
2
–
–
–
–
–
–
3566
–
–
1
–
–
–
1
2
–
–
–
–
–
2
2
5
–
–
–
–
2
–
–
–
2
2
–
–
–
–
–
3876
–
2
2
2
2
–
–
–
–
–
–
–
–
3
4
5
–
–
–
–
–
–
2
1
2
–
–
–
–
–
–
3876
–
–
–
–
–
–
–
–
–
–
1
2
5
–
–
–
–
–
–
–
2
–
–
–
–
–
–
–
–
–
–
45
–
–
–
5
–
–
–
–
–
–
–
–
2
–
2
2
–
–
–
–
3
–
–
–
2
–
–
–
–
–
–
2137
2
–
2
4
–
2
2
–
–
–
–
–
–
–
2
4
–
2
–
2
3
2
–
–
4
–
–
–
–
2
–
3350b
Birdseye Creek
_____________________
245A
301
174A
_____________________
aFoliage of Juniperus osteosperma from this midden was AMS–dated at 1719 radiocarbon yr BP (1675 calendar yr BP). This is regarded as a contaminant; the rest of the midden contents are assigned the age obtained from the Neotoma fecal pellets.
bFoliage of Juniperus scopulorum from this midden yielded a slightly younger age (see Table 1).
467
Sample age (yr BP)
Sample number
Wind River Canyon
_____________________________________________________________________________________________________
207B
240
208
241B
212
242
183
181C
171A
173
182
181A
172
_____________________________________________________________________________________________________
TABLE 2. Relative abundances of plant macrofossil types in woodrat middens from Wind River Canyon and Birdseye Creek sites. Organ types abbreviated as follows: lf = leaf, n =
needle, fl = floret, se = seed, inv = involucre, fr = fruit, st = stem.
2002]
411
412
have been skipped in long-distance dispersal
events during the mid-Holocene, leading to
the initial establishment well north of the region. Under this scenario, the region was first
colonized by J. osteosperma ~2000 yr BP, either
by back-colonization from the Bighorn Basin
to the north or by additional dispersal events
from populations to the south (SE Wind River
Mountains, SW Wyoming).
Alternatively, Wind River Canyon may have
been colonized by J. osteosperma populations
before 5400 yr BP, and did in fact serve as an
intermediate site in northward invasion into the
Bighorn Basin region. Under this scenario, early
populations of J. osteosperma underwent local
extinction some time before 3900 yr BP, with
recolonization ca 2000 yr BP. This hypothesis
cannot be excluded given the absence of middens predating 3900 yr BP.
Lyford et al. (2002b) identified a punctuated
pattern of J. osteosperma migration in northern Wyoming in which establishment occurred
during periods of relatively dry climate (ca
5400 yr BP, 2800–1000 yr BP) and colonization
of new sites ceased during relatively wet periods (5400–2800 yr BP). Our data are consistent with this pattern; J. osteosperma is absent
from all middens representing the wet period
(3900–2800 yr BP), and the 1st documented
occurrences (2100–1900 yr BP) occurred during the dry period.
The populations established 5400 yr BP at
southern Bighorn Canyon and Mahogany Butte
persisted throughout the subsequent wet period
(Lyford et al. 2002a, 2002b). Because the Mahogany Butte site is higher (1800 m) and wetter than Wind River Canyon and Birdseye
Creek, Mahogany Butte populations might be
expected to be more sensitive to a moisture
increase than those in the Wind River Canyon
region. Juniperus osteosperma populations
established in Wind River Canyon at or before
5400 yr BP may have declined but persisted
locally after 5400 yr BP, and then expanded in
response to the drying trend after 2800 yr BP.
The absence of J. osteosperma from the 3 middens dating from 2750 to 2850 yr BP as well as
the 5 middens dating 3350–3900 yr BP suggests, however, that J. osteosperma was absent
from the region during the wet period and
that the species invaded the canyon by dispersal from distant sources (e.g., in the eastern
Bighorn Basin). Woodrats of Neotoma cinerea
[Volume 62
show a strong preference for collecting Juniperus foliage, particularly J. osteosperma (Nowak
et al. 2000, Lyford 2001), increasing our confidence that absence of J. osteosperma from the
middens predating 2100 yr BP indicates its
absence from the surrounding vegetation.
Our paleological data provide constraints
on the history of J. osteosperma in the Wind
River Canyon region and suggest strongly that
current populations were established ca 2000
yr BP from distant sources to the north or south.
Those populations may carry a genetic signature of their establishment sources and history.
Genetic analyses of the Wind River Canyon
populations and potential source populations
in the Wind River foothills and the Bighorn
Basin might reveal the precise pathway and
sequence of invasion of Wind River Canyon.
Juniperus scopulorum populations persisted
in Wind River Canyon after the rapid transition to dominance by J. osteosperma, in contrast to sites in the Bighorn Basin (e.g., Pryor
Mountain and Big Horn Canyon; Lyford et al.
2002a). Wind River Canyon is more mesic than
the northern sites, owing to shading by the
west canyon wall and to greater surface drainage and groundwater seepage from upper
slopes. The locally moist habitats have allowed
persistence of J. scopulorum over much of the
canyon. We cannot determine from the midden
records whether J. scopulorum was displaced
from drier sites when J. osteosperma invaded.
Rhus trilobata and Opuntia occurred in
Wind River Canyon both before and after the
establishment of J. osteosperma. Rhus trilobata
is concentrated around locally moist sites
(gulches, seeps, cliff bases) and was thus buffered against changes in moisture of the magnitude experienced in the past 4000 years. Similarly, Opuntia occurs widely in dry, exposed
microsites in the canyon, which have been
available regardless of climate regime. The reason for the disappearance of Artemisia nova
from the canyon between 2500 and 2000 yr
BP is unclear. The species did not occur near
any of our study sites and is uncommon in the
canyon. Its disappearance may have resulted
from increasing density of woodlands as J.
osteosperma populations expanded. The late
Holocene appearance of Yucca glauca at Wind
River Canyon matches a pattern observed in
the northern Bighorn Basin by Lyford et al.
(2002a).
2002]
ACKNOWLEDGMENTS
This research was supported by grants to
the University of Wyoming and the University
of Arizona from the National Science Foundation (Ecology and Paleoclimatology Programs).
Invaluable assistance in the field and laboratory was provided by Gabe Cisneros, Mark
Betancourt, Jodi Norris, Camille Holmgren,
Kate Rylander, and Rob Eddy. The manuscript
was improved by comments from 2 anonymous reviewers.
LITERATURE CITED
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the Colorado Plateau. Pages 259–292 in J.L. Betancourt, T.R. Van Devender, and P.S. Martin, editors,
Packrat middens: the last 40,000 years of biotic
change. University of Arizona Press, Tucson.
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through the Grand Canyon. Pages 240–258 in J.L.
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DRIESE, K.L., W.A. REINERS, E.H. MERRILL, AND K.G.
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USA: a tool for vegetation analysis. Journal of Vegetation Science 8:133–146.
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Wyoming landscapes. Yale University Press, New
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States Department of Agriculture, Forest Service,
Miscellaneous Publication 1146.
LYFORD, M.E. 2001. The roles of dispersal, climate, and
topography in the Holocene migration of Utah juniper
into Wyoming and southern Montana. Doctoral dissertation, University of Wyoming, Laramie.
LYFORD, M.E., J.L. BETANCOURT, AND S.T. JACKSON. 2002a.
Holocene vegetation and climate history of the
northern Bighorn Basin, southern Montana. Quaternary Research: In press.
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GRAY. 2002b. Anatomy of a late Holocene plant
migration: influence of environmental heterogeneity
and climate variability. Ecology: In review.
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196 in S.S. Beus, editor, Geological Society of America Centennial field guide—Rocky Mountain section. Geological Society of America, Boulder, CO.
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1994. A 30,000 year record of vegetation dymanics at
a semi-arid locale in the Great Basin. Journal of Vegetation Science 5:579–590.
NOWAK, R.S., C.L. NOWAK., AND R.J. TAUSCH. 2000. Probability that a fossil absent from a sample is also
absent from the paleolandscape. Quaternary Research
54:144–154.
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County, Utah, and Dinosaur National Monument,
Moffat County, Colorado. Master’s thesis, Northern
Arizona University, Flagstaff.
SPAULDING, W.G., J.L. BETANCOURT, L.K. CROFT, AND K.L.
COLE. 1990. Packrat middens: their composition and
methods of analysis. Pages 59–84 in J.L. Betancourt,
T.R. Van Devender and P.S. Martin, editors, Packrat
middens: the last 40,000 years of biotic change. University of Arizona Press, Tucson.
STUIVER, M., AND P.J. REIMER. 1993. Extended 14C database and revised CALIB 3.0 14C age calibration program. Radiocarbon 35:215–230.
THADEN, R.E. 1980. Geologic map of the Birdseye Pass
Quadrangle, showing chromolithofacies [sic] and coal
beds in the Wind River Formation, Fremont and Hot
Springs counties, Wyoming. U.S. Geological Survey
Geologic Quadrangle Map GQ-1537. Washington,
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THOMPSON, R.S. 1990. Late Quaternary vegetation and
climate in the Great Basin. Pages 200–239 in J.L.
Betancourt, T.R. Van Devender, and P.S. Martin, editors, Packrat middens: the last 40,000 years of biotic
WEST, N.E. 1999. Juniper-piñon savannas and woodlands
of western North America. Pages 288–308 in R.C.
Anderson, J.S. Fralish, and J.M. Baskin, editors,
Savannas, barrens, and rock outcrop plant communities of North America. Cambridge University Press.
Received 6 June 2001
Accepted 11 March 2002
Western North American Naturalist 62(4), ©2002, pp. 414–422
VEGETATION RESPONSE TO PRESCRIBED
FIRE IN DINOSAUR NATIONAL MONUMENT
Barry L. Perryman1, Richard A. Olson2, Stephen Petersburg3, and Tamara Naumann3
ABSTRACT.—Much of western North America is dominated by dense, monotypic, late seral stands of big sagebrush
(Artemisia tridentata Nutt.). These stands often have depauperate understories with limited species richness, diversity,
and herbaceous cover. The National Park Service at Dinosaur National Monument, Colorado, is using both strategic and
natural prescribed fire in Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis Beetle and Young) communities to foster intra-community (α-scale) and landscape diversity. This study analyzed an accumulated foliar cover data set
between paired burn and control areas on 6 different sites during the last 20 years. Across the monitoring period, mean
total vegetation cover of all combined sites was 44% control and 42% burn. Total vegetation cover in burn areas was
higher than or equal to paired control areas within 2–3 years post-burn. Shrubs were essentially eliminated in burn
areas, but perennial grass cover was 10–35% higher. Mean number of species on all sites and years combined was 17
control and 18 burn. Species richness was different on only 1 site-year, Dry Woman 1995 (P = 0.001, 15 control, 9 burn).
Species similarity by site and between treatments ranged from 44% to 75%. Differences in Shannon-Weiner diversity
index values between paired sites occurred in 6 of 20 years (P < 0.05). Index value differences on these 6 sites were due
to a large annual grass component in burn areas. Prescribed burning successfully shifted late successional sagebrushdominated communities to earlier herbaceous-dominated successional stages without lowering total vegetation cover,
while maintaining α-scale diversity and species richness.
Key words: prescribed fire, vegetation cover, community diversity, species richness.
More than 54 million ha in western North
America are dominated by big sagebrush (Artemisia tridentata Nutt.; Whitson and Alley 1984).
Late seral or climax sagebrush stands are often
dense and monotypic with limited species richness, diversity, and herbaceous cover in the
understory (Johnson et al. 1996, West 1999), a
condition due in large part to fire suppression
activities and/or poor grazing management. In
either case decadent, late seral sagebrush communities are left in a stable ecological state that
requires energy input to shift the stand from a
shrub- to a grass-dominated domain (Archer
1989, Laycock 1991). Brush control measures
in decadent sagebrush stands foster intra-community (α-scale) species richness and diversity
(Olson et al. 1994, Johnson et al. 1996) as well
as landscape patch richness and diversity.
Anecdotal accounts by early residents of the
Dinosaur National Monument, Colorado, area
suggest large areas of the monument were
dominated at that time by native perennial
grasslands. Domestic livestock began grazing
the monument area in the late 1800s, increased
beginning in the 1920s, reached a peak autho-
rized level of 35–40,000 AUMs on 56,000 ha in
the 1940s, and, based on rangeland surveys,
reduced to 5000 AUMs by 1973. Resource
managers believe that intensive grazing pressure during those 2 decades, in coincidence
with the drought of the 1930s and the initiation
of effective fire suppression activities circa
1940, all contributed to a shift from grassland
to big sagebrush–dominated communities.
Under a 1916 Congressional mandate to conserve and protect the natural resources of
National Park Service lands, resource managers initiated a complex fire management
program in the early 1980s. The program includes both strategic prescription burns and
prescribed natural fires. Prescribed natural
fires are defined as natural ignitions allowed
to burn within strict constraints of location,
proximity to monument boundaries, and threats
to life and property. Both ignition sources reintroduced fire into stable, decadent Wyoming
sagebrush (Artemisia tridentata ssp. wyomingensis Beetle and Young) stands. The goal is
to shift community composition from brush
to perennial grassland dominance and foster
1School of Veterinary Medicine, College of Agriculture, Biotechnology, and Natural Resources, University of Nevada–Reno, MS 202,
2Department of Renewable Resources, College of Agriculture, University of Wyoming, PO Box 3354, Laramie, WY 82071.
3USDI, National Park Service, Dinosaur National Monument, 4545 East Hwy 40, Dinosaur, CO 81610.
414
Reno, NV 89557.
2002]
VEGETATION RESPONSE TO PRESCRIBED FIRE
α-scale and landscape diversity in these plant
communities. A post-fire monitoring program
was implemented in 1984 after the 1st prescribed burn with follow-up sampling in succeeding years. In 1998 the National Park Service decided to analyze the post-fire monitoring data collection.
The specific objectives of this project were
to (1) assess plant cover trends in paired burn
and control areas, (2) assess intra-community
(α-scale) plant species richness and diversity
between paired burn and control areas, and
(3) assess species richness similarity between
burned and control species suites.
STUDY AREA
Study sites are within the boundary of Dinosaur National Monument, Colorado/Utah, on
the east end of the Uinta Mountains in the
northwest corner of Colorado with a small portion in northeastern Utah. The monument is
centered on the confluence of the Green and
Yampa rivers and best described as bench and
canyon topography. Elevations range from
1700 to 2740 m. Annual precipitation ranges
from 20 to 38 cm.
Predominant vegetation type varies by elevation. At lower elevations near springs and
water courses, riparian tree and shrub communities are represented by willows (Salix
sp.), boxelder (Acer negundo L.), narrowleaf
cottonwood (Populus angustifolia James), water
birch (Betula occidentalis Hook.), and Wood’s
rose (Rosa woodsii Lindl.). On drier sites predominant plant species are basin big sagebrush, Wyoming big sagebrush, rubber rabbitbrush (Chrysothamnus nauseosus [Pallus ex
Pursh] Britt.), serviceberry (Amelanchier alnifolia Nutt.), black greasewood (Sarcobatus vermiculatus [Hook.] Torrey), plains prickly pear
(Opuntia polyacantha Haw.), winterfat (Ceratoides lanata Pursh), sand verbena (Verbena
stricta Vent.), and buckwheat (Eriogonum sp.).
Rocky canyon slopes are characterized as a
pinyon pine (Pinus edulis Engelm.) / juniper
( Juniperus osteosperma [Torrey] Little) woodland with mountain mahogany (Cercocarpus
montanus Raf.) and Mormon tea (Ephedra viridis Coville). High-elevation areas are typically
dominated by ponderosa pine (Pinus ponderosa Dougl. ex Laws.) and Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco), with scattered
pockets of aspen (Populus tremuloides Michx.).
415
Six sites (East Cactus, West Cactus I, West
Cactus II, Success, Iron Springs Bench, and
Dry Woman) having similar elevation, annual
precipitation amounts, soils, and vegetation
were selected for study. Elevation at these
sites is approximately 1950 m with mean annual
precipitation of 20–25 cm. Soils are primarily
Mollisols, formed under grasslands, with a
dominant Wyoming big sagebrush and mixedgrass community. Forward rates of fire spread
on all 6 sites ranged between 60 and 100 chains
hour –1, with residence time over perennial
grass root crowns less than 20–30 seconds and
surface temperatures less than 120°C. All burns
were initiated during the late growing season
when fine fuel loads were at annual maximum.
METHODS
Accumulated monitoring data sets from the
6 prescribed/natural burn sites in Dinosaur
National Monument were analyzed to determine fire effects on plant foliar cover and
diversity. Sites and sample years included in
the analysis are listed in Table 1. Percent cover
categories include total vegetation, herbaceous
(combined grass and forb cover), shrub, grass,
forb, litter, and bare ground. For each site
paired burn and control sites were compared
using a paired t test. Cover data were collected
during the peak of the growing season by the
National Park Service and generated from
both Daubenmire quadrat (Daubenmire 1959)
prior to 1992 and optical projection point measurements beginning in 1992 (Winkworth and
Goodall 1962; single or 1st hits were recorded
with the optical device). Percent values were
conditioned for analyses using the arcsine transformation procedure to achieve normal distribution (Zar 1999). Arcsine values were used in
all statistical analyses, but cover is reported as
percent values. Differences were determined
at P ≤ 0.1 on all cover analyses.
Species richness (number of species) was
analyzed with paired t tests and by determining percent of similarity ([number of shared
species/total species] × 100) between paired
areas on each site by year and all years combined. Diversity (species richness and evenness) assessments were performed using cover
data for each identified species in each set of
paired plots by year. A Shannon-Weiner index
value (Shannon and Weaver 1949) was calculated for each site pair by year and a modified
Year
1988
1989
1991
1992
1994
1987
1988
1990
1993
1989
1990
1995
1988
1993
1996
1997
1984
1987
1990
1995
Site
East Cactus
West Cactus I
West Cactus II
Success
Iron Springs
Bench
Dry Woman
1
1
1
5
4
4
2
4
2
2
5
2
2
2
5
2
2
2
3
3
n
48
46
36
58(±1)
45(±2)
71(±2)
24(±9)
29(±2)
31(±5)
41(±1)
60(±2)
79(±4)
51(±3)
53(±5)
47(±2)
47(±6)
34(±1)
34(±0)
47(±3)
49(±3)
31
66
44
66(±4)
33(±3)
83(±7)
34(±18)
43(±2)
27(±2)
36(±1)
61(±3)
56(±4)
35(±3)
51(±1)
47(±1)
14(±3)
26(±1)
16(±0)
60(±3)
42(±2)
0.079
0.021
0.145
0.427
0.01
0.373
0.172
0.908
0.212
0.250
0.748
0.987
0.187
0.129
0.013
0.008
0.262
Total vegetation
_______________________________
Control
Burn
P-value
10
11
5
23(±1)
34(±3)
36(±2)
12(±2)
16(±2)
3(±0)
6(±1)
20(±2)
41(±1)
32(±3)
30(±1)
25(±1)
25(±5)
14(±1)
22(±1)
16(±2)
28(±3)
0
1
2
0(±0)
0(±0)
0(±0)
0(±0)
0(±0)
0(±0)
0(±0)
3(±1)
0(±0)
0(±0)
0(±0)
0(±0)
0(±0)
0(±0)
0(±0)
1(±0)
0(±0)
0.001
0.001
0.001
0.5
0.002
0.058
0.079
0.001
0.001
0.05
0.009
0.001
0.10
0.022
0.016
0.014
0.011
Shrub
_______________________________
Control
Burn
P-value
TABLE 1. List of sites, years, sample size (n), mean cover (%), P-values and standard errors of vegetation cover classes by treatment.
37
36
30
33(±2)
10(±1)
34(±1)
13(±2)
13(±1)
28(±5)
36(±1)
37(±2)
37(±4)
18(±1)
23(±4)
21(±3)
22(±2)
20(±1)
12(±1)
30(±2)
19(±2)
31
66
43
62(±4)
33(±3)
82(±6)
34(±18)
42(±3)
27(±2)
36(±1)
57(±3)
56(±4)
35(±3)
51(±1)
47(±1)
14(±3)
26(±1)
16(±1)
57(±3)
42(±3)
0.002
0.005
0.005
0.469
0.005
0.642
0.844
0.002
0.248
0.21
0.124
0.001
0.338
0.189
0.030
0.002
0.033
Herbaceous
_______________________________
Control
Burn
P-value
416
[Volume 62
19(±3)
23(±2)
40(±2)
31(±18)
37(±2)
25(±3)
60(±4)
15
50
28
38(±3)
18(±2)
22(±2)
24(±1)
10(±1)
8(±1)
9(±1)
23(±1)
23
20
15
20(±1)
0.008
0.018
0.004
0.469
0.001
0.716
0.806
0.005
0.161
0.042
0.026
0.007
15
16
16
13(±2)
2(±1)
11(±1)
2(±1)
5(±1)
10(±3)
14(±2)
13(±1)
12±(4)
7(±2)
12(±4)
9(±1)
41(±4)
27(±2)
41(±1)
37(±4)
16
16
16
24(±2)
7(±2)
23(±2)
4(±1)
5(±1)
8(±1)
13(±1)
18(±3)
15(±1)
8(±1)
10(±0)
11(±3)
0.008
0.067
0.019
0.479
0.883
0.663
0.795
0.141
0.571
0.795
0.781
0.584
0.175
0.033
0.451
0.031
0.774
26(±1)
11(±1)
12(±1)
11(±2)
2(±0)
4(±0)
4(±0)
17(±4)
8(±3)
5(±1)
7(±0)
5(±1)
10(±5)
7(±1)
12(±3)
23(±1)
11(±1)
42(±6)
34(±1)
17(±3)
12(±1)
8(±1)
16(±3)
12(±2)
0.572
0.133
0.129
0.022
0.021
Forb
_______________________________
Control
Burn
P-value
Grass
_______________________________
Control
Burn
P-value
TABLE 1. Continued.
40
36
28
25(±1)
35(±1)
18(±1)
20(±3)
16(±1)
28(±4)
25(±4)
23(±2)
32(±2)
41(±1)
22(±1)
32(±1)
38(±5)
22(±1)
28(±8)
20(±3)
23(±3)
18
39
19
23(±1)
3(±0)
13(±3)
36(±14)
43(±2)
19(±1)
9(±1)
30(±2)
23(±4)
15(±7)
28(±2)
36(±2)
14(±1)
12(±1)
11(±2)
21(±1)
42(±4)
0.372
0.001
0.131
0.529
0.001
0.283
0.167
0.042
0.078
0.154
0.112
0.182
0.140
0.125
0.301
0.872
0.011
Litter
_______________________________
Control
Burn
P-value
53
30
45
18(±1)
18(±3)
11(±2)
45(±9)
49(±1)
33(±4)
42(±4)
18(±1)
38(±2)
45(±4)
42(±0)
22(±2)
41(±7)
46(±1)
57(±3)
28(±1)
31(±5)
68
29
44
23(±1)
63(±2)
18(±1)
39(±5)
15(±1)
50(±4)
53(±7)
14(±1)
50(±8)
54(±6)
40(±5)
17(±3)
84(±1)
52(±4)
74(±1)
23(±1)
16(±4)
0.64
0.001
0.039
0.777
0.001
0.272
0.503
0.079
0.291
0.523
0.775
0.358
0.09
0.38
0.29
0.124
0.128
Bare ground
_______________________________
Control
Burn
P-value
2002]
417
418
t test performed for each paired set of index
values (Zar 1999). When differences in paired
index values occurred, a Bray-Curtis polar
ordination (Gauch 1982) was performed to
assess similarity/dissimilarity of the paired species assemblages. Diversity differences were
determined at P < 0.05.
Statistical analyses were performed on all
sites (and years) where data existed for more
than 1 transect. Where only 1 transect was sampled, exact values were recorded and summarized.
RESULTS
Vegetation Cover
Across the monitoring period, mean total
vegetation cover of all combined sites was
44% (control) and 42% (burn). Total vegetation
cover in burn areas was higher than or equal
to paired control areas on all sites within 2–3
years post-burn. The greatest species composition shifts occurred between shrub and perennial grass components. Shrubs were essentially eliminated in burn areas, but perennial
grass cover was 10%–35% higher compared to
paired control plots. Overall shrub cover was
21% in control areas and zero in burn areas,
while mean grass cover was 32% in burn areas
and 15% in control areas. Foliar cover results
by site and year are shown in Table 1.
EAST CACTUS.—In 1988 (1st post-fire growing season), the only differences in cover by
vegetation class between paired sites were
bare ground and shrub. In 1989 forb and
shrub cover were both greater in control than
burn areas; no other class comparisons were
significant. Results for 1991 indicate that total
vegetation cover was higher in control than
burn areas, while shrub cover was again greater
in the control area. However, herbaceous cover
was higher in burn areas. By 1992 total vegetation cover was higher in the burn area than
control. Herbaceous, forb, and grass cover were
also higher in burn areas than control. The
large increase of herbaceous cover in the burn
area from the previous year (16% to 60%) was
a response to April 1992 precipitation (9.5
cm), 6.5 cm above normal. Shrubs were still
more prevalent in control areas. Results for
1994 show continued trends from 1992. There
was no difference in total vegetation cover;
however, herbaceous and grass cover were
higher in burn areas. Litter was also more
[Volume 62
abundant in burn areas due to increased grass
production from the previous 2 years.
WEST CACTUS I.—The 1987 analysis (1st
post-burn growing season) indicated that the
burn was effective in reducing shrub cover and
that the litter component was reduced on the
burn area. There were no differences between
the other classes. In 1988 shrub cover was still
lower in the burn area; however, burn area
grass cover increased over that in the control
area. There was no difference in litter cover,
unlike the previous year. The same trend was
apparent in the 1990 results. Shrub cover was
lower in the burn area and grass cover was
higher. In 1993 shrub cover was again higher
in control areas, and both grass and total
herbaceous cover were higher in burn areas.
The greater herbaceous cover was largely due
to an increase in the grass component.
WEST CACTUS II.—In 1989 and 1990 (1st
and 2nd years post-burn), the only cover differences were in the shrub class. However, by
1995 herbaceous and litter cover were higher
in burn than control areas. Bare ground and
shrub cover were greater in the control areas.
SUCCESS.—There were no differences between cover classes in 1988 (1st year postburn). However, variability of mean values and
only 1 degree of freedom probably reduced
the power of the statistical analyses. A visual
comparison of paired means shows that total
vegetation, herbaceous, grass, and litter cover
means are substantially higher in burn areas
while shrub cover is higher in the control area
(Table 1). In 1993 variability was lower (4 df)
and statistical analyses show differences in all
classes except forbs.
IRON SPRINGS BENCH.—In the 1st post-burn
year (1996), analyses indicated differences in all
cover classes. Total vegetation, shrub, and litter
cover were greater in the control area. All other
classes were higher in the burn area. In 1997
herbaceous, grass, forb cover, and bare ground
were higher in the burn area. Shrub cover was
greater in the control area.
DRY WOMAN.—In 1984, 1987, and 1990,
only 1 transect was sampled in each paired area,
making it impossible to perform a statistical
analysis since the lack of replication precluded
calculating a measure of variance. Actual transect values of each paired area are listed in
Table 1. The burn treatment was successful in
reducing shrub cover, and trends for other
cover classes were similar to the other 5 sites.
2002]
In 1995 we sampled 5 transects in each paired
site, making analyses possible. Results indicated
that total vegetation, herbaceous, grass, and forb
cover were higher in the burn area. Shrub cover
was higher in the control area.
Diversity
Mean numbers of species on combined
control and burn areas were 17 and 18, respectively. Statistical comparisons (t tests) of species
numbers (richness) are displayed in Table 2.
Dry Woman 1995 was the only site where a
difference in species number occurred (P <
0.001) between the control (15) and burn (9)
area.
A summary of percent species similarity
(percent of species shared) between paired
areas by site and year is shown in Table 3.
Actual shared species fluctuated over time at
each site. East Cactus similarity ranged from a
high of 67% in 1988 to a low of 43% in 1992;
West Cactus I ranged from 40% in 1990 to 59%
in 1987; and West Cactus II ranged from 59%
similarity in 1989 to 78% in 1995. Success 1988
and 1993 were 50% and 47% similar, respectively. Iron Springs 1996 was 55% similar, while
Iron Springs 1997 was 71% similar. The Dry
Woman site ranged from 22% similarity in 1984
to 57% in 1990. Combining all years on each of
the 6 sites, East Cactus was 62% similar; West
Cactus I, 44%; West Cactus II, 72%; Success,
44%; Iron Springs, 75%; and Dry Woman, 62%.
419
Differences in Shannon-Weiner index values
(P < 0.05) occurred in 6 of 20 paired analyses
(Table 4: West Cactus I, 1997; Success 1993;
Iron Springs 1996 and 1997; Dry Woman 1984
and 1995). A Bray-Curtis ordination of these 6
sites is displayed in Figure 1. Even though
these 6 sites have different diversity index values between paired burn and control areas,
they do ordinate along shrub (x)- and grass (y)dominated axes. In all but a single instance
(Dry Woman 1995), control sites ordinate to
the right of each paired burn site, indicating
similar community composition. Dry Woman
1995 again was the only pair exhibiting different species richness values (Table 2).
Further investigation of the raw data indicated that community differences fell into 2
general categories. In 5 instances there was a
major component of annual grass species, cheatgrass (Bromus tectorum L.) and/or six weeks
fescue (Vulpia octoflora [Walt.] Rydb.) measured
in the burn areas at the time of sampling (Dry
Woman 1984, Dry Woman 1995, West Cactus I,
1987, and Iron Springs 1996, 1997). The Dry
Woman site has a continuing history of heavy
spring/summer grazing by domestic animals;
the Iron Springs Bench site also had a spring/
summer grazing history prior to removal in
1985. These grazing histories may have been
partially responsible for the initial invasion and
could have exacerbated proliferation of the
annual brome. The other site, Success 1993,
TABLE 2. Species richness (mean number of species) and standard errors of paired sites for all areas and sample years.
Site
Control
Burn
df
P-value
West Cactus I, 1987
West Cactus I, 1988
West Cactus I, 1990
West Cactus I, 1993
East Cactus, 1988
East Cactus, 1989
East Cactus, 1991
East Cactus, 1992
East Cactus, 1994
Iron Springs, 1996
Iron Springs, 1997
Success, 1988
Success, 1993
Dry Woman, 1984
Dry Woman, 1987
Dry Woman, 1990
Dry Woman, 1995
West Cactus II, 1989
12 (±1)
13 (±0)
18 (±1)
12 (±7)
11 (±0)
9 (±1)
15 (±0)
10 (±1)
9 (±1)
10 (±1)
16 (±1)
8 (±1)
9 (±1)
18
16
24
15 (±1)
13 (±1)
23 (±2)
19 (±1)
15 (±0)
16 (±1)
16 (±1)
8 (±0)
10 (±1)
8 (±1)
16 (±1)
13 (±1)
9 (±0)
11 (±1)
15 (±1)
7 (±1)
7 (±1)
12
15
23
9 (±1)
12 (±0)
23 (±2)
18 (±1)
1
1
1
4
1
1
1
2
2
3
3
1
3
n/a
n/a
n/a
4
1
1
4
0.09
0.126
0.295
0.018
0.5
0.705
0.05
0.015
0.184
0.297
0.613
0.5
0.103
n/a
n/a
n/a
0.001a
0.5
0.5
0.587
aSignificant at α = 0.05.
420
[Volume 62
TABLE 3. Summary of percent plant species similarity ([# shared species / # total species] × 100) between burn and
control areas by site and sample year.
Site
West Cactus I, 1987
West Cactus I, 1988
West Cactus I, 1990
West Cactus I, 1993
East Cactus, 1988
East Cactus, 1989
East Cactus, 1991
East Cactus, 1992
East Cactus, 1994
Iron Springs, 1996
Iron Springs, 1997
Success, 1988
Success, 1993
Dry Woman, 1984
Dry Woman, 1987
Dry Woman, 1990
Dry Woman, 1995
Total species
Species shared
Percent similarity
17
22
25
19
15
14
20
21
19
18
24
16
19
18
20
23
23
17
33
36
10
10
10
11
10
9
12
9
9
10
17
8
9
4
10
13
12
10
20
28
59
45
40
58
67
64
60
43
47
56
71
50
47
22
50
57
52
59
61
78
TABLE 4. Shannon-Weiner diversity index (H′) and t-test summary between all paired sites and sample years.
Site
West Cactus I, 1987
West Cactus I, 1988
West Cactus I, 1990
West Cactus I, 1993
East Cactus, 1988
East Cactus, 1989
East Cactus, 1991
East Cactus, 1992
East Cactus, 1994
Iron Springs, 1996
Iron Springs, 1997
Success, 1988
Success, 1993
Dry Woman, 1984
Dry Woman, 1987
Dry Woman, 1990
Dry Woman, 1995
Control H′
Burn H′
df
P-value
0.72
0.52
0.66
0.63
0.68
0.55
0.05
0.71
0.55
0.41
0.95
0.52
0.63
0.94
0.90
0.88
0.81
0.55
0.90
0.89
0.94
0.72
0.72
0.59
0.60
0.36
0.44
0.77
0.71
0.85
0.76
0.52
0.38
0.71
0.77
0.86
0.58
0.39
0.83
0.90
80
49
54
42
28
27
23
55
48
37
65
33
40
45
70
43
64
30
39
48
0.008a
0.09
0.579
0.669
0.439
0.084
0.667
0.408
0.098
<0.001a
0.024a
0.988
0.033a
0.01a
0.06
0.841
0.012a
0.205
0.384
0.899
aSignificant at α = 0.05.
exhibited a very large difference in the amount
of cover from needle-and-thread grass (Stipa
comata Trin. & Rupr.) in the burn area.
DISCUSSION
Prescribed burning caused a shift from late
successional, sagebrush-dominated communities
to earlier successional, grassland communities.
This occurred without reducing total vegetation cover, while improving forage quantity for
large grass-preference ungulates such as elk
(Cervus elaphus) that occur in the monument.
Intra-community (α-scale) diversity has been
maintained. Overall species richness in burn
and control plots was not different, and only a
single instance (Dry Woman 1995) had different
species richness. However, the proportionality
2002]
421
Fig. 1. Bray-Curtis ordination of 6 paired sites with significant Shannon index values (B–burn, C–control; IS–Iron Springs,
Suc–Success, DW–Dry Woman, WC1–West Cactus I).
of vegetation class has changed from shrub- to
perennial grass–dominated communities. Percent similarity of species richness between
paired areas is somewhat variable by year, but
expected. Similarity should fall within a natural
range of variation since most plant communities are dynamically affected by amount and
timing of precipitation events that differentially favor selected species (Sharp et al. 1990).
Consequently, a species presence can shift to
absence within and between burn and control
areas across years. Differences in ShannonWeiner index values where species numbers
are greater in the burn areas than control are
exceptions to the general trend. A few species
difference may not indicate a difference at all,
but may be an artifact of observer bias, different cover assessment methods, and species
area curve influences.
The National Park Service is intervening with
fire before these sagebrush/grass communities
cross a threshold into a very late, decadent,
successional stage with characteristic depauperate understories (Laycock 1991, West 1999).
Otherwise, the immediate post-burn response
of the herbaceous component would not be
observed. Landscape diversity has also been
improved by creating an alternating, patchy
sequence of communities rather than the original, continuous sagebrush matrix. National Park
Service goals of maintaining fire as an active
ecological process and fostering diversity while
protecting soil and vegetation resources have
been achieved, and implementation of periodic
prescribed burns in these shrub/grassland
communities should be continued. Without
this intervention in the successional process,
these sagebrush/grassland communities will
eventually cross a threshold into a stable state
dominated by shrubs to the near exclusion of
understory vegetation. If wildfire or prescribed
fire is introduced into this community stage,
additional inputs such as reseeding along with
other cultural practices will be necessary. The
Park Service should continue this aggressive,
proactive management approach.
LITERATURE CITED
ARCHER, S. 1989. Have southern Texas savannas been converted to woodlands in recent history? American
DAUBENMIRE, R. 1959. A canopy-coverage method of vegetational analysis. Northwest Science 33:43–64.
GAUCH, H.G., JR. 1982. Multivariate analysis in community
ecology. Cambridge Press, New York. 298 pp.
JOHNSON, K.H., R.A. OLSON, AND T.D. WHITSON. 1996.
Composition and diversity of plant and small mammal communities in Tebuthiuron-treated big sagebrush (Artemisia tridentata). Weed Technology 10:
404–416.
LAYCOCK, W.A. 1991. Stable states and thresholds of range
condition on North American rangelands: a viewpoint. Journal of Range Management 44:427–433.
OLSON, R.A., J. HANSEN, T. WHITSON, AND K.H. JOHNSON.
1994. Tebuthiuron to enhance rangeland diversity.
Rangelands 16:197–201.
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SHANNON, C.E., AND W. WEAVER. 1949. The mathematical
theory of communication. University of Illinois Press,
Urbana.
SHARP, L.A., K. SANDERS, AND N. RIMBEY. 1990. Forty years
of change in a shadscale stand in Idaho. Rangelands
12:313–328.
WEST, N.E. 1999. Synecology and disturbance regimes of
sagebrush steppe ecosystems. Pages 15–26 in P.G.
Entwhistle et al., compilers, Proceedings: Sagebrush
Steppe Ecosystem Symposium. Bureau of Land
Management, Boise, ID. Publication BLM/ID/PT001001-1150.
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WHITSON, T.D., AND H.P. ALLEY. 1984. Tebuthiuron effects
on Artemisia ssp. and associated grasses. Weed Science 32:180–184.
WINKWORTH, R.E., AND D.W. GOODALL. 1962. A crosswire
sighting tube for point quadrat analysis. Ecology 48:
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Western North American Naturalist 62(4), ©2002, pp. 423–436
MORPHOLOGICAL AND GENETIC VARIATION AMONG POPULATIONS
OF THE RARE KACHINA DAISY (ERIGERON KACHINENSIS)
FROM SOUTHEASTERN UTAH
Loreen Allphin1 and Michael D. Windham2
ABSTRACT.—Erigeron kachinensis, the Kachina daisy, is a rare species restricted to canyons in southeastern Utah.
The species is known to exhibit low fecundity due to low percent fertilization of ovules and high percent abortion of fertilized ovules. Previous reproductive studies suggest that low fecundity is a consequence of small population size and
inbreeding depression. This study examines genetic diversity within and among populations of E. kachinensis in Natural
Bridges National Monument using enzyme electrophoresis. Field populations are found to have significantly different
morphologies. However, morphological differences were less pronounced among populations grown in the greenhouse.
The Kachina daisy exhibits levels of genetic variability in its populations similar to that of other outcrossed species.
Genetic diversity statistics demonstrate that only 22.8% of genetic variation is distributed among populations. Genetic
distance could not be correlated with geographic distance. Most of the populations showed significant deviation of fixation indices from zero for multiple loci. Observation of genotype frequencies demonstrates that populations are fixing on
different genotypes and may be experiencing initial stages of genetic drift. Mean observed heterozygosity was 0.166 and
was found to increase with increasing size and/or age in populations.
Key words: Erigeron, heterozygosity, genetic variability, gene flow, fleabane.
Ecological factors and life history traits may
affect the amount and distribution of genetic
diversity within and among natural populations
(Hamrick et al. 1979, Nevo et al.1984, Hamrick
and Godt 1990). The pool of genetic diversity
in wild populations is critical for a species’
survival if environmental pressures exceed its
limits of developmental plasticity (Frankel 1983).
Moreover, genetic variability is associated with
increased fitness in populations of many plant
and animal species (Hamrick et al. 1979, Wills
1981, Danzmann et al. 1986, Ledig 1986). For
example, increased levels of heterozygosity have
been shown to increase longevity in long-lived
perennial plant species (Schaal and Levin 1976,
Hamrick et al. 1979). In addition, greater levels of genetic variation may buffer genotypes
against environmental challenges. Conversely,
the loss of genetic variability might render a
population more vulnerable to extinction in
cases of habitat perturbation, reproductive
bottlenecks, etc. (Wright 1933, O’Brien et al.
1985, Lacy 1987, Simberloff 1988, Barrett and
Kohn 1991).
According to Wright’s (1943, 1946) isolationby-distance models, mating is dependent on the
distance between individuals and their ability
to disperse propagules. Based on these models, inbreeding may occur if isolation increases
and prevents gene flow between small patch
populations. Rare and/or endangered plant
species often occur in small, disjunct populations with reduced genetic diversity due to
increased habitat fragmentation and inbreeding (Stebbins 1980, Simberloff 1988, Barrett
and Kohn 1991, Godt et al. 1996, Sun 1996).
Erigeron kachinensis Welsh and Moore
(Asteraceae), the Kachina daisy, is a rare endemic of the Colorado Plateau regions of southeastern Utah and Colorado. The species exists
in alcove seeps that arise along walls of deep
canyons in the Cedar Mesa Sandstone. As water
percolates through sandstone, it will eventually reach an impermeable layer and exude out
of the canyon wall forming seeps. Plants adhere
to the surface of the rock in such seeps, forming what are known as hanging gardens. The
Kachina daisy is limited in distribution to
these hanging gardens, which are often rare or
sporadic in distribution (May 1995). Because
this species is restricted to seeps and alcoves
that are small in size, typically along a single
seepline within a canyon, there is concern for
its preservation.
1Department of Integrative Biology, Brigham Young University, Provo, UT 84602.
2Garrett Herbarium, Utah Museum of Natural History, University of Utah, Salt Lake City, UT 84112.
423
424
Several factors contribute to rarity within
E. kachinensis and elucidate its need for
preservation. The Kachina daisy is an obligately outcrossing species with a homomorphic,
sporophytic type self-incompatibility system
(Allphin et al. 2002). The primary pollinators
of the species are small generalist flies. The
species exhibits low reproductive success due
to low levels of successful fertilization events
and a high percentage abortion of developing
fruits (Allphin and Harper 1997). Its low reproductive success has been attributed at least
in part to genetic causes, specifically inbreeding and small population size (Allphin et al.
2002). Reproductive success in the species
appears to be limited by both the species’
inability to find compatible mates due to its
incompatibility system, and recessive lethals
exposed during sexual recombination in its
small populations (Allphin et al. 2002).
Increased tourism in the canyon country of
southeastern Utah poses another serious threat
to the species. Hanging garden plant communities in the canyons of the Colorado Plateau
are fragile (Fowler et al. 1995, May 1995). Shady
alcoves that are habitat for the Kachina daisy
are also favorite areas for hikers because they
provide refuge from the hot summer sun. Many
of these alcoves contain ancient Anasazi Indian
ruins that also attract tourists. Populations of the
Kachina daisy are also potentially threatened
by flash floods, alcove roof rock spall, and other
natural catastrophes. For example, if drought
caused seeplines occupied by this species to
dry up, many populations would be eliminated.
The Kachina daisy was proposed as “endangered” by the U.S. Fish and Wildlife Service
(USFWS) in 1976 (U.S. Department of Interior 1975, 1976). Later proposals downgraded
the original recommendation to “threatened”
status (U.S. Department of Interior 1988). The
Kachina daisy was more recently listed by the
USFWS as a category 2 species, a species for
which more information is needed before assigning final designation. However, all category 2
species are now officially deleted from federal
lists (U.S. Department of Interior 1996). The
Bureau of Land Management in Utah currently
lists the Kachina daisy as a sensitive species.
Although the species was originally known
only from hanging garden communities in Natural Bridges National Monument, San Juan
County, Utah, 3 potential races or clusters of
populations have been recognized for this
[Volume 62
species: hanging garden, high elevation, and
Colorado (Welsh et al. 1993, Allphin and Harper
1994). New collections of the species at high
elevations and Colorado have significantly increased the number of known populations of
the Kachina daisy and have raised the question of its necessity for protection. However, a
previous genetic evaluation of the group suggests that the high-elevation and Colorado
races likely represent different taxa from E.
kachinensis and thus will not be discussed further in this manuscript (Allphin et al. 1996).
Nevertheless, the typical Kachina daisy, restricted to seeps and alcoves, is extremely rare
and known only from a small number of populations and drainages in the Colorado Plateau
region of Utah (Allphin 1992).
The maintenance of genetic diversity in populations of plant species such as the Kachina
daisy is an important aspect of many conservation programs (Marshall and Brown 1975,
Frankel and Soulè 1981, Dole and Sun 1992,
Rieseberg and Swensen 1996). Since the Kachina daisy occurs in small, isolated populations exhibiting reduced fitness likely due to
inbreeding (Allphin et al. 2002), it is important
to understand how genetic diversity is distributed within and among its natural populations.
Therefore, the research presented in this
manuscript will examine the amount and distribution of genetic diversity within and among
populations of E. kachinensis and assess the
amount of gene flow among its relatively isolated populations. Significant morphological
and pathogenic differences have been observed
among populations of this species (Allphin and
Harper 1997). Therefore, we will also determine
if genetic differentiation among populations,
rather than phenotypic plasticity across varying environments, might account for observed
morphological and pathogenic differences.
MATERIALS AND METHODS
Study Site
Natural populations of E. kachinensis sampled for this study occur in 6 alcove seeps
located in Natural Bridges National Monument,
San Juan County, Utah (Allphin and Harper
1994, 1997). The alcoves all occur at the same
elevation (1768 m), traverse Cedar Mesa Sandstone, and are shaded for much of the day by
high canyon walls. However, the study alcoves
differ with respect to the amount and timing of
GENETIC VARIATION IN THE KACHINA DAISY
2002]
425
TABLE 1. Population names, abbreviations, and sizes; various population characteristics; and number of individuals
sampled by population and size-class for E. kachinensis.
Population
North-facing A
North-facing B
South-facing A
South-facing B
West-facing A
West-facing B
Population
number
Abbreviation
1
2
3
4
5
6
NFA
NFB
SFA
SFB
WFA
WFB
aData taken from Allphin and Harper (1994, 1997).
bRepresents the number of individuals sampled in natural
Aspect(°)a
Densitya
(ind./m2)
Estimated
population
size
Number of
individuals
sampledb
Number of
individuals
sampled/size-classc
302
328
160
197
252
225
15.1
12.7
6.1
15.7
7.9
27.4
200–300
100–200
50–100
50–100
100–150
50–100
25
25
25
25
25
25
5–8
5–8
5–8
5–7
5–7
5–7
populations and used to compute diversity statistics. This number does not include the seedlings that
were sampled for life stage comparisons.
cRepresents the range in number of individuals sampled per size-class (1–4) in natural populations.
direct sunlight received and soil moisture. Two
of the alcoves face north, 2 face south, and 2
face west (Table 1). Each alcove, within the
pairs that share the same aspect, differs with
respect to the amount of soil moisture, ranging
from 5.8% to 25.6% (Allphin and Harper 1994,
1997; Table 1, 9).
Morphological and
Pathogenic Comparisons
To determine if observed morphological
differences among populations are due to
genetic differences or environmental plasticity, we assessed morphological differences
across 6 study populations from both field- and
greenhouse-grown individuals. For assessment
of morphological differences among the 6 field
populations, we randomly selected 100 previously tagged individuals from each of 6 study
alcoves (Allphin and Harper 1994, 1997). The
following morphological measurements were
taken from previous field studies for comparison across the alcoves: leaf length (average of
3 longest leaves), clump diameter, and number
of flower heads (characteristics which showed
significant differences among populations; Allphin and Harper 1994, 1997). Statistical comparisons among populations were performed
using a 1-way analysis of variance and Tukey
multiple means comparison (Computing Resource Center 1992).
The same morphological characteristics, in
addition to leaf width and head diameter (these
characteristics were not measured in field
study but were observed to vary in the greenhouse), were also measured across greenhousegrown progeny from each of the 6 study populations. To facilitate greenhouse comparisons,
we randomly collected seed from all 6 natural
populations, germinated it in the greenhouse,
and allowed it to grow to maturity. Note that
all greenhouse individuals received optimal
resources during the experimental period.
Greenhouse-grown individuals allowed for
morphological comparisons under a commongarden situation. Because morphological differences in field populations may be attributed
to abiotic variability in habitat, greenhouse
comparisons allowed us to determine whether
or not observed morphological differences in
field populations were environmentally or
genetically controlled. Statistical comparisons
among populations were performed for these
morphological characteristics using 1-way analysis of variance and Tukey multiple means
comparison (SAS Institute, Inc. 1994).
The incidence of pathogens, fungal infections, and herbivory was recorded for all 100
individuals in each of the 6 study populations
in an earlier study of E. kachinensis (Allphin
and Harper 1997). The data compared the percent of infected individuals in each population
across the 6 study alcoves. For this study we
reexamine these data for evidence of differential resistance to these stressors among study
populations and determine if differential resistance is related to genetic differentiation among
populations.
Enzyme Electrophoresis
The level of genetic diversity was assessed
within and among the 6 study populations
from Natural Bridges National Monument using enzyme electrophoresis (Soltis et al. 1983,
Odrzykoski and Gottlieb 1984, Allphin et al.
1998). Approximately 4–5 young leaves were
426
collected from 25 individuals in each population (Table 1). The approximate proportion of
the total population size that this sample represents can be estimated from Table 1. Samples
from each population consisted of 6 or 7 individuals in each of 4 size-classes based upon
number of rosettes: (1) a single rosette, (2) two
rosettes, (3) three rosettes, and (4) ≥ four rosettes per plant (Table 1). Seedling leaf samples
were obtained from (8–10) seedlings grown
from germinated seed that had been randomly
collected from each study population and sown
in the greenhouse. Because the number of
rosettes in these populations has been shown
to increase with age in this species (Allphin
and Harper 1997), the separation of samples
with respect to size allowed us to indirectly
evaluate whether genetic variation, heterozygosity, was correlated with increased age in
this species (Schaal and Levin 1976, Hamrick
et al. 1979).
All leaf tissue samples were ground in a
phosphate-PVP grinding buffer (Soltis et al.
1983) using a mortar and pestle. Ground material was absorbed into wicks made of Whatman 3 MM filter paper and stored in an ultra
cold freezer (–70°) prior to electrophoresis.
Electrophoresis was performed using a variety
of gel and electrode conditions (Soltis et al.
1983, Odrzykoski and Gottlieb 1984; Table 2).
Samples were analyzed using 12% starch gels
that were sliced and stained for allozyme
markers following standard protocols (Soltis et
al. 1983). Twenty enzymes were surveyed for
variability. Allozyme markers from 12 polymorphic enzyme loci that provided consistent,
interpretable results were analyzed (Table 2).
The presence or absence of each allele detected
was recorded for all populations.
Genetic data collected from the 12 variable
loci were used to compute a variety of genetic
diversity statistics for each study population
following Hamrick et al. (1979), Hedrick
(1985), and Hamrick and Godt (1990) using
LYNSPROG, a genetic statistics program written by M.D. Loveless (College of Wooster,
Wooster, OH). These genetic diversity statistics include mean observed heterozygosity
(Ho, a direct estimate), expected heterozygosity (He, based on Hardy-Weinberg equilibrium model), polymorphic index (P, mean proportion of polymorphic loci), number of alleles
per locus (A), mean number of effective alleles
per locus (Ae, number of equally frequent alle-
[Volume 62
les in an ideal population that would produce
the same homozygosity as the actual population; Hartl 1988), and Nei’s genetic distance
(D; Nei 1972). Correlation was performed
in the form of linear regression to determine
whether there was a significant positive relationship between geographic distance and
genetic distance (SAS Institute, Inc. 1994).
Mean observed heterozygosity (Ho) was also
computed by size-class (life stage) for the
Kachina daisy. Because of the small sample
sizes for each life stage in each population,
populations were pooled. Therefore, mean Ho
values for this analysis represent a mean across
all individuals of a particular life stage across
all populations. Significant differences in these
values were assessed using a 1-way analysis of
variance (Computing Resource Center 1992).
A fixation index (F) was computed for each
locus and population using methods of Wright
(1965). Chi-square analysis of Wright’s fixation
indices for study populations was used to test
significance of deviations from Hardy-Weinberg (H-W) expectations (0). Fixation indices
and deviations from H-W expectations were
computed following Hamrick et al. (1979),
Hedrick (1985), and Hamrick and Godt (1990)
using LYNSPROG.
Distribution of genetic variation among
populations was estimated using Nei’s (1973)
genetic diversity statistics employing LYNSPROG. These statistics include total genetic
diversity (HT), intrapopulation genetic diversity (HS), diversity among populations (DST),
the proportion of genetic variation distributed
among populations (GST), and inter- to intrapopulation gene diversity (RST). The average
number of diploid migrants exchanged among
local populations per generation, Nm (Wright
1951, Slatkin and Barton 1989), was also estimated (Nm = (1/GST – 1)).
Ruzicka’s (1958) index of similarity was
computed for all pairwise comparisons of populations based on allele frequency data for all
loci. Average linkage clustering was performed
on this similarity matrix using the unweighted
pair-group method (UPGMA; Krebs 1989).
Cluster analysis permitted the construction of
a dendrogram, or cluster diagram, illustrating
genetic similarities among populations. This was
compared to a 2nd dendrogram based upon various abiotic/site characteristics. In order to generate the 2nd dendrogram, a similarity matrix
was computed based upon Ruzicka’s index of
2002]
427
TABLE 2. Enzymes used in population genetic analyses of E. kachinensis.
Enzyme
Esterase
Isocitrate dehydrogenase
Leucine aminopeptidase
Malate dehygrogenase
NADH-diaphorase
Phosphoglucoisomerase
Phosphoglucomutase
6-Phosphogluconate dehydrogenase
Shikimate dehydrogenase
Triosephosphate dehydrogenase
TOTAL NUMBER OF LOCI
Acronym
Gel/electrode
buffersa
No. loci
scored
EST
IDH
LAP
MDH
NADH
PGI
PGM
6-PGD
SKDH
TPI
8
11 & M
8
M
8
6&8
6
11 & M
11 & M
6&8
1
1
1
1
1
2
2
1
1
1
12
aSystems 6, 8, 11 after Soltis et al. (1983); system M a 7.5 pH version of the morpholine citrate system after Odrzykoski and Gottlieb (1984).
similarity (Ruzicka 1958) for various abiotic/
site characteristics for each population of E.
kachinensis as reported in an earlier study (Allphin and Harper 1994). Average linkage clustering using UPGMA was also performed on this
similarity matrix (Krebs 1989) and presented
in the form of a dendrogram to demonstrate
habitat similarities among study populations.
RESULTS
Significant differences for morphological
characters among populations from field and
greenhouse individuals are given in Table 3.
Most morphological differentiation among populations is apparently due to environmental
factors. From the field data, alcoves 2 and 5
had the smallest leaves and smallest clump
diameters (Table 3; Allphin and Harper 1997).
Alcoves 4 and 6 exhibited the largest leaves
and clump diameters (Table 3; Allphin and
Harper 1997). These data are consistent with
environmental data. Alcoves 2 and 5 are the
driest sites and alcoves 4 and 6 the wettest
(Allphin and Harper 1994; Tables 1, 9). Greater
morphological differences were observed for
field populations than among cultivated individuals (Table 3). Actual morphological differences due to genetic factors were more rare,
yet more pronounced, for greenhouse-grown
individuals. Population (alcove) 1 exhibits significantly smaller leaves, leaf widths, clump diameters, and head diameters than other populations grown in common-garden conditions
(Table 3).
Levels of genetic variability for allozyme
loci are summarized in Table 4. Mean observed
heterozygosity varies among the 6 populations
(alcoves). Populations 4 (south-facing B) and 6
(west-facing B) exhibited the highest mean
observed heterozygosity and polymorphic indices (Ho = 0.201, 0.195, P = 0.187, 0.231;
Table 3). Conversely, population 5 showed the
lowest level of genetic variability and lowest
polymorphic index (Ho = 0 .120, P = 0.155).
Mean heterozygosity across all populations was
moderate (Ho = 0.166; Table 4). Expected
heterozygosity was different from observed
heterozygosity in most populations (Table 4).
Increased number of rosettes for individuals in populations of the Kachina daisy is consistent with increasing age in field populations
(Allphin and Harper 1997). Mean observed
heterozygosity (computed as a mean value across
all loci for individuals in the same age/size
class in the population) increases with increasing size-class (based on number of rosettes) in
the Kachina daisy (Fig. 1). Seedlings (from year
sampled) exhibit relatively high diversity (Ho
= 0.266). Diversity is significantly lower for
size-class 1 than the seedling class and continues to increase across size-classes. The youngest size-class, SC1, is significantly less heterozygous than size-classes 3 and 4 at P ≤ 0.05.
Genetic diversity statistics for all populations
are summarized in Table 5. Total gene diversity
is fairly high (Ht = 0.311). Most of this diversity is distributed within populations (Hs =
0.249). GST was estimated as 0.228, and thus
22.8% of genetic variation is distributed among
populations (Table 5). This results in a relatively high estimate of Nm, the number of
migrants exchanged between local populations
per generation. Gene flow (Nm) was estimated
to be 0.847. This is interpreted as an exchange of
between 8 and 9 migrants every 10 generations
428
[Volume 62
TABLE 3. Mean measurements for several morphological characteristics of 6 study populations of the Kachina daisy in
Natural Bridges National Monument. Cultivated populations were sown from seed of the 6 field populations and grown
under greenhouse conditions at the University of Utah. Means for any population which do not share the same letter are
significantly different at P ≤ 0.05. Columns with lines indicate that data are unavailable. Morphological data from field
populations were taken from Allphin and Harper (1997).
Population
(alcove)
Leaf length
(cm)
Leaf width
(cm)
Clump diam.
(cm)
No. flower
heads
Head diam.
(cm)
2.7 b
(0.07)
1.9 a
(0.06)
2.5 b
(0.06)
3.6 c
(0.08)
2.1 a
(0.06)
3.4 c
(0.08)
—
4.8 b
(0.13)
3.6 a
(0.10)
6.2 c
(0.18)
7.3 d
(0.18)
4.4 b
(0.1 0)
6.6 c
(0.18)
1.3 a
(0.08)
0.9 b
(0.10)
1.9 c
(0.19)
2.3 c
(0.20)
0.6 b
—
1.4 a
(0.14)
—
6.9 a
(2.33)
18.0 b
(5.66)
16.9 b
(2.79)
13.6 bc
(8.42)
9.8 c
(6.89)
11.9 bc
(6.30)
67.9 a
(32.69)
80.0 a
(28.28)
39.0 b
(15.25)
34.0 b
(17.75)
63.62 a
(41.81)
47.0 ab
(21.49)
1.0 a
(0.18)
1.3 b
(0.21)
1.3 b
(.180)
1.1 a
(0.08)
1.3 b
(0.19)
1.4 b
(0.14)
Field
1
2
3
4
5
6
Cultivated
1
—
—
—
—
(0.12)
—
3.7 a
(1.03)
5.8 b
(1.77)
5.4 b
(1.27)
5.2 b
(1.52)
4.9 b
(0.82)
5.4 b
(1.49)
2
3
4
5
6
0.7 a
(0.19)
1.3 b
(0.15)
1.2 b
(0.25)
1.2 b
(0.32)
1.2 b
(0.36)
1.4 b
(0.43)
—
—
—
—
TABLE 4. Level of intrapopulation allozyme variation as measured across 25 individuals in each of 6 study populations
of the Kachina daisy in Natural Bridges National Monument. Values are given as means across 12 enzyme loci for each
population.
Population
(alcove)
Number of
unique
genotypesa
NFA (1)
NFB (2)
SFA (3)
SFB (4)
WFA (5)
WFB (6)
ALL POPULATIONS
1
1
1
2
2
2
Mean observed
heterozygosity
(Ho)
Mean expected
heterozygosity
(He)
Polymorphic
index
(PI)
Mean no.
alleles
(A)
Mean no.
effective alleles
(Ae)
0.150
0.177
0.154
0.201
0.120
0.195
0.166
0.157
0.179
0.181
0.187
0.155
0.230
0.182
0.157
0.179
0.181
0.187
0.155
0.231
0.182
1.58
1.58
1.67
1.42
1.67
1.83
1.63
1.29
1.30
1.32
1.34
1.26
1.47
1.33
aLines illustrate that the unique genotype(s) is (are) shared by both populations in the species pair but no other populations.
(Slatkin and Barton 1989). However, this
appears to be an overestimate of gene flow
when genetic distances are compared with geographic distance (Table 6). Genetic distance is
not significantly correlated with geographic
distance (r2 = 0.0592). Distant populations are
just as likely to have alleles in common as pop-
ulations in close proximity. Gene flow (Nm)
values lower than 1 can lead to population differentiation (Wright 1951).
Although the genetic diversity statistics demonstrate that most variability is distributed within populations in the Kachina daisy, genotypic
frequencies among populations for 2 enzyme
2002]
429
Fig. 1. Mean observed heterozygosity across seedlings and 4 other size/age classes (as described in Methods) in E.
kachinensis. Seedlings were sown from seed collected in all natural populations of E. kachinensis and grown in the
greenhouse. Mean observed heterozygosity for the seedling class is significantly higher than for size-class 1. Size-class 1
exhibits significantly lower mean observed heterozygosity than size-classes 3 and 4 at P = 0.05.
loci in particular suggest that populations
are fixing on different genotypes. Table 7 summarizes genotype frequencies for 6-PGD-2
and EST-1. At EST-1 higher genotype frequencies are observed for AA and AB genotypes for populations 1, 2, and 6, and yet populations 3 and 4 generally lack allele A. At 6PGD-2 populations 1 and 2 (north-facing populations) exhibit higher frequencies for genotypes AB and BB, while populations 3 and 4
(south-facing) are missing allele B entirely.
Moreover, 2 of the populations (3, 4) have
unique genotypes not found in other populations. Two of the population pairs (both northfacing or both west-facing) had unique genotypes not found in other sampled populations.
Furthermore, at PGI-2 population 3 (SFA) carries an allele that cannot be found in any other
population (Fig. 2, Table 4).
Fixation index (F) was determined for all
loci in each population of E. kachinensis. Chisquare analysis demonstrates significant deviation from Hardy-Weinberg expectations for
some of the populations and loci (Table 8).
West-facing B, one of the smallest and most
genetically variable of the study populations,
showed significant deviation from H-W expectations at 4 loci. All populations showed significant deviation from H-W expectations for 1 or
more loci (Table 8). Since 6 populations and 12
TABLE 5. Genetic diversity statistics from 6 populations
of E. kachinensis in Natural Bridges National Monument,
San Juan County, Utah. Data given are mean values for 12
enzyme loci.
Total gene diversity (HT)
Within populations (HS)
Among populations (DST)
Coefficient of gene differentiation
(GST = DST / HT)
Inter- to intrapopulation gene diversity (RST)
0.311
0.249
0.066
0.228
0.441
loci were sampled, there are 72 determinations for F deviations from H-W equilibrium.
For E. kachinensis, 7 of the 72 determinations
(~10%) deviated significantly (P = 0.05).
The dendrogram based on allele frequency
data demonstrates genetic similarities among
populations (Fig. 3A). Populations 1 and 2 are
the most similar (86.2%). These populations
are only approximately 100 m from each other,
and gene flow likely occurs. Populations 3 and
4 are also very similar (83.7%). These 2 populations are approximately 250 m apart; thus,
gene flow is also likely between these populations. However, population 6 (WFB) clusters
with populations 1 and 2, but it is geographically the most distant of the populations (over
2 miles). West-facing A and B (5 and 6) are
located only 500 m from one another but are
genetically dissimilar (Fig. 3A).
430
[Volume 62
TABLE 6. Geographic distance and genetic distances between sampled populations of E. kachinensis. Geographic distances (m) are given above the diagonal, and Nei’s (1972) genetic distances (D) are given below the diagonal.
Population
NF-A
1
NF-B
2
SF-A
3
SF-B
4
WF-A
5
WF-B
6
1
2
3
4
5
6
—
0.0192
0.0989
0.1122
0.1126
0.0376
100
—
0.1448
0.1446
0.1307
0.0355
1610
1519
—
0.284
0.0516
0.0376
1860
1769
250
—
0.0457
0.0553
3980
3880
2360
2110
—
0.0799
3480
3380
1860
1610
500
—
TABLE 7. Genotype frequencies at 2 of the enzyme loci (6-PGD-2 and EST) for study populations of E. kachinensis in
Natural Bridges National Monument.
Population
EST
N-facing A (1)
N-facing B (2)
S-facing A (3)
S-facing B (4)
W-facing A (5)
W-facing B (6)
6-PGD-2
N-facing A (1)
N-facing B (2)
S-facing A (3)
S-facing B (4)
W-facing A (5)
W-facing B (6)
AA
AB
BB
0.167
0.480
0
0
0
0.292
0.667
0.480
0.093
0
0.417
0.583
0.167
0.040
0.917
1.00
0.583
0.125
0.160
0.158
1.00
1.00
0.750
0.375
0.480
0.210
0
0
0.250
0.250
0.360
0.632
0
0
0
0.375
When this dendrogram is compared with
another based on abiotic/site characteristics of
the 6 study alcoves (Allphin and Harper 1994),
a different clustering pattern results (Fig. 3B).
Populations are only ~63% similar at best.
Abiotic/site characteristics differ the most among
populations that are most similar genetically
(populations 1, 2). Environmental variability has
been shown to select for allozyme markers (Mitton 1994), but these data show that allozyme
variability is independent of environment.
DISCUSSION
The results presented in this manuscript
have many evolutionary and conservation implications. Population size is often correlated with
an increase in genetic diversity (heterozygosity) in natural populations (Billington 1991,
Stangel et al. 1992). However, in the Kachina
daisy, 2 of the smallest populations (SFB and
WFB) have the highest heterozygosity and polymorphic indices. These data are consistent with
findings in other rare species (Sherwin et al.
1991, Allphin et al. 1998, Maki and Morita
1998). In addition, genetic similarities among
populations were found to be independent of
ecological (abiotic) similarities among populations.
In the Kachina daisy, most genetic variability appears to be distributed within rather than
among populations. This would not appear to
support the idea of historic, long-term population isolation and genetic drift. The distribution
of genetic diversity may be affected by spatial
and historical factors (Kimura and Maruyama
1971, Slatkin 1987, Sheely and Meagher 1996).
Because the historical range and past population sizes of the Kachina daisy are unknown,
interpretation of the data presented in this
manuscript is difficult. However, most populations have fixation indices that are significantly different from zero (no fixation) for at
least 1 or more loci. These data suggest that
most populations of the Kachina daisy do not
randomly cross. Maki (1999) found similar findings in a threatened insular endemic in the
genus Aster.
2002]
431
Fig. 2. Photograph of an allozyme showing variability at the PGI-2 locus. Notice only a single population (alcove 3)
has allele 2.
TABLE 8. Fixation index (F) values for selected enzyme loci of each population generated using LYNSPROG. Population F values exhibiting significant chi-square deviations from zero are followed by a symbol. Chi-square values for fixation index values were generated using LYNSPROG, following the methods of Hedrick (1985). **P = 0.01; * P = 0.05;
†P = 0.10.
Population
6PGD-1
MDH-2
IDH-2
NADH-1
SKDH-1
NFA (1)
NFB (2)
SFA (3)
SFB (4)
WFA (5)
WFB (6)
0.0200
0.4714*
0.0000
0.0000
–0.1190
0.5104*
–0.1190
–0.3056†
–0.2162
0.1048
–0.1463
–0.4242*
0.2833
0.0882
0.4778*
0.2656
0.1188
0.3354†
0.7873**
0.0000
–0.0387
–0.3824†
0.6543**
0.5131**
0.0000
0.0559
0.5000†
0.0143
0.0571
0.2781
High fixation indices could result from inbreeding, selection pressures, random genetic
drift, founder events, etc. We suggest that populations of the Kachina daisy do not show
signs of historic, long-term isolation and drift
because habitat fragmentation and reduction
in population size are of fairly recent origin.
However, other additional lines of evidence
suggest that populations of the Kachina daisy
might be in the early stages of genetic differentiation and genetic drift. For example, populations of the Kachina daisy appear to have
differential ability to resist pathogens and predators (Allphin and Harper 1997). In addition,
populations differentially fix alternate genotypes
at some loci (Table 7).
Habitat fragmentation might be a recent
phenomenon in the Kachina daisy. Natural
habitat fragmentation in this species occurs
when seeps and hanging gardens become more
isolated from one another, as seeplines in a
canyon begin to dry. Drying along seeplines in
Natural Bridges National Monument appears
to be a relatively recent event (Brough et al.
1987). We compared soil moisture data from
1991–92 (Allphin and Harper 1994) with soil
moisture data obtained in 1997 at all study
populations. These data demonstrate that all
seeps with alcove overhangs (NFA and NFB)
were drier in 1997 than in 1992 (Table 9). The
populations without alcove overhangs (those
populations receiving additional moisture from
local precipitation) were the only ones that did
not show a drier condition in 1997. Although
we have only 2 data points and cannot conclusively say that the alcoves are drying, observations made in these alcoves over 10 years of
monitoring suggest drying in these seeps. For
example, we have observed an increase in
rodent burrows at seeplines that were originally too wet to support habitation. Moreover,
population size has gradually dwindled in the
study populations over the past 10 years of
study (Allphin and Harper 1994, 1997).
432
[Volume 62
TABLE 9. Mean soil moisture (% of dry weight) for
1990–1992 (mid-June) and 1997 (mid-June).
Population
Percent
soil moisture
(1990-1992)a
Percent
soil moisture
(1997)
1
2
3
4
5
6
Mean
12.3
5.8
18.4
12.2
12.7
25.6
14.5
15.0
8.3
18.4
6.1
8.7
22.8
13.2
aData taken from Allphin and Harper (1994).
Fig. 3. A, Dendrogram illustrating similarities (percent)
among sampled populations of E. kachinensis based upon
allele frequencies for allozyme loci; B, dendrogram illustrating similarities (percent) among sampled populations
of E. kachinensis based upon abiotic/site characteristics
(Allphin and Harper 1994).
Since some populations still contain over 100
individuals, we may be only beginning to see
the effects of isolation and small population
size as a result of habitat fragmentation in these
populations (Table 1). Although there was a
moderately high estimate of gene flow between
populations of the Kachina daisy based upon
genetic distances, we could find no correlation
between geographic and genetic distance in
this species. This is not as expected under the
isolation-by-distance model (Wright 1943, 1946).
One might expect a lack of correlation between
geographic and genetic distance if panmixia
(or regular random mating) were occurring
among populations (Wright 1946). This explanation seems unlikely for these populations of
the Kachina daisy because of the long distances separating many of these populations,
the bi-directionality of canyon systems, and
the species’ limited dispersal capabilities, due
to its short-distance seed dispersal (primarily
gravity) and small pollinators. A lack of corre-
lation between genetic distance and geographic distance has been found for other rare
species including the family Asteraceae (Allphin et al. 1998, Matthews and Howard 1999).
Therefore, we suggest that Nm may not be
reflective of actual gene flow events in the
Kachina daisy, but possibly alleles shared in
common through common ancestry. For example, if small population sizes were due to a
recent habitat fragmentation, many alleles
would still be shared among populations and
not yet lost through genetic drift (Wright 1933,
1943, 1946).
Most morphological variation among natural
populations of the Kachina daisy is apparently
due to environmental factors. Morphological
differences among populations reflecting genetic
differences are evidenced by a few, very pronounced morphological differences between
some of the populations of greenhouse-grown
individuals (primarily alcove 1). Morphological
characteristics appear to be very plastic in this
species and not always indicative of genetic
differences.
Additional patterns emerge from the genetic
data that have biological and conservation implications. First, there is a positive relationship
between heterozygosity and longevity in our
data. If we ignore the seedling size-class, heterozygosity increases with increasing size-class
in the Kachina daisy. Increased size of individuals in populations of the Kachina daisy has
been shown to be consistent with increasing
age in field populations (Allphin and Harper
1997). It is, therefore, probable that greater
genetic variation increases the potential to
achieve larger size (number of rosettes) or age,
and is best interpreted as a correlation between
heterozygosity and increasing longevity of
2002]
individuals, as has been demonstrated for trees
and other long-lived perennials (Schaal and Levin
1976, Hamrick 1979. Hamrick et al. 1979). The
Kachina daisy is a relatively long-lived species.
Demographic monitoring (Allphin and Harper
1997 and unpublished data) has demonstrated
no mortality in large individuals over 10 years
of monitoring. This demographic research suggests that individuals may live as long as 20
years in the absence of environmental catastrophe. We suggest that the relationship between heterozygosity and longevity in the
Kachina daisy might help to explain why this
nonwoody perennial is so long-lived.
Another pattern emerges in the comparison
of heterozygosity in seedlings with other sizeclasses of the Kachina daisy. Greenhouse-grown
seedlings exhibited unusually high levels of
heterozygosity compared to the size/age classes
(1–3). This is greater genetic variability than
would have been observed in seedlings in the
natural populations, suggesting that the high
level of heterozygosity in seedlings is an artifact of being grown in a greenhouse setting.
For example, environmental constraints could
have eliminated some genotypes from natural
populations.
Additionally, there is evidence of low reproductive success due to high levels of embryo
abortion in the Kachina daisy that can be
explained by inbreeding and genetic load (Allphin and Harper 1997, Allphin et al. 2002).
Genetic load, recessive deleterious or lethal
genes present in a population, should be expected to increase in out-crossed species (Wiens
1984, Wiens et al. 1987, Charlesworth 1989a,
1989b). Such deleterious recessive genes appear
in homozygous form as chromosomes are independently assorted in meiosis and produce
lethal or sublethal combinations that result
in the abortion of developing embryos or
endosperm, or death of developing seedlings.
Because the most reproductively active sizeclasses of the Kachina daisy are the largest
size-classes (Allphin and Harper 1997), and
this obligately outcrossing species exhibits relatively high levels of genetic variability for
these size-classes, there is a greater potential
for inbreeding due to genetic load because
more recessive lethals can be maintained in
the recessive state. As the populations of the
Kachina daisy become isolated and individuals
are forced to mate with close relatives, lethals
433
may be exposed in the homozygous state,
causing the high level of embryo abortions
observed in this species (Allphin and Harper
1997). Genetic data presented in this manuscript support the reproductive studies that
suggest inbreeding (Allphin et al. 2002). In this
study all populations showed significant deviation of fixation indices from zero for 1 or more
loci (Table 8). These data suggest that populations of the Kachina daisy are not mating randomly and that inbreeding may be occurring
in at least some of the natural populations.
To determine how genetic diversity in the
Kachina daisy compares with other outcrossing, perennial endemic species, we compared
our data with a synthesis paper by Hamrick
and Godt (1990) summarizing levels of diversity across a wide sampling of plants. Mean
observed heterozygosity in E. kachinensis is
similar to that found in other outcrossing plant
species and perennial dicots (Table 10; Hamrick and Godt 1990). However, genetic diversity in E. kachinensis is higher than the level
of variation found in other narrow endemic
species (Table 10; Hamrick and Godt 1990).
Other species in Asteraceae show similar patterns (Maki and Morita 1998, Ayres and Ryan
1999). For example, Aster spathulifolius, a narrow endemic from Japan, also showed high
genetic diversity for its endemism (Maki and
Morita 1998).
Allozyme markers, as used to assess genetic
diversity in the Kachina daisy, have been considered “neutral” genes reflecting evolutionary processes affecting the entire genome (Avise
1994). Mitton (1994), however, proposed that
environmental factors may occasionally select
for allozyme markers. He considers that allozymes are, therefore, not always indicative of
random genetic change. However, our 2 dendrograms for similarity among populations of
the Kachina daisy based upon genetic and
environmental characteristics provide different topologies from one another (Fig. 3). This
suggests that environmental conditions likely
are not responsible for genetic differences
across the study alcoves (populations) and that
allozymes can be used as neutral markers in
this species.
The patterns presented in this manuscript
suggest some important conservation implications. Because populations show signs of inbreeding depression and are beginning to show
signs of population differentiation and loss of
434
[Volume 62
TABLE 10. Level of allozyme variation at species level for species in different categories. The data were extracted from
Hamrick and Godt (1990). These data are compared with allozyme variation at the species level for E. kachinensis. Standard error values are given in parentheses below the means. Statistical differences among means for species or groups
were determined using multiple means comparison (Snedecor and Cochran 1967). Means followed by the same letter in
a column are not significantly different at P = 0.05.
Categories
or species
Number
taxa
Mean no.
of pop.
Mean no.
of loci
Dicots
329
Short-lived
perennials
Outcrossing
annuals
Endemic
152
11.9
(1.7)
8.8
(1.2)
10.7
(2.1)
6.5
(0.9)
6
16.8
(0.4)
17.2
(0.7)
17.7
(0.7)
17.8
(0.6)
12
E. kachinensis
172
81
1
diversity, land managers may need to bring
pollen or seed from other populations, or genetically more distant sources, to increase reproductive capacity in local populations of the
Kachina daisy. However, we caution the movement of genes among populations if populations are beginning to become differentiated
from one another.
Alcove seeps and hanging gardens that support Kachina daisy populations are fragile habitats. Backcountry recreation is increasing in
the Colorado Plateau area of Utah. Trails into
canyon bottoms run near populations of the
Kachina daisy. One trail in Natural Bridges
National Monument comes very close to alcove
6, which despite theoretic expectations was
shown to have the highest genetic diversity
and the smallest population size. Land managers should take special care to maintain this
population. Trails may need to be rerouted to
avoid human impact to Kachina daisy populations. With growing human impact to the canyon
country, protection of all populations may prove
necessary for preservation of genetic diversity
within the Kachina daisy.
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Ae
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Western North American Naturalist 62(4),© 2002, pp. 437–450
TEMPERATURE RESPONSES AND HABITAT SHARING IN
TWO SYMPATRIC SPECIES OF OKANAGANA
(HOMOPTERA: CICADOIDEA)
Allen F. Sanborn1,2, Jessica H. Breitbarth1,3, James E. Heath1,4, and Maxine S. Heath1,4
ABSTRACT.—Okanagana striatipes and O. utahensis are species synchronous in location of activity and utilization of
host plants. They possess similar acoustic behavior. Analysis of calling songs shows that calls overlap in frequency but
differ in temporal pattern. Based on characteristics of the cicada auditory system and the species recognition mechanism, the potential for acoustic interference exists. Both species are ectothermic behavioral thermoregulators. Measurements of thermal preference and body temperature during singing show that although thermal preferences are similar,
O. utahensis sings at a significantly higher body temperature. Differences in body temperature required to coordinate
singing in the 2 species provide a partial temporal separation of acoustic signaling. We suggest the physiological mechanisms that permit synchronous utilization of a habitat by the 2 species are the production of calling songs of different
temporal patterns and the presence of different thermal requirements, which may permit and/or facilitate temporal separation of the acoustic environment during the day.
Key words: Okanagana striatipes, Okanagana utahensis, temperature, thermal adaptation, communication, song,
cicadas.
Okanagana striatipes (Haldeman) and Okanagana utahensis Davis share sagebrush fields
of the western United States. Okanagana utahensis is described as resembling O. striatipes
but is slightly larger and darker in color (Davis
1919). The 2 species are active in adult form
during June and July (Davis 1919). Both O.
striatipes (Davis 1930) and O. utahensis (Davis
1919) have been associated with sagebrush
(Artemisia spp.). Although the cicadas may sing
from other plant species, a species of sagebrush
is always present in the habitat and appears to
be the host plant for both species. Cryptic coloration makes the cicadas very difficult to see
when perched on sagebrush (Davis 1932).
Acoustic behavior of the 2 species is also
similar. Both species, for example, are solitary
animals when calling. Frequency ranges of the
calling songs appear to overlap. Okanagana
striatipes produces a calling song of medium
pitch and average duration (Beamer and Beamer
1930, Davis 1930), whereas the song of O. utahensis is a long, shrill cry (Davis 1919) or a continuous song (Davis 1921).
Most male cicadas produce acoustic signals
to attract females. Acoustic interference between
species inhabiting the same environment has
been shown in insects (Perdeck 1958, Ulagaraj
and Walker 1973, Morris and Fullard 1983,
Latimer and Broughton 1984, Bailey and Morris 1986, Greenfield 1988, Römer et al. 1989,
Schatral 1990), frogs (Schwartz and Wells 1983,
Schwartz 1993), and birds (Cody and Brown
1969, Ficken et al. 1974, Popp et al. 1985).
Cicada calls also have been shown to cause
acoustic interference in frogs (Paez et al. 1993).
It has been suggested that temporal separation
(Wolda 1993, Gogala and Riede 1995, Riede
1995, Riede 1997) or frequency separation
(Gogala and Riede 1995, Riede 1996) occurs in
cicada communities to decrease acoustic interference.
The important song parameter in cicada
long-distance communication has been shown
to be call frequency (Doolan and Young 1989)
or call intensity (Daws et al. 1997). Because
the songs of O. utahensis and O. striatipes
appear to overlap in frequency and the calls
are of similar intensity (Sanborn and Phillips
1995), the potential for acoustic interference
exists between these cicadas sharing the same
habitat.
1University of Illinois, Department of Molecular and Integrative Physiology, 524 Burrill Hall, 407 South Goodwin Avenue, Urbana, IL 61801.
2Present address: Barry University, School of Natural and Health Sciences, 11300 NE Second Avenue, Miami Shores, FL 33161.
3Present address: 23769 Monte Carlo Place NW, Paulsbo, WA 98370.
4Present address: 104 Hummingbird Circle, Buchanan Dam, TX 78609.
437
438
Although the songs of O. striatipes and O.
utahensis appear to differ in their temporal
patterns, the potential for acoustic interference between the species still exists due to
characteristics of the cicada auditory system
and the species recognition process. The auditory system of cicadas usually shows a peak
sensitivity at the frequency of the species calling song but is sensitive to a wide range of frequencies (Katsuki and Suga 1958, 1960, Hagiwara and Ogura 1960, Katsuki 1960, Enger et
al. 1969, Popov 1969, 1981, Simmons et al.
1971, Young and Hill 1977, Schildberger et al.
1986, Huber et al. 1990; but see Popov et al.
1985, Popov and Sergeeva 1987, Fonseca 1993
for exceptions). In fact, Huber et al. (1990)
showed that the auditory system in Magicicada
cassinii (Fisher) is more sensitive to the call of
M. septendecim (L.) than the auditory system
of M. septendecim. Physical properties of the
sound-production system prevent O. striatipes
and O. utahensis from altering the frequency
of their calling songs to prevent acoustic interference. Since one species is probably capable
of hearing the other quite well, temporal patterns of the song must act to separate the
species.
Popov and Shuvalov (1974) described cicada
auditory receptors as a specialized system in
the analysis of amplitude-modulation patterns.
However, these receptors respond to a wide
range of temporal patterns; they are not an
integral part of the conspecific signal recognizer (Huber 1983). Pringle (1954, 1956) suggested the frequency of a cicada song acts as
an information carrier, while the speciesspecific information is carried in temporal patterns of the song (Hagiwara and Ogura 1960,
Moore 1961, Frings and Frings 1977, Huber
1984, Joermann and Schneider 1987). Since
auditory receptors respond to a wide variety of
signals, auditory neural pathways must be responsible for filtering out species-specific calls.
The primary response to a conspecific song
is based on the spectral content of the song
(Huber et al. 1979). Nerve fibers respond to
natural calling and courtship sounds with a
specificity dependent on carrier frequency,
rhythm, and transient content of the presented sound (Huber et al. 1980). Cicada auditory nerves respond synchronously to the temporal pattern of a conspecific song while the
response to allospecific calls is not clearly
[Volume 62
related to song activity (Pringle 1954, Katsuki
and Suga 1960, Schildberger et al. 1986, Huber
et al. 1980, 1990). Cicada auditory receptors
are sensitive to intensity changes (Hagiwara
and Ogura 1960, Katsuki 1960, Katsuki and Suga
1960) and are especially sensitive to transient
stimuli found in calling songs (Huber et al.
1979, 1980). Amplitude modulations within the
call elicit groups of spikes in the auditory nerve
(Huber et al. 1980). Interneurons are responsible for filtering the auditory input to the brain,
and apparently these interneurons react only
to conspecific calls (Huber et al. 1980, Huber
1984).
Okanagana striatipes and O. utahensis are
synchronous in time of activity during the
year, location of activity, utilization of host
plants, and possession of similar acoustic behavior. These similarities expose the 2 species to
interspecific competition for physical and
acoustic resources within their environment.
We try to determine with this study whether
there are differences in acoustic signals produced by the species and the possible role of
thermal requirements for singing in the 2
species that may act as physiological mechanisms to permit synchronous sympatry.
MATERIALS AND METHODS
Animals
The species Okanagana striatipes and O.
utahensis were studied in Cortez, Montezuma
County, Colorado, USA. Animals were randomly sampled for data collection in the field and
for specimen collection for laboratory experimentation. Experiments were performed in
early July, approximately in the middle of the
emergence period for each species, during
1982, 1983, 1984, 1986, 1988, and 1989. The
species were active in an almost pure flat of
Great Basin sagebrush (Artemisia tridentata).
We placed the animals captured for laboratory
experimentation in a cardboard carton on ice
with a plant specimen and a wet paper towel
to prevent dehydration. Live weights were
measured with a Cent-O-Gram triple beam
balance sensitive to ± 5 mg.
Song Analysis
We recorded calling songs of both species
on 1/4 inch audio tape using a Uher 4000
Report Monitor portable tape deck and an
Electro-Voice RE 55 dynamic microphone.
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HABITAT SHARING IN OKANAGANA SPP.
Songs were recorded at a tape speed of 19 cm
⋅ sec–1. Recordings were analyzed with MacSpeech Lab II (GW Instruments, Somerville,
MA) and a Macintosh computer. Recordings
were digitized at a sampling rate of 40 kHz,
and a narrow band FFT was used to determine peak frequency. Figures 1–4 were generated with a Kay Elemetrics Corporation Digital Sona-graph 7800 using an intermediate
bandwidth analysis filter and a Sona-graph
Printer 7900.
Temperature Responses
In the laboratory we recorded insect body
temperatures at the minimum temperature for
controlled flight, maximum voluntary tolerance
or shade-seeking temperature, and temperature of heat torpor. Minimum flight and heat
torpor temperatures represent body temperature limits of full activity since cicadas with
body temperatures beyond this range are torpid. Maximum voluntary tolerance temperature
represents a thermoregulatory point (Heath
1970). Procedures we used to determine thermal responses were the same as those used in
previous cicada studies (Heath 1967, Heath
and Wilkin 1970).
Temperatures were measured with a Physitemp Model BAT-12 digital thermocouple
thermometer and a type MT 29/1 29 gauge
hypodermic microprobe copper/constantan thermocouple that had been calibrated to a National
Institute of Standards and Technology mercury
thermometer. Body temperatures were measured by inserting the probe midway into the
dorsal mesothorax. When an animal was oriented for insertion of the probe, we handled it
by grasping the wingtips between the thumb
and forefinger. This procedure prevented conductive heat transfer between the insect and
the experimenter. All body temperatures were
recorded within 5 seconds of the insect performing the activity of interest.
To determine the minimum flight temperature, we repeatedly tossed a pre-cooled animal
vertically 1–2 m. As the animal warmed, it
began attempts at flight. Initially, it produced
small wing movements with the wings folded
against the body. As body temperature increased, the wings were extended and normal
flight movements of the wings began. The animals sometimes glided as they warmed before
they could fly efficiently. When an animal made
439
a controlled flight or landing, we recorded body
temperature.
Maximum voluntary tolerance was determined by placing a pre-cooled animal on a
vertical surface and warming the insect with a
heat lamp. The heat lamp was placed 45–50
cm from the vertical surface, and the insect
was placed in the center of the beam emanating from the lamp. Animals basked in the heat
produced by the heat lamp until their body
temperature reached the maximum voluntary
tolerance temperature. When body temperature
corresponding to maximum voluntary tolerance
was reached, the animals walked or flew out of
the central portion of the heat lamp. When an
animal began to move, we measured body
temperature.
Temperature of heat torpor was determined
by placing an animal in a cardboard container
and heating the insect with a heat lamp. The
container prevented the specimen’s escape during heating. Body temperature of the insect was
measured when motor control ceased due to
increase in body temperature. Heat torpor
temperature is not a lethal temperature, and
animals recover after their body temperature
has decreased to the temperature range normally experienced.
Field Temperatures
We recorded body temperatures of singing
animals in the field. Animals were captured in
an insect net, which contracted around the
animal to prevent movement. The temperature
probe was inserted through the net into the
dorsal mesothorax of the animal to measure
body temperature. This procedure prevented
conductive heat transfer between the experimenter and the animal that could have altered
insect body temperature. All body temperature
measurements were made within 5 seconds of
capture. Species identification of each specimen
was made after measuring body temperature.
Thermoregulation in ectothermic cicadas
can be modeled as a coupled on-off regulator
(Heath et al. 1971a). When body temperature
is below a certain set point, the animal remains
exposed to solar radiation. Whenever the body
temperature exceeds this set point, the cicada
retreats to shade. The degree of radiant heating
is altered by changing activity location. The
insect can obtain a similar result by changing
body orientation with respect to the sun.
440
We collected behavioral data on body orientation at the same time that body temperatures
were recorded in the field to determine if the
cicadas behaviorally thermoregulate. Orientation of the animals with respect to the sun can
be interpreted as an indication of “preferred”
thermal state. Animals with their bodies positively oriented to the sun are positioned to
maximize radiant heat gain and can be thought
of as attempting to elevate body temperature.
Negatively oriented animals have minimized
heat gain from solar radiation and may be
viewed as trying to maintain or decrease body
temperature. An animal that is oriented with
the side of the body toward the sun may be
thought of as being near its “preferred” body
temperature. Side-orientation permits the animal to increase or decrease body temperature
slowly, depending upon the rate of radiant heat
input and the rate of heat loss to the environment.
Information on calling activity was obtained
by determining which species were singing at
different times of the day. Species determination was made through animals captured for
body temperature measurements and from
calls being produced by uncaptured animals.
All statistics are reported as mean ± 1 standard error.
RESULTS
The species involved in the present study
are medium-sized cicadas. Live weight determined for O. striatipes (386.76 ± 10.510 mg,
n = 17) is significantly smaller (t = –7.905, df
= 52, P << 0.0001) than measured weight of
O. utahensis (582.70 ± 16.025 mg, n = 37).
The calling song of O. striatipes is a continuous train of constant-amplitude sound pulses
(Fig. 1C). The song begins as a train of syllables of varying duration and interburst intervals (Fig. 1A). Syllables begin to fuse together
(Fig. 1B) until sound pulses become a continuous train, producing the calling song. Frequency spread of the song is approximately 7
kHz to 12 kHz. Peak sound energy in the power
spectrum is 9.74 ± 0.345 kHz (n = 7, range
8.56–10.27 kHz). Expansion of the time wave
(Fig. 2) shows sound pulses are produced at a
rate of approximately 247 ± 42 pulses ⋅ sec–1
(n = 7, range 181.6–321.9).
The calling song of O. utahensis is composed
of a train of syllables (Fig. 3). Each syllable
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(Fig. 4) is composed of about 26 individual
sound pulses (26.1 ± 0.34, n = 21, range 23–29).
The syllables are 87.0 ± 1.6 msec in duration
(n = 21, range 75.6–107.2) and separated by
6.3 ± 0.62 msec (n = 20, range 3.6–12.57). The
pulse repetition rate is 300.96 ± 17.81 sec–1
within each syllable (n = 21, range 251.9–
325.8). Sound energy of the call is distributed
between 6 kHz and 11 kHz. There is an increase in intensity and a change in the emphasized frequency midway through the syllable.
Peak energy is at 8.85 kHz (8.85 ± 0.07 kHz,
n = 21, range 8.20–9.36) near the beginning of
a syllable and 9.13 kHz (9.13 ± 0.07 kHz, n =
21, range 8.36–9.64) when the intensity increases midway through a syllable. Sound
energy is concentrated in a more narrow frequency range or is more sharply tuned during
the earlier portion of the call.
Table 1 summarizes temperature responses
of O. striatipes and O. utahensis. Minimum flight
temperatures are approximately equal (t =
–0.0888, df = 52, P = 0.4986). Mean maximum voluntary tolerance and heat torpor temperatures are not significantly different (t =
–0.7983, df = 54, P = 0.2142 and t = –0.8137,
df = 52, P = 0.2098, respectively).
Figure 5 compares the number of each
species singing in a given body temperature
range. Mean body temperatures of singing
animals for each species are significantly different (t = –7.0385, df = 56, P << 0.00001),
with O. utahensis singing at higher body temperatures than O. striatipes. Body temperatures of singing animals range from 33.5°C to
37.8°C in O. striatipes and from 34.9°C to
40.2°C in O. utahensis.
The relationship between maximum voluntary tolerance temperatures and mean singing
temperatures is different for each species
(Table 2). Okanagana striatipes sings at a body
temperature approximately equal to and not
significantly different from (t = –0.6773, df =
34, P = 0.2514) the maximum voluntary tolerance temperature of the species. However, O.
utahensis sings at a body temperature significantly greater than the maximum voluntary
tolerance temperature determined for the
species (t = –5.3755, df = 72, P << 0.0001).
Both O. striatipes and O. utahensis are ectothermic behavioral thermoregulators. Solar
radiation is used to elevate body temperature
for activity. Shuttling movements between
sunny and shaded perches and changes in
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441
Fig. 1. Okanagana striatipes calling song. Upper trace in each pair is the time wave and lower trace is the sona-gram.
Frequency spread of the song is approximately 7 kHz to 12 kHz. The song begins as a train of syllables varying in duration and interburst interval (A). Syllables become longer and begin to fuse together (B) until the sound pulses produce
the continuous calling song (C).
body orientation are then used by both species
to regulate body temperature during activity.
O. striatipes at lower body temperatures (33–
34°C) illustrated in Figure 5 were positively
oriented to the sun. Animals with highest body
temperatures (36–37°C) were negatively oriented or side-oriented. Animals in the central
body temperature range (34–36°C) were positively oriented, side-oriented, or negatively
oriented. Most (7 of 9) O. striatipes with body
temperatures below the recorded mean singing
temperature (35.87°C) for the species were
positively oriented. Animals with body temperatures greater than the mean were found in
all possible states of orientation. These data
suggest the animals were actively regulating
body temperature around the mean temperature recorded.
Okanagana utahensis showed a similar pattern of orientation with respect to the sun.
Animals with lower body temperatures (34–
37°C) were positively oriented while animals
442
Fig. 2. Expanded time wave of the Okanagana striatipes
calling song. Sound pulses are produced at a rate of about
182 sec–1 during the full song.
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Although these generalizations of the calling activity in the 2 species hold true, activity
patterns can be altered by ambient conditions.
Singing activity in both species is inhibited by
extremely high ambient temperatures. Similarly, on a mostly cloudy day, O. striatipes sang to
a greater degree than normal, and O. utahensis sang to a lesser degree than normal in
the early afternoon. Clouds not only prevented
Ta from rising to a level that would passively
raise body temperature high enough for singing,
but also prevented O. utahensis from using radiant heat to elevate body temperature. On the
other hand, O. striatipes was able to elevate
body temperature a sufficient amount and continued calling while activity was suppressed in
O. utahensis due to the ambient conditions.
DISCUSSION
at the upper end of the distribution (39–41°C)
showed negative or side-orientation. The difference between the species is that O. utahensis remains positively oriented at a temperature range (36–37°C) when O. striatipes has
positioned itself to decrease radiant heat gain.
Although both species are behavioral thermoregulators, O. utahensis regulates body temperature at a higher temperature than O. striatipes.
Acoustic activity of the 2 species differs
throughout the day, producing a partial temporal separation of acoustic activity that is
dependent on species-specific thermal preferences. Both species show an initial burst of
activity in the morning when ambient conditions are sufficient to elevate body temperature to the species-specific level required for
singing. Okanagana striatipes begins to sing
before O. utahensis. The lower body temperature required for singing gives O. striatipes a
period of about 20–30 minutes in the morning
when it is the only cicada species calling. When
ambient conditions are sufficient to elevate
body temperature in O. utahensis, this species
begins to sing as well. Singing in O. striatipes
begins to decline as O. utahensis proceeds
through its initial peak of acoustic signaling,
which lasts 2.5 to 3 hours. As ambient temperature (Ta) continues to rise during the afternoon, O. utahensis continues sporadic signaling while activity in O. striatipes decreases.
Okanagana striatipes then resumes activity to
a greater degree as Ta falls in the late afternoon
while activity in O. utahensis is suppressed by
falling body temperature.
Thermal requirements of O. striatipes and
O. utahensis represent a possible mechanism
to decrease the potential for acoustic interference. Laboratory temperature responses of the
2 species are approximately equal (Table 1).
This would be expected in 2 animals sharing a
habitat because they are exposed to the same
environmental conditions. However, the mean
body temperature of singing O. utahensis is
significantly greater than the mean body temperature of singing O. striatipes (Table 2). It
appears O. utahensis “prefers” or requires a
higher body temperature to coordinate singing
activity.
Crawford and Dadone (1979) suggested that
temperature sets limits on the ability of cicadas
to coordinate motor control of singing. The
rate of action for potential firing in the timbal
nerve is temperature dependent (Wakabayashi
and Hagiwara 1953, Wakabayashi and Ikeda
1961). Raising thoracic temperature during
activity in Cystosoma saundersii causes the song
cycle period to change (Josephson and Young
1979), and the change in body temperature in
Tibicen winnemanna (Davis) during endothermic warming is responsible for changes in
acoustic activity of that species (Sanborn 1997).
These data suggest the ability of the cicada
nervous system to coordinate calling songs is
temperature dependent.
Cicadas perform complex activities, such as
singing, over a small temperature range. The
temperature range may represent the maximum range over which the cicada can adjust
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443
Fig. 3. Okanagana utahensis calling song. The song is composed of a train of syllables with sound energy between 6
kHz and 11 kHz.
its rate of activity to compensate for the direct
effect of temperature on metabolic processes
(Heath et al. 1971b). The body temperature
range of singing ectothermic cicadas has been
reported as 25.0–31.8°C in Magicicada cassinii
(Heath 1967), 32.2–41.5°C in Tibicen chloromerus (Walker) (Sanborn 2000), 35.0–40.8°C
in Diceroprocta olympusa (Walker) (Sanborn
and Maté 2000), 33.5–43.0°C in Cacama valvata (Uhler) (Heath et al. 1972), 38.0–41.9°C
in Okanagodes gracilis Davis (Sanborn et al.
1992), and 39.0-41.8°C in Okanagana hesperia
(Uhler) (Heath 1972). Body temperatures of
singing O. striatipes were measured between
33.5°C and 37.8°C. Body temperatures of calling O. utahensis ranged from 34.9°C to 40.9°C,
both within the range of body temperatures
reported for other cicadas to coordinate singing activity. The range is also similar to the
body temperature range of singing in endothermic cicada species (Sanborn et al. 1995a,
1995b, Sanborn 2000).
Comparison of the data in the singing temperature histogram (Fig. 5) suggests the species
select different body temperature ranges when
singing. Orientation of the species with respect
to the sun also suggests different thermal preferences. Okanagana striatipes begins thermoregulatory behaviors to decrease body temperature while at the same body temperatures O.
utahensis continues to maximize radiant heat
gain. Comparison of maximum voluntary tolerance temperatures and mean singing tem-
peratures of each species illustrates a difference in thermal activity of the species. Okanagana striatipes sings at a body temperature
approximately equal to the maximum voluntary
tolerance temperature of the species, while O.
utahensis sings at a body temperature significantly greater than the maximum voluntary
tolerance temperature determined for the
species. Although maximum voluntary tolerance temperatures are approximately equal
between species, O. utahensis selects a higher
body temperature range for activity. Singing at
body temperatures greater than the maximum
voluntary tolerance temperature has also been
described in the cicada Diceroprocta apache
(Davis) (Heath and Wilkin 1970).
Singing at temperatures above an upper
thermoregulatory point suggests O. utahensis
requires an elevated body temperature for the
singing mechanism to function properly. Okanagana utahensis may require a higher body
temperature due to the song parameters of the
species. Amplitude modulations within syllables, production of syllables themselves, and/or
greater pulse repetition rate of the O. utahensis song may require a higher body temperature for coordination than that required by
O. striatipes to coordinate a continuous, unmodulated song. Okanagana utahensis produces
sound pulses at a rate of approximately 300
sec–1 compared to 250 sec–1 in O. striatipes.
Since timbal muscle contraction kinetics are
temperature dependent (Josephson and Young
444
Fig. 4. Expanded time wave of the Okanagana utahensis
calling song. Syllables are composed of approximately 26
individual sound pulses produced at the rate of 300 sec–1.
Syllables are about 87 msec in duration and separated by
6.3 msec. Intensity increases and dominant frequency
changes midway through the syllable produced by lateral
abdominal movements.
1979, 1985, Josephson 1981, Young and Josephson 1983, Sanborn 2001), the greater pulse
repetition rate may require a higher timbal
muscle temperature for the timbal to contract
at the frequency necessary to produce the calling song.
Temperature requirements affect daily activity cycles of the 2 species. It is through this
temporal separation of species activity that differences in singing body temperature become
important. The lower body temperature of singing O. striatipes permits the species to sing in
the morning and late afternoon when O. utahensis is potentially unable to raise body temperature to the range necessary for acoustic
activity. Similarly, the higher body temperature required by singing O. utahensis permits
acoustic activity during the heat of the day
when O. striatipes is forced to retreat to shaded
sites. Thus, thermal requirements act to separate reproductive activity temporally and to
reduce or eliminate acoustic interference between the species. Thermal separation of activity has been described in ants (Cros et al.
1997), beetles (Colombini et al. 1994, Fallaci
et al. 1997), and flies (Gaugler and Schutz 1989,
Schutz and Gaugler 1992) that share a habitat.
Cicadas have developed several behavioral
and physiological methods to minimize acoustic
[Volume 62
interference, competition for environmental resources, and interspecific interactions. Cicadas
using similar songs for communication or similar host plants can avoid interspecific competition through geographic separation (Pringle
1954, Fleming 1971), microhabitat segregation
(Schedl 1986, Riede 1997), temporal separation
of calling times (Hayashi 1975a, Wolda 1993,
Gogala and Riede 1995, Riede 1995, 1997) or
time of year the species are active (Young
1981a, 1981b), or maximizing communicatory
differences (Fleming 1971, Walker 1974).
Okanagana striatipes and O. utahensis, however, are similar morphological species using
the same host plant; they are active in the
same place at the same time of year; males of
both species are solitary animals when calling;
they produce mating calls that overlap in frequency; and both are diurnally active. They
are species that contradict the general patterns
used by cicadas to avoid interspecific competition, and yet they are able to share an environment while using the same resources, both
physical and acoustic.
The songs of cicadas act as an isolating
mechanism between species (Alexander 1957,
Alexander and Moore 1958, Moore and Alexander 1958, DuMortier 1963, Haskell 1974,
Bennet-Clark 1975, Fleming 1975, 1984). When
related sympatric species share an environment, selection should minimize signal differences within a species and maximize differences between species (Alexander 1967, Walker
1974, Young 1981a). In general, sympatric species differ markedly in calling song structure
and/or frequency (Pringle 1954, Alexander
1956, 1957, 1967, Moore and Alexander 1958,
Alexander and Moore 1962, DuMortier 1963,
Fleming 1971, 1984, Walker 1974, Young 1981a).
However, O. striatipes and O. utahensis possess
similar songs and acoustic behavior.
The overlap of calling song frequency in O.
striatipes and O. utahensis is probably due to
the similar size of the animals. The frequency
of a cicada’s song is determined by the natural
period of timbal vibration, which is then modified by several body parts (Pringle 1954, Moore
and Sawyer 1966, Popov 1975, Popov et al.
1985, Huber et al. 1990, Bennet-Clark and
Young 1992, Fonseca 1996, Bennet-Clark 1997,
1999) and scaled to body size (Daniel et al.
1993, Bennet-Clark and Young 1994). Because
the 2 cicadas are physically similar in size,
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445
TABLE 1. Temperature responses (°C) of Okanagana striatipes and Okanagana utahensis from Cortez, Colorado.
O. striatipes
(mean ± sx–)
O. utahensis
(mean ± sx–)
20.66 (± 0.348)
n = 17
20.71 (± 0.313)
n = 37
35.46 (± 0.571)
n = 17
36.03 (± 0.404)
n = 37
45.52 (± 0.605)
n = 17
46.08 (± 0.375)
n = 37
Behavior
Minimum flight temperature
Maximum voluntary tolerance
temperature
Heat torpor temperature
All interactions P > 0.2.
timbal size is probably similar in the species
and call frequencies also should be similar.
The slightly larger size, together with a slightly larger timbal, of O. utahensis is probably responsible for the lower emphasized frequency
of the call.
Calling song temporal patterns have been
suggested as a means of separating many sympatric species of cicadas. Four species of Maoricicada are thought to remain isolated by the
pulse-repetition frequency of their calling songs
(Fleming 1971). Jiang (1985) suggested that
the number of sound pulses, pulse length, and
repetition frequency of amplitude-modulated
pulse trains separate Acutivalva choui Yao, Aola
bindusara (Distant), and Linguvalva sinensis
Chou and Yao. All 3 species inhabit the same
location, sing only from 0630 to 0645 hours,
and produce calls of similar frequency. The
sympatric sibling species Platypleura maytenophila Villet and P. hirtipennis (Germar) (Villet
1987) overlap in calling song frequency but
differ in temporal pattern (Villet 1988). Nakao
and Kanmiya (1988) showed that there are significant differences in the songs produced by
the cicada Meimuna kuroiwae Matsumura over
its entire range. Meimuna kuroiwae is a synonymized species of what were originally 7 independent species (Hayashi 1975b) and should
probably be classified as separate species based
on their calling songs.
Temporal patterns of the songs of the 2
species we studied are markedly different.
Okanagana striatipes produces a continuous
train of constant-amplitude sound pulses (Fig.
1). The song of O. utahensis is a train of syllables that exhibit an amplitude-modulation pattern within each syllable (Fig. 3). Temporal
patterns of the songs probably facilitate segregation of the 2 species during interspecific
interactions.
Fig. 5. Distribution of body temperatures of singing
Okanagana striatipes (solid bars) and O. utahensis (striped
bars). Mean body temperature of singing O. striatipes is
35.87 ± 0.257°C (mean ± sx–, n = 19). Mean body temperature of singing O. utahensis is 38.62 ± 0.242°C (n = 39).
Mean singing temperatures are significantly different (t =
–7.0385, df = 56, P << 0.0001).
Abdominal movements also may produce
the amplitude modulation seen in O. utahensis
syllables. The cicada abdomen acts as a resonating structure, increasing the volume of the
sound produced (Pringle 1954, Moore and Sawyer 1966, Young 1972, Simmons and Young
1978, Bennet-Clark 1999). When the abdomen
is tuned to the natural period of timbal vibration, the intensity of the song increases (Pringle
1954). The observed increase in intensity may
also be facilitated by changes in abdominal
position. By altering the gap between the
opercula and the tympana, the tension placed
on the timbals, tympana, and folded membrane
is changed, causing an increase in sound
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TABLE 2. Comparison of maximum voluntary tolerance temperatures (°C) and field singing temperatures (°C) of
Okanagana striatipes and Okanagana utahensis.
Maximum voluntary
tolerance temperature
(mean ± sx–)
Singing
temperature
(mean ± sx–)
Okanagana striatipes
35.46 (± 0.571)
n = 17
35.87 (± 0.257)b
n = 19
Okanagana utahensis
36.03 (± 0.404)a
n = 37
38.62 (± 0.242)ab
n = 39
Species
a,bP << 0.001.
intensity (Pringle 1954, Weber et al. 1987, Villet 1988, Young 1990). Another possibility for
the increased intensity could be increased activity in the timbal tensor muscle during the syllable (Hennig et al. 1994). Altering the size of
the opercula-tympanal distance may also produce change in the emphasized frequency within O. utahensis syllables as described in many
species of cicadas (Allard 1946, Young 1972,
Joermann and Schneider 1987, Sanborn 1997).
Although cicada auditory receptors react to
allospecific calls, the response of the auditory
neurons to different portions of the calls could
help to separate O. striatipes and O. utahensis.
Amplitude modulation, intensity changes, and
syllables of the O. utahensis calling song represent stimuli to which the cicada auditory
system has already been shown to be sensitive
(Hagiwara and Ogura 1960, Katsuki 1960, Katsuki and Suga 1960, Huber et al. 1979, 1980).
Production of syllables by O. striatipes prior to
production of the calling song may negate any
benefit O. utahensis has in possessing a song
constructed of syllables.
Temporal patterns of the full calling songs
may be sufficient in isolating O. striatipes and
O. utahensis, but the overlap in frequency could
cause acoustic interference between the 2 species. Walker (1986) collected 2 species of cicadas
attracted to synthetic cricket calls. Doolan and
Young (1989) showed that the call frequency of
Cystosoma saundersii Westwood is important
in eliciting steering behavior in tethered females. The correct temporal pattern of the
species song is necessary for the females to
exhibit courtship behavior. If O. striatipes and
O. utahensis possess a similar 2-step recognition process, simultaneous calling could cause
females to waste time and energy in interspecific interactions. In addition, females flying to
males of the wrong species could be exposing
themselves to predation. Flying C. saundersii
females fall victim to bird predation when flying to a calling male (Doolan and MacNally
1981). Different thermal requirements of each
species decrease the chance of acoustic interference between these closely related species
of Okanagana.
We therefore suggest that O. striatipes and
O. utahensis are able to share the same environment by (1) producing calling songs of different temporal patterns and (2) utilizing different thermal requirements that permit and/or
facilitate temporal separation of the day.
ACKNOWLEDGMENTS
The authors wish to acknowledge the assistance of Polly K. Phillips in the field. The study
was supported in part by a traineeship from
USPHS GMSO7143 to AFS. Aaron Ellinson
made helpful comments on the manuscript.
LITERATURE CITED
ALEXANDER, R.D. 1956. A comparative study of sound
production in insects with special reference to the
singing Orthoptera and Cicadidae of the eastern
United States. Doctoral dissertation, Ohio State University, Columbus.
______. 1957. Sound production and associated behavior
in insects. Ohio Journal of Science 57:101–113.
______. 1967. Acoustical communication in arthropods.
Annual Review of Entomology 12:495–526.
ALEXANDER, R.D., AND T.E. MOORE. 1958. Studies on the
acoustical behavior of seventeen-year cicadas (Homoptera, Cicadidae, Magicicada). Ohio Journal of Science 58:107–127.
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Received 18 January 2001
OREOHELICES OF UTAH, II. EXTANT STATUS OF THE BRIAN HEAD
MOUNTAINSNAIL, OREOHELIX PARAWANENSIS GREGG, 1941
(STYLOMMATOPHORA: OREOHELICIDAE)
George V. Oliver1 and William R. Bosworth III1
ABSTRACT.—The Brian Head mountainsnail, Oreohelix parawanensis, is reported for the 1st time as a living species,
and for the 1st time its habitat is described. Preliminary determination of the very limited distribution of this species
(≤2.3 ha inhabited in ∼11 ha overall area) is presented. Morphometric data previously had been reported only for the
holotype and for 1 topotype; measurements from 37 new specimens as well as 20 paratypes are provided here, and these
data show that the lost holotype was not typical of the species. Sizes of reproductive snails and of embryos are also
reported.
Key words: Oreohelix parawanensis, Brian Head mountainsnail, gastropods, mollusks, Parowan Mountains, Utah.
Oreohelix parawanensis, the Brian Head
mountainsnail, was described by Gregg (1941a)
based on 31 specimens, “all dead and many of
them immature,” that he collected “from a
rock slide on the southwest slope of Brian
Head, Parawan [sic] Mountains, Iron County,
Utah, altitude about 11,000 [ft].” It is likely
that he collected these specimens during the
summer of 1935, when he spent “nearly three
months at Cedar Breaks National Monument”
(Gregg 1941b), which is only about 2 km from
Brian Head (Peak) and where he extensively
collected mollusks.
Bickel (1977) searched for O. parawanensis
and reported: “The type locality was collected
June 27, 1975 at which time the rock slide
holding the population was still covered by
several feet of snow, and only a few empty shells
were collected from its margin.” Clarke and
Hovingh (1994), too, searched for O. parawanensis at the type locality, in 1992 (Clarke
1993), noting that “no live specimens have
ever been collected.” They reported that they
“searched this rock slide carefully and excavated the surface from one side to the other”
and conducted “[c]areful searches elsewhere
. . . in the vicinity,” but they were able to find
only a single empty shell of O. parawanensis,
and their efforts “failed to produce any live
specimens.”
Despite the fact that the Brian Head mountainsnail, O. parawanensis, had never been
demonstrated to be extant and could have
been extinct before the arrival of Europeans in
North America, state and federal management
agencies have shown considerable interest in
the conservation of this species (as discussed
below). Efforts to determine its current status
had been inadequate, field investigations having been poorly timed seasonally and of insufficient duration, one such search (Bickel 1977)
having been conducted too early in the season
when the locality was under snow and apparently carried out for only part of a day, and the
other search (Clarke 1993) having been made
too late in the season—October, under conditions of “very strong wind & cold”—evidently
involving only a little more than 4 person-hours
of effort. Furthermore, none of the previous
work (Gregg 1941a, Bickel 1977, Clarke and
Hovingh 1994) had revealed any aspects of the
biology of this species. As part of our continuing studies of mountainsnails in Utah (Oliver
and Bosworth 2000), our goals were to attempt
to find living examples of O. parawanensis, to
make a preliminary assessment of its distribution and habitat, and to examine available
specimens morphometrically.
METHODS
We scheduled our fieldwork to search for
living examples of O. parawanensis to coincide
with conditions favorable for finding active
1Utah Natural Heritage Program, Utah Division of Wildlife Resources, 1594 West North Temple, Salt Lake City, UT 84114.
451
452
terrestrial mollusks, especially mountainsnails
(Oreohelix). Oreohelices of other species are
known to hibernate by burrowing beneath the
surface of the soil ( Jones 1935, 1940), and
moderately warm conditions prevail for only
a few weeks each year on Brian Head. We
searched for O. parawanensis on 11, 12, and
13 August 1998 on Brian Head. During this
time daytime temperatures ranged from ∼15°C
to ∼20°C, and summer storms produced brief,
light rain showers each day. We established 14
collecting stations on the southern and western slopes of Brian Head, all within 0.7 km of
the summit.
We made an effort to sample a variety of
habitat conditions, but, because most members of the genus Oreohelix are strongly calciphilic (Henderson and Daniels 1916, Pilsbry
1916, 1939, Jones 1940), we focused efforts in
areas where we could locate exposed limestone.
From the summit of Brian Head, we scanned
the slopes, using binoculars, looking for surface limestone to aid us in choosing suitable
collecting stations. For each collecting station
we noted plant association (dominant plant
species present), substrate type, and other variables such as slope and aspect, as well as all
mollusks that we found.
RESULTS AND DISCUSSION
We found O. parawanensis at 7 of our 14
collecting stations on Brian Head. At these 7
stations we found empty shells of O. parawanensis, and at 4 of the stations we discovered
the 1st living examples of this species. Oreohelix parawanensis was overall the most common
gastropod at the stations where it was detected.
We found and collected 49 empty shells (as
well as 5 embryos) of O. parawanensis, and we
found 18 live individuals, of which we took 8.
One of the living examples of O. parawanensis
was active and extended when found under a
rock between 0938 and 0948 hours on 13
August 1998; less than an hour earlier (at 0857
hours), air temperature at the locality was 16°C.
Of the new specimens, we have deposited
5 in the collection of the Academy of Natural
Sciences of Philadelphia (ANSP 401984) and 5
in the collection of the Los Angeles County
Museum (LACM 152567), these being the 2
collections that house all of Gregg’s type
material of this species that we have located.
[Volume 62
Type Locality
The type locality of O. parawanensis stated
by Gregg (1941a), while adequate and, for its
time, rather precise, can now be stated even
more precisely. The only problematic part of
Gregg’s (1941a) locality is his ambiguous phrase
“a rock slide.” While others (e.g., Bickel 1977,
Clarke and Hovingh 1994) have referred to
“the rock slide” as though there were no doubt
about the location, many parts of the southern
and western faces of Brian Head consist of talus
material of varying dimensions, and various
terms could be applied to these geological features, including the term “rock slide.” Although
the exact site of Gregg’s collection of O. parawanensis may never be known, we believe the
following is the most probable location: Utah,
Iron County, Brian Head (Peak), below summit
on SW face, T36S, R9W, section 13, NW1/4 of
NW1/4 of NW1/4.
Distribution
The distribution of O. parawanensis, as revealed by our fieldwork, can be summarized
as follows:
Utah, Iron County, Brian Head (Peak), below summit on SW face; T36S, R9W, section 13 (NW1/4 of
NW1/4 of NW1/4), section 14 (E1/2 of NE1/4 of
NE1/4), and section 11 (E1/2 of SE1/4 of SE1/4).
The 7 collecting stations in which we found
this species are contained within an irregular,
but somewhat triangular, polygon of about 11
ha. However, only small patches within this
overall distributional area appeared to provide
suitable habitat, and we found O. parawanensis in only 7 of these patches. The total area
that we determined to be inhabited by this
species is only about 2.3 ha or less, although
this is a rather crude estimate and one that
should be considered the minimal area of its
occurrence. Because we found only dead shells
of O. parawanensis at 3 of the collecting stations, it is possible that these represented
extinct colonies, and the currently inhabited
area may be even smaller than that stated
above.
Habitat
The habitat of O. parawanensis has not previously been described (e.g., Gregg 1941a,
Pilsbry 1948, Bickel 1977, Clarke 1993, Clarke
and Hovingh 1994). Of the 7 stations wherein
we found O. parawanensis, 4 were on limestone
2002]
EXTANT STATUS OF BRIAN HEAD MOUNTAINSNAIL
substrates; 2 were in areas of primarily basaltic
rock with some limestone and, in 1 case, a little sandstone; and 1—the highest location and
probably the type locality—was almost entirely
basaltic rock. Slope in the places where we
found O. parawanensis varied from almost none
to about 40°, and elevations of the inhabited
patches ranged from 3255 m to 3340 m. All 18
live individuals of O. parawanensis were under
surface rocks, mostly single individuals but
rarely more, 4 being the largest number of live
O. parawanensis we found under 1 rock. Most
dead shells of this species that we found were
also under rocks, though a few were lying exposed on the ground.
The stations inhabited by O. parawanensis
almost without exception contained dense
clumps of currants of 2 species, wax currant
(Ribes cereum) and gooseberry currant (Ribes
montigenum). Meadow rue (Thalictrum cf.
fendleri) and Indian paintbrush (Castilleja sp.)
were also typical of the places where we found
O. parawanensis. Another plant that characterized several of the stations where O. parawanensis was present was ground juniper ( Juniperus communis). Limber pine (Pinus flexilis),
western bristlecone pine (Pinus longaeva), and
Engelmann’s spruce (Picea engelmannii) were
present at some of the stations where we found
O. parawanensis, which were at or slightly
above tree line. Other species of forbs, a few
grasses, and a few other trees also were present at some of the stations that yielded O.
parawanensis.
Associated Gastropods
The diversity of gastropod species that we
found on Brian Head was rather low, as could
be expected at such a high, cold, and barren
location. Most, but not all, other gastropod
species we found on Brian Head were at stations shared with O. parawanensis. The gastropods we found at the same collecting stations
as O. parawanensis were the Rocky Mountain
column (Pupilla blandi), the crestless column
(Pupilla hebes), a vallonia (Vallonia cf. cyclophorella), the Rocky mountainsnail (Oreohelix
strigosa), and the western glass-snail (Vitrina
pellucida). Most of these associated gastropods
were found in small numbers. We found O.
strigosa to be the most common associate of O.
parawanensis, the 2 congeners occurring together in similar numbers.
453
All of these species were reported by Gregg
(1941a) as associates of O. parawanensis, along
with the multirib vallonia (Vallonia gracilicosta),
the spruce snail (Microphysula ingersolli), the
quick gloss (Zonitoides arboreus), and the forest disc (Discus cronkhitei [= Discus whitneyi]),
although Gregg (1941a) did not indicate how
close their association with O. parawanensis
was. In the above list we have not included
other species we found on Brian Head that
were not at the same collecting stations that
yielded O. parawanensis.
Conservational Considerations
O. parawanensis was formerly a Category 2
candidate for listing by the U.S. Fish and
Wildlife Service, under provisions of the Endangered Species Act, until 28 February 1996,
when Category 2 was eliminated. Furthermore,
even though O. parawanensis had never been
reported to be living and thus was not known
to be extant, it was indicated as “declining”
(U.S. Fish and Wildlife Service 1994). Oreohelix parawanensis is listed as a species of special concern by the state of Utah (Utah Division of Wildlife Resources 1998).
The area inhabited by O. parawanensis is
within the Dixie National Forest. Because the
stations supporting this species are at or above
tree line, timber harvest is not a threat. Nearby ski resort operations (lifts, runs, buildings,
etc.) to the west and northwest do not currently threaten the snail; however, if resort operations were to expand into the area where O.
parawanensis occurs, its entire population could
be destroyed. The high elevation of the site
and the rather barren nature of the terrain
afford relatively good protection for the snail
from most other anthropogenic threats. An
unpaved road, however, loops around the
south side of Brian Head and up to the summit, where a small pavilion stands. Hikers and
mountain bikers utilize the area, and we encountered rock collectors near some of the inhabited stations. Bones, apparently those of a
domestic goat or sheep, were present near an
inhabited station, and we observed large numbers of domestic sheep on U.S. Forest Service
land less than 10 km away.
Type Specimens
Although Gregg (1941a) stated that the holotype of O. parawanensis had been deposited in
454
the collection of the Academy of Natural Sciences of Philadelphia (no. 176907), this specimen could not be located and is presumed lost
(Edward S. Gilmore, ANSP, personal communication, 1998). Gregg (1941a) also indicated
that the paratypes (presumably all of the other
30 specimens of O. parawanensis he collected)
were retained in his own collection (no. 324).
We located 20 paratypes in the collection
of the Los Angeles County Museum (LACM
1660) and 1 paratype in the collection of the
Academy of Natural Sciences of Philadelphia
(ANSP 340315). Labels associated with both of
these lots of specimens indicate they are paratypes, and the lot in the Los Angeles County
Museum also has Gregg’s catalogue number
(324) on 1 of the labels. Although the Los
Angeles County Museum catalogue indicates
19 paratypes are housed in that collection
(Lindsey Groves, LACM, personal communication, 1999) and there are 19 specimens of
moderate size in the lot, an additional tiny
specimen, an embryonic shell, is contained in
the same lot but is separated and protected in
a clear gelatin capsule. Thus 20—or 21, if one
includes the embryo—of Gregg’s 30 paratypes
have been located and examined.
Although Gregg (1941a) stated that all specimens of O. parawanensis he collected were
dead, examination of the specimens in the Los
Angeles County Museum suggests to us that
some of the paratypes may actually have been
alive when they were collected despite assertions (Gregg 1941a, Bickel 1977, Clarke 1993,
Clarke and Hovingh 1994) that no living examples had ever been found. Of the paratypes we
examined, 7 showed evidence of an epiphragm,
and 1 other shell contained dried soft tissue.
Such evidence, however, does not prove they
were alive when Gregg collected them; these
specimens may in fact be exactly as they were
discovered at the time of collection.
Morphometric Data
Previously published morphometric data are
available for only 2 specimens of O. parawanensis, the holotype (Gregg 1941a) and a shell
collected by Clarke and Hovingh (1994), for
which they reported “diameter 8.5 mm, height
5.0 mm, whorls 41/4.”
Gregg (1941a) considered many of his 31
specimens to be immature, presumably based
on size. However, we know of no easily applied
criterion that will distinguish mature from
[Volume 62
immature shells of O. parawanensis (but see
discussion below), there being no reflection of
the lip and no apertural teeth in this species.
Thus, we measured available intact shells large
enough that they could be assigned to this
species with confidence.
We measured shell diameter, height, and
umbilicus width and counted whorls of 20
paratypes (19 LACM, 1 ANSP) and 37 intact
examples among the new specimens of O.
parawanensis that we collected (Table 1). Our
sample of 37 new specimens and the sample of
20 paratypes were very similar for the 3 mensural characters (shell diameter, height, and
umbilicus width), whorl counts, and the 2 proportions calculated from the mensural data;
the similarity of the 2 data sets is apparent if
one compares ranges and means of characters
in Table 1. However, when morphometric data
for the paratypes and the new specimens are
compared with Gregg’s (1941a) data for the
now-lost holotype (Table 1), it can be seen that
the holotype was quite dissimilar from the
paratypes and new topotypes in its measurements and proportions. The holotype was larger
in diameter and more depressed (low-spired,
as shown by the diameter/height proportion)
than any specimen examined by us. Also, the
umbilicus was larger—both absolutely and relative to its diameter—in the holotype than in
any examined shell. The exceptionally large
umbilicus of the holotype is of particular interest since Gregg (1941a), in defining this species,
claimed that the larger umbilicus of O. parawanensis is of importance in distinguishing it
from the 2 species that he considered to be its
closest relatives, O. handi and O. eurekensis.
It would, of course, be desirable to reexamine the holotype, and especially to remeasure
its umbilicus width, but this unfortunately is
not possible, the holotype having been lost.
However, Gregg (1941a) did publish 4 photographs of O. parawanensis, an umbilical (ventral) view and an apertural (frontal) view, both
at approximately life size, and the same 2 photographs at greater than natural size. Pilsbry
(1948) reproduced these photographs. The
photographs were published again by Clarke
and Hovingh (1994), who incorrectly credited
them to Pilsbry (1948). Also, Clarke and Hovingh (1994) labeled the photographs as illustrations of the holotype, despite the fact that
neither the original (Gregg 1941a) nor the secondary source (Pilsbry 1948) had indicated this
2002]
455
TABLE 1. Morphometric data for Oreohelix parawanensis.
Character
Whorls (no.)
Diameter (mm)
Height (mm)
Umbilicus width (mm)
Diameter/umbilicus
Diameter/height
New specimens (n = 37)
____________________________
Range
Mean
sx–
3.00–5.00
3.72–9.62
2.34–6.48
0.78–2.25
3.93–5.78
1.44–1.90
4.14
6.81
4.15
1.50
4.66
1.65
0.07
0.20
0.14
0.06
0.09
0.02
Paratypes (n = 20)
____________________________
Range
Mean
sx–
3.50–4.38
5.28–8.26
3.14–4.72
0.85–1.78
3.79–6.21
1.56–1.83
4.09
6.79
3.95
1.37
5.06
1.72
0.06
0.19
0.11
0.06
0.13
0.02
Holotypea
4.33
10.5
5.4
3.0
3.50
1.94
aMeasurements and whorl count of the holotype reported by Gregg (1941a) and proportions derived from the reported measurements.
to be so—or even that the photographs represented only a single specimen.
Using Gregg’s (1941a) published photographs, we measured shell diameter and umbilicus width for the shell shown in umbilical
view and shell diameter and height for the
shell shown in apertural view. Although we do
not consider any of the originally published 4
photographs to be exactly natural size, our
measurements of the photographed shells did
allow us to calculate their proportions accurately. Using the photograph of the umbilical
view, we arrived at a diameter/umbilicus proportion of 3.55. While not within the range for
this proportion that we observed in 37 topotypes or in 20 paratypes, it is not quite so far
out of the range as the proportion (3.50) calculated using the measurements for the holotype
published by Gregg (1941a). In the photograph of the apertural view, diameter/height
= 1.77, which does fall within the range observed in both the new specimens and the
paratypes, unlike the proportion (1.94) calculated from Gregg’s (1941a) reported measurements of the holotype.
When we scaled the umbilical photographic
view to the diameter reported by Gregg (1941a)
for the holotype, 10.5 mm, the umbilicus then
measured 3.00 mm, exactly the umbilicus measurement reported by Gregg (1941a), which
strongly suggests that the photograph represents the holotype, as Clarke and Hovingh
(1994) assumed. However, scaling the apertural-view photograph to a diameter of 10.5
mm, the height became 5.93 mm as opposed
to 5.4 mm as reported by Gregg for the holotype. The specimen shown in apertural view,
then, either was not the holotype or, if it was
the holotype, was slightly mismeasured by
Gregg—off by ∼0.5 mm. Not only is height the
most difficult of standard shell measurements
to make with accuracy, but good tools for measuring small specimens may not have been
readily available to Gregg. Thus, we believe
that possibly the photograph showing frontal
view also may be of the holotype.
Embryos and Size
at Maturity
Four (10.8%) of the 37 intact dead shells of
O. parawanensis we collected contained embryonic shells, 3 with 1 embryo each and 1
with 2 embryos. These 5 embryos were 1.8 to
2.4 mm in diameter (mean = 2.2 mm), with 2
to 2.25 whorls (mean = 2.1). The embryo that
was with the paratypes in the collection of the
Los Angeles County Museum (LACM 1660)
measured 2.3 mm in diameter and had 2.25
whorls, in close agreement with the 5 we found.
The discovery of embryos inside 4 dead shells
provides information on the size of reproductive O. parawanensis. The 4 gravid snails measured 5.68 to 9.32 mm (mean = 6.98 mm) in
diameter and had 3.67 to 4.75 whorls (mean =
4.14 whorls). This shows that maturity in O.
parawanensis is reached by the time the snails
are 5.68 mm in diameter or have 3.67 whorls.
Only 3 of 20 paratypes (excluding the single
embryo) we examined were smaller than the
smallest of the gravid shells we found, and
these 3 measured 5.28, 5.30, and 5.42 mm in
diameter and had 3.67, 3.625, and 3.50 whorls,
respectively—almost the size of the smallest
shell we found that contained an embryo. Unless
most or all of the 10 paratypes that we could
not locate were smaller than the 20 that we
have seen, Gregg’s (1941a) statement that many
of his 31 specimens were immature was not
correct.
456
Nomenclature and
Systematic Placement
Clarke and Hovingh (1994), noting Gregg’s
(1941a) misspelling—at least by current spelling
convention—of the Parowan Mountains, declared: “Since the proper spelling is Parowan
the species name is here emended to Oreohelix parowanensis.” The alternate spelling
“Parawan,” however, has been used by various
authors and has appeared in other malacological literature (e.g., Herrington and Roscoe
1953). Most importantly, Gregg’s spelling of
the specific epithet, parawanensis, is consistent
with his spelling of the “Parawan Mountains”
in the original publication (Gregg 1941a), and
there is no evidence to suggest that it was not
Gregg’s intent to spell parawanensis as he did.
Clarke and Hovingh’s (1994) alteration of the
name, then, is a violation of Article 32(a), concerning correct original spelling, and an unjustified emendation under Article 33(a) of the
International Code of Zoological Nomenclature (Ride et al. 1999). Thus, the name correctly remains Oreohelix parawanensis.
Bickel (1977) stated that O. parawanensis is
taxonomically “probably invalid, a synonym of
O. strigosa depressa,” arguing that “O. parawanensis is a stunted population of the widespread, O. strigosa depressa.” As noted in the
type description (Gregg 1941a), Oreohelix
strigosa depressa occurs with O. parawanensis, and our work has corroborated this. To this
we can add that the 2 species maintain their
morphological distinctiveness where they coexist. These facts render implausible Bickel’s
(1977) suggestion that O. parawanensis could
be a synonym of O. strigosa. However, Gregg’s
(1941a) assertion that O. parawanensis “seems
nearest related to O. eurekensis . . . and O.
handi,” both of which it closely resembles in
size and shell morphology, is reasonable and
almost certainly correct. Pilsbry (1948) placed
O. parawanensis near O. handi, making it a
member of the O. yavapai species group and
not the O. strigosa species group as the genus
was organized by Pilsbry (1933, 1939), who
described the genus Oreohelix. Bickel’s (1977)
suggestion that O. parawanensis represents
merely a “dwarfed colony” of O. strigosa, moreover, raises the question of whether his “few
empty shells” were in fact O. parawanensis at
all and not shells of immature O. strigosa, which
possibly could be confused with O. parawanensis upon superficial inspection.
[Volume 62
The atypical size and proportions of the
holotype (discussed above), however, suggest
that the taxonomic distinction of O. parawanensis from such species as O. handi and O.
eurekensis should be reexamined.
ACKNOWLEDGMENTS
Adonia R. Henry’s assistance in the field
contributed importantly to the findings reported
here, and we extend to her our special thanks.
This work was supported in part by a 1998
Canon Exploration Grant from Canon USA,
Inc., administered by The Nature Conservancy,
and we are grateful to Canon USA, Inc., and
The Nature Conservancy for this support.
Additional financial support for this work
came from the Utah Reclamation Mitigation
and Conservation Commission, under the
Central Utah Project Completion Act. Lindsey
Groves of the Los Angeles County Museum,
Edward S. Gilmore and Gary Rosenberg of
the Academy of Natural Sciences of Philadelphia, and Eric A. Rickart of the Utah Museum
of Natural History provided help with curatorial matters—searches, specimen loans, and
cataloguing new material. The encouragement
of Larry Dalton, Michael Canning, and Bill
James, all of the Utah Division of Wildlife
Resources, throughout the course of this work
is much appreciated. We thank Art Metcalf,
University of Texas–El Paso, and an anonymous reviewer for suggestions that improved
the manuscript.
LITERATURE CITED
BICKEL, D. 1977. A survey of locally endemic Mollusca of
Utah, Colorado, Wyoming, Montana, North Dakota,
and South Dakota. Unpublished report prepared for
the U.S. Fish and Wildlife Service. Minot State College, Minot, ND. ii + 60 pp.
CLARKE, A.H. 1993. Status survey of fifteen species and
subspecies of aquatic and terrestrial mollusks from
Utah, Colorado, and Montana. Unpublished report
prepared for the U.S. Fish and Wildlife Service by
Ecosearch, Inc., Portland, TX. 77 pp. + appendices.
CLARKE, A.H., AND P. HOVINGH. 1994. Studies on the status of endangerment of terrestrial mollusks in Utah.
Malacology Data Net 3:101–138.
GREGG, W.O. 1941a. A new Oreohelix from southern Utah.
Nautilus 54:95–96 + plate 8.
______. 1941b. Mollusca of Cedar Breaks National Monument, Utah. Nautilus 54:116–117.
HENDERSON, J., AND L.E. DANIELS. 1916. Hunting Mollusca
in Utah and Idaho. Proceedings of the Academy of
Natural Sciences of Philadelphia 68: 315–339 + plates
XV–XVIII.
2002]
HERRINGTON, H.B., AND E.J. ROSCOE. 1953. Some Sphaeriidae of Utah. Nautilus 66:97–98.
JONES, D.T. 1935. Burrowing of snails. Nautilus 48:1–3.
______. 1940. A study of the Great Basin land snail[,] Oreohelix strigosa depressa (Cockerell). Bulletin of the
University of Utah 31 (4) [Biological Series 6]:1–43.
OLIVER, G.V., AND W.R. BOSWORTH III. 2000. Oreohelices
of Utah, I. Rediscovery of the Uinta mountainsnail,
Oreohelix eurekensis uinta Brooks, 1939 (Stylommatophora: Oreohelicidae). Western North American Naturalist 60:451–455.
PILSBRY, H.A. 1916. Notes on the anatomy of Oreohelix,
with a catalogue of the species. Proceedings of the
Academy of Natural Sciences of Philadelphia 68:
340–359 + plates XIX–XXII.
______. 1933. Notes on the anatomy of Oreohelix,—III,
with descriptions of new species and subspecies.
Proceedings of the Academy of Natural Sciences of
Philadelphia 85:385–410.
______. 1939. Land Mollusca of North America (north of
Mexico). Academy of Natural Sciences of Philadelphia, Monographs 3(1):I–XVII + 1–573 + i–ix.
457
______. 1948. Land Mollusca of North America (north of
Mexico). Academy of Natural Sciences of Philadelphia, Monographs 3(2): XLVII + 521–1113.
RIDE, W.D.L., H.G. COGGER, C. DUPUIS, O. KRAUS, A.
MINELLI, F.C. THOMPSON, AND P.K. TUBBS, EDITORS.
1999. International code of zoological nomenclature.
4th edition. International Trust for Zoological Nomenclature, London. XXIX + 306 pp.
U.S. FISH AND WILDLIFE SERVICE. 1994. Endangered and
threatened wildlife and plants; animal candidate review for listing as endangered or threatened species;
proposed rule. Federal Register 59:58982–59028.
UTAH DIVISION OF WILDLIFE RESOURCES. 1998. Utah sensitive species list. Salt Lake City, UT. 48 pp.
Received 26 January 2001
Accepted 5 October 2001
GEOGRAPHIC VARIATION IN PELAGE COLOR
OF PIÑON MICE (PEROMYSCUS TRUEI)
IN THE NORTHERN GREAT BASIN AND ENVIRONS
Leslie N. Carraway1 and B.J. Verts1
ABSTRACT.—Cluster analyses of values for hue, value, and chroma (based on Munsell soil-color charts) obtained at 6
points on pelages of 202 adult piñon mice (Peromyscus truei) from the northern Great Basin and environs produced dendrograms with specimens grouped into 5 clusters. In most instances distribution of specimens forming clusters reflected
those published for nominal races. In instances in which previous distributions of nominal races were not supported and
for specimens previously unclassified, geographic distribution of groups of color morphs was logical and suggested
avenues for additional research on geographic variation in the species.
Key words: Peromyscus truei, piñon mice, pelage color, geographic variation, northern Great Basin.
Piñon mice (Peromyscus truei) occur from
southwestern and central Oregon, southwestern Idaho, southwestern Wyoming (likely extirpated), eastern Colorado, extreme western
Oklahoma, and western Texas, south to central
Baja California, Mexico, central Arizona, and
southern New Mexico (Gafur et al. 1980, Hoffmeister 1981, Modi and Lee 1984, Clark and
Stromberg 1987, Caire et al. 1989, Carraway
et al. 1993, Davis and Schmidly 1994, RamírezPulido et al. 1996). Also, a disjunct population
occurs near the southern tip of Baja California
Sur (Hall 1981). In the northern Great Basin,
the species usually is associated with rocky
habitats vegetated by old-growth piñon pine
(Pinus edulis) or junipers (Juniperus; Hall 1946,
Hoffmeister 1951, Carraway et al. 1993), but
those in southwestern Oregon and northwestern California are associated with oak (Quercus kelloggii and Q. garryana) or redwood
(Sequoia sempervirens) and Douglas-fir (Pseudotsuga menziesii; Hoffmeister 1951, Fisher 1976).
Dorsal pelage of P. truei usually is some
shade of grayish brown, but color varies considerably geographically (Hall 1981). Five
nominal subspecies occur in the northern
Great Basin and adjacent areas: P. t. gilberti, P.
t. nevadensis, P. t. preblei, P. t. sequoiensis, and
P. t. truei. Pelage color originally was involved
in differentiating among these races (Shufeldt
1885, Allen 1893, Bailey 1936, Hall and Hoffmeister 1940, Hoffmeister 1941) and was used
extensively as a diagnostic character in the most
recent taxonomic treatment of the species (Hoffmeister 1951). The latter author described dorsal pelage color for the 5 races in the northern
Great Basin and environs based on Ridgway’s
(1912) classification as “Sepia,” “Buffy Brown,”
“Mummy Brown,” “Brussels Brown,” and
“Tawny-Olive to Saccardo’s Umber,” respectively. Nevertheless, no quantitative analysis
of pelage color as a character for distinguishing these races has been conducted.
In 1992 we collected specimens of P. truei
in southeastern Oregon about 225 km from
the nearest previously published record in the
state. Subsequently, we collected other specimens and located other records from regions
in Oregon and Idaho north and east of the
known range of the species (Carraway et al.
1993). Because of the proximity of several nominal races to these collection sites (Hoffmeister
1951), we chose not to assign the newly acquired
specimens to a subspecies at that time. To
assign newly acquired specimens to appropriate subspecies and to delineate more precisely
ranges of nominal subspecies of P. truei in the
northern Great Basin and environs, we analyzed pelage color of 202 museum specimens
collected in Oregon, Idaho, Nevada, northern
California, and western Utah.
1Department of Fisheries and Wildlife, Nash 104, Oregon State University, Corvallis, OR 97331-3803.
458
PELAGE COLOR IN PIÑON MICE
2002]
459
METHODS
For analyses we used 105 complete (skin
and skull) museum specimens of Peromyscus
truei collected at 43 sites in April–August
(summer) and 97 collected at 35 sites in September–March (winter). These groups include
specimens selected from within ranges of the
5 recognized subspecies of P. truei in the
northern Great Basin and environs and specimens unclassifed to subspecies from Oregon
and Idaho. All specimens are adults in full,
clean pelage free of visible grease and exhibiting no molting. Adults were identified by molars
being fully erupted and exhibiting wear, and
all skull sutures being completely fused. Collection locality (Appendix 1), date of collection, and pelage color were recorded for each
specimen.
Pelage color was measured on specimens
by comparing color at each of 7 points (Fig. 1)
with those of Munsell soil-color chips; hue,
value, and chroma were recorded for each
color (Munsell Color 1975). We used this
color-comparison technique in preference to
more precise electronic methods because necessary equipment was easily transported to
various systematics collections, cost less, and
had the potential of being used in the field.
The 3 hues found in P. truei (5YR, 7.5YR,
10YR) were coded 1, 2, and 3, respectively,
but actual numbers for color value and chroma
read from color chips were used in analyses
(Appendix 2). For example, for a specimen
with a measured color of 10YR3/2 at a point,
the data for that point appeared as 3 3 2.
Because values for midventer color (Fig. 1)
were the same for all specimens (10YR8/1),
this character was not used in statistical analyses; thus, each specimen was represented by
18 numerals.
Separate data matrices were formed for
specimens collected in summer and winter.
We analyzed each data matrix by use of a dissimilarity association matrix from Euclidean
distance with unweighted pair-group method
using arithmetic averages (UPGMA) method
within Hierarchical Cluster Analysis in BioΣtat
II (Pimentel 1994). Clusters were designated
A, B, C, D, and E, and collection localities for
specimens within each cluster were superimposed on a map of Hoffmeister’s (1951) distributions of nominal subspecies occurring in the
northern Great Basin and adjacent regions.
Fig. 1. Sketch of a Peromyscus truei illustrating points of
reference for determining pelage color: a, pate; b, anterior
dorsum; c, posterior dorsum; d, dorsal tail stripe; e, side
above lateral stripe; f, lateral stripe; and g, midventer.
RESULTS
All characters, other than midventer color,
exhibited some variation among both summerand winter-caught specimens (Table 1). Most
variation was in value and chroma, as the most
common hue recorded was yellowish brown
(10YR). Despite this variation, at sites at which
multiple specimens were available, all individuals grouped in the same cluster.
Analyses of data matrices for P. truei collected in summer resulted in 5 distinct clusters, whereas those collected in winter resulted
in only 4 clusters (Figs. 2a, 2b). Cophenetic
correlation coefficients (rcs) for these dendrograms were 0.967 and 0.963, respectively,
indicating a high level of agreement between
similarity values implied by dendrograms and
those of original similarity matrices.
Specimens forming cluster A were from
western Josephine County, Oregon, and coastal
California; those in cluster B were from Shasta
Valley, Siskiyou County, California; those in
cluster C were from eastern Nevada, western
Utah, southern Idaho, and Crook, Deschutes,
Jefferson, and Harney counties, Oregon; those
in cluster D were from Nevada; and those in
cluster E were from Jackson, eastern Josephine, Klamath, and Lake counties, Oregon (Fig.
3). Because geographic distributions of clusters A, C, D, and E for winter-caught specimens
were in accord with those of summer-caught
specimens, seasonal groups were combined
for plotting (Fig. 3). No winter-caught specimens from Siskiyou County, California, with
requirements for inclusion were available for
study, accounting for absence of a cluster B
produced by that analysis (Fig. 2b).
We also reanalyzed our matrices, eliminating
data for 1 point on the pelage at a time to
ascertain whether relationships among clusters
460
[Volume 62
TABLE 1. Munsell color determinations of pelage color of Peromyscus truei from the northern Great Basin and adjacent areas at 6 points of reference. Vertical listings of colors indicate variation possible within a cluster (geographic
region); thus, horizontal listings do not indicate colors for any particular individual.
Anterior
dorsum
Posterior
dorsum
Dorsal
tail
Lateral
side
Lateral
stripe
10YR2/1
10YR2/1
10YR7/4
10YR8/3
10YR8/6
10YR2/1
10YR2/1
10YR7/4
10YR8/3
10YR2/1
10YR2/1
10YR7/4
7.5YR6/6
10YR8/6
7.5YR3/2
7.5YR3/2
10YR3/2
5YR3/4
7.5YR3/4
7.5YR7/6
7.5YR6/6
10YR6/8
10YR7/4
10YR7/6
Winter
10YR8/6
10YR7/3
10YR8/3
10YR7/6
10YR8/6
Summer
7.5YR7/6
10YR8/3
10YR8/4
7.5YR7/6
10YR8/3
10YR8/4
7.5YR7/6
7.5YR8/4
10YR8/4
5YR3/4
7.5YR3/4
10YR2/1
10YR2/1
10YR3/2
7.5YR7/6
7.5YR7/6
10YR6/6
10YR7/4
10YR7/6
10YR8/4
10YR8/4
10YR8/6
7.5YR7/8
7.5YR7/6
10YR7/6
Winter
10YR7/4
10YR8/3
10YR8/4
10YR8/3
10YR8/4
Summer
10YR7/4
10YR7/4
7.5YR7/6
7.5YR8/4
10YR7/4
10YR8/4
7.5YR6/6
Winter
10YR7/4
10YR7/6
10YR8/6
10YR7/4
10YR7/6
7.5YR6/6
10YR7/6
7.5YR7/6
7.5YR8/6
10YR7/6
10YR8/4
10YR8/6
7.5YR7/6
10YR7/3
10YR7/6
10YR8/6
10YR6/8
10YR7/8
7.5YR7/6
7.5YR7/8
10YR6/8
Cluster
Season
Pate
A
Summer
Winter
Summer
Summer
B
C
D
E
were affected. Results were not altered greatly
by sequentially eliminating data points (Fig.
2). Such suggests that the set of variables was
correlated over the body.
DISCUSSION
Distribution of specimens in cluster A conforms most closely with that depicted for P. t.
sequoiensis at the northern extent of its range
(Fig. 3). The greater Euclidean distance separating cluster A from other clusters, combined
with the larger body size, shorter tail, and
shorter ears reported for specimens from the
region (Hoffmeister 1951), suggests that morphometric and genetic analyses might indicate
that sequoiensis is specifically different from
the remaining P. truei. Whether sequoiensis
and gilberti are sympatric or remain parapatric
in that region is unknown.
Based on analyses of matrices containing
all 6 characters, distribution of all specimens
forming cluster D (Fig. 3) conforms with that
depicted for P. t. truei (Hoffmeister 1951, Hall
1981). However, southwestern Utah contains
localities from which specimens group into
cluster C (Fig. 3). Such suggests that the zone
of intergradation between clusters C and D is
10YR2/1
10YR3/1
10YR3/1
7.5YR2/0
10YR3/2
10YR2/1
10YR3/1
10YR3/2
10YR7/6
10YR8/6
7.5YR6/6
10YR7/6
10YR6/8
7.5YR7/6
7.5YR7/8
10YR6/8
slightly farther west and south than depicted
for truei and nevadensis by Hoffmeister (1951).
Specimens forming cluster E were all collected at localities in Oregon: 3 in Josephine
County, 3 in Jackson County, 1 in Klamath
County, and 3 in Lake County. Those from
Josephine County suggest that the range of P.
t. gilberti extends farther west than previously
believed and those from Klamath and Lake
counties extend the range farther north and
eastward (Fig. 3). Hoffmeister (1951) and Hall
(1981) indicated that the range of P. t. truei
extended from Nevada into south central Oregon, encompassing localities at which the
Lake County specimens were collected. However, as specimens from that region were not
available to them, the depicted ranges for
these 2 races must have been based on conjecture. Winter-caught specimens forming cluster E separated into 2 clusters when matrices
were analysed after removal of color of pate,
anterior dorsum, or posterior dorsum (Figs. 2f,
2j, 2n). We consider the separation spurious
because both clusters so formed included individuals from more than 1 of the geographic
groups of specimens forming cluster E (Fig.
3). Several other species of small mammals
and races thereof associated primarily with
2002]
461
Fig. 2. Dendrograms with accompanying cophenetic correlation coefficients (rcs) produced from data matrices formed
for specimens of Peromyscus truei collected in summer and winter with: a–b, all characters included; c–d, with dorsal
tail color removed; e–f, with anterior dorsum color removed; g–h, with lateral stripe color removed; i–j, with posterior
dorsum color removed; k–l, with lateral side color removed; and m–n, with pate color removed.
habitats in either eastern or western Oregon
have ranges that extend through a gap in the
Cascade Range near the southern border of
that state (Verts and Carraway 1998). Thus,
classification of specimens of P. truei from Klamath and Lake counties as gilberti is not surprising.
Summer-caught specimens forming cluster
B grouped with those in cluster E in analyses
in which 1 of 3 color characters (dorsal tail,
posterior dorsum, or lateral side) was excluded
(Figs. 2c, 2i, 2k). However, when 1 of the other
3 characters (pate, anterior dorsum, or lateral
stripe) was excluded and when all 6 characters
were included, dendrograms indicated that
cluster E was distinct from cluster B and, that
other than cluster A, was the most divergent
group from the remaining clusters (Figs. 2e,
2g, 2m). Also, specimens in cluster B grouped
with those in cluster C when colors of either
the anterior dorsum or the lateral stripe were
excluded (Figs. 2e, 2g). One locality in Siskiyou County, California, from which specimens
forming cluster B were taken, was within the
range of P. t. gilberti depicted by Hoffmeister
(1951); 3 other localities were only slightly to
the east of the depicted range (Fig. 3). From
geographic proximity involved (Fig. 3) and
from combining of specimens in 3 dendrograms (Figs. 2c, 2i, 2k), specimens in cluster
B seem to be consubspecific with those in
cluster E. However, the Euclidean-distance
462
[Volume 62
Fig. 3. Distribution of localities of specimens of Peromyscus truei forming the 5 clusters overlain on a distribution map
of subspecies (stippled areas) following Hoffmeister (1951): 1, P. t. gilberti; 2, P. t. nevadensis; 3, P. t. preblei; 4, P. t.
sequoiensis; and 5, P. t. truei. Superscript numerals following cluster-letters indicate number of specimens from that
locality; absence of numeral indicates only 1 specimen. Dashed-line trapezoid in central Oregon contains 23 localities.
Solid square in Harney Co., Oregon, is locality at which KU 157813 was collected. Names of counties referred to in text
are abbreviated.
relationship between clusters B and E in 4
dendrograms (Figs. 2a, 2e, 2g, 2m) and the fact
that all individuals forming cluster B were
from a restricted area (Shasta Valley) suggest
that these groups are taxonomically distinct.
Color analysis combined with morphometric
and genetic analyses of additional specimens,
including some caught in winter, from central
Siskiyou County, California, might aid significantly in clarifying the relationship of this somewhat tenuous colormorph.
Except for 2 groups of localities in southwestern Utah and extreme eastern Nevada near
the periphery of the depicted range of P. t.
2002]
truei, the distribution of specimens forming
cluster C includes regions depicted by Hoffmeister (1951) and Hall (1981) as occupied by
P. t. preblei and P. t. nevadensis (Fig. 3). In addition, cluster C also includes specimens from
eastern Oregon (other than those from Lake
County) and southwestern Idaho collected north
and east of the previously described geographic
range of the species (Fig. 3). Distributions of P.
t. preblei and P. t. nevadensis (Hoffmeister 1951,
Hall 1981), combined with localities of more
recently collected specimens in cluster C, form
a band in a more-or-less continuous arc from
central Oregon, through southwestern Idaho,
and along the Nevada–Utah state boundary
(Fig. 3). Despite this logical grouping of 2
nominal subspecies, we are reluctant to synonymize these races in the absence of morphometric and genetic analyses.
Absence of available specimens from much
of southeastern Oregon and the northern portion of Nevada (Fig. 3; Hoffmeister 1951, Hall
1981) may not represent the true distribution
of the species. In August 1998 we collected a
specimen (KU 157813) near the center of this
broad region at Juniper Spring (T40S, R31E,
SE1/4 SE1/4 Sec. 12), Harney County, Oregon
(Fig. 3). Unfortunately, it was a juvenile and so
could not be included in the present analyses.
However, its occurrence in a region heretofore
considered not occupied by the species, combined with the extremely restricted habitat
occupied by the species (Carraway et al. 1993),
suggests that additional effort to acquire specimens from the region might be productive.
Scattered stands of either Pinus edulis or Juniperus, the appropriate vegetative habitats, occur
over much of the region (Johnson 1995). Specimens from this region might provide additional insight into the relationship between
those presently classified as P. t. preblei and P.
t. nevadensis.
Overall, distributions of nominal races of P.
truei in the northern Great Basin and environs
based on our analyses of pelage color tend to
support conclusions of previous researchers.
In instances in which previous conclusions
were not supported and for specimens from
areas outside the previously known range of
the species, distributions based on our color
analyses are logical and suggest avenues for
additional research on geographic variation in
the species.
463
ACKNOWLEDGMENTS
We thank curators and collection managers
at Museum of Natural History, Albertson College, Caldwell, Idaho (CIMNH); Eastern Oregon University, La Grande (EOSC); National
Museum of Natural History, Washington, D.C.
(USNM); Portland State University, Portland,
Oregon (PSU); Department of Biology, Boise
State University, Boise, Idaho (BSU); Department of Fisheries and Wildlife, Oregon State
University, Corvallis (OSUFW); Museum of
Vertebrate Zoology, University of California,
Berkeley (MVZ); Natural History Museum,
University of Kansas, Lawrence (KU); and
Burke Museum, University of Washington,
Seattle (UW) for loans or access to collections
they curate. N.A. Slade provided statistical
advice. L.J. Rielly aided in finding a particularly obscure locality. Earlier drafts of the manuscript were reviewed by J.L. Patton, T.L. Best,
and S.T. Alvarez-Castañeda. This is Technical
Paper 11,618, Oregon Agricultural Experiment
Station.
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BAILEY, V. 1936. The mammals and life zones of Oregon.
North American Fauna 55:1–416.
CAIRE, W., J.D. TYLER, B.P. GLASS, AND M.A. MARES. 1989.
Mammals of Oklahoma. University of Oklahoma
Press, Norman.
CARRAWAY, L.N., E. YENSEN, B.J. VERTS, AND L.F. ALEXANDER. 1993. Range extension and habitat of Peromyscus truei in eastern Oregon. Northwestern Naturalist 74:81–84.
CLARK, T.W., AND M.R. STROMBERG. 1987. Mammals in
Wyoming. University of Kansas Museum of Natural
History, Public Education Series 10:1–314.
DAVIS, W.B., AND D.J. SCHMIDLY. 1994. The mammals of
Texas. Texas Parks and Wildlife Press, Austin.
FISHER, J.L. 1976. An autecological study of the pinyon
mouse, Peromyscus truei, in southwestern Oregon.
Unpublished master’s thesis, General Studies–Science/Mathematics, Southern Oregon State College,
Ashland.
GAFUR, Z., D. HOYLE, K. KEECH, B. ROBERTS, L.R. POWERS,
AND E. YENSEN. 1980. Records of the pinyon mouse,
Peromyscus truei, from southwestern Idaho. Journal
of the Idaho Academy of Science 16:1–2.
HAFNER, M.S., W.L. GANNON, J. SALAZAR-BRAVO, AND S.T.
ALVAREZ-CASTAÑEDA. 1997. Mammal collections in
the Western Hemisphere: a survey and directory of
existing collections. American Society of Mammalogists, Lawrence, KS.
464
HALL, E.R. 1946. Mammals of Nevada. University of California Press, Berkeley.
______. 1981. The mammals of North America. 2nd edition.
John Wiley & Sons, New York. Volume 2:601–1181
+ 90.
HALL, E.R., AND D.F. HOFFMEISTER. 1940. The pinyon
mouse (Peromyscus truei) in Nevada, with description of a new subspecies. University of California
Publications in Zoology 42:401–406.
HOFFMEISTER, D.F. 1941. Two new subspecies of the
piñon mouse, Peromyscus truei, from California. Proceedings of the Biological Society of Washington 54:
129–132.
______. 1951. A taxonomic and evolutionary study of the
piñon mouse, Peromyscus truei. Illinois Biological
Monograph 21(4):1–104.
______. 1981. Peromyscus truei. Mammalian Species 161:
1–5.
JOHNSON, F.D. 1995. Wild trees of Idaho. University of
Idaho Press, Moscow.
MODI, W.S., AND M.R. LEE. 1984. Systematic implications
of chromosomal banding analyses of populations of
Peromyscus truei (Rodentia: Muridae). Proceedings
of the Biological Society of Washington 97:716–723.
MUNSELL COLOR. 1975. Munsell soil color charts. Kollmorgen Corporation, Baltimore, MD.
PIMENTEL, R.A. 1994. BioΣtat II: a multivariate statistical
toolbox. Version 3.5. Sigma Soft, San Luis Obispo, CA.
RAMÍREZ-PULIDO, J., A. CASTRO -CAMPILLO, J. ARROYO CABRALES, AND F.A. CERVANTES. 1996. Lista taxonómica de los mamiferos terrestres de México.
Occasional Papers, The Museum, Texas Tech University 158:1–62.
RIDGWAY, R. 1912. Color standards and color nomenclature. Privately published, Washington, DC. 53 plates.
SHUFELDT, R.W. 1885. Description of Hesperomys truei, a
new species belonging to the subfamily Murinae.
Proceedings of the United States National Museum
8:403–408.
VERTS, B.J., AND L.N. CARRAWAY. 1998. Land mammals of
Oregon. University of California Press, Berkeley.
APPENDIX 1. Specimens of Peromyscus truei examined. Type specimens indicated by an asterisk [*].
Localities followed by a plus [+] were not plotted
on Figure 3. Museum acronyms follow Hafner et al.
(1997).
U.S.A.—CALIFORNIA, Del Norte Co.: Gasquet
(USNM 91540). Siskiyou Co.: 1.5 mi SW Edgewood
(MVZ 69422, 69424–69426); 10 mi E Montague (MVZ
69413); 11 mi NE Weed (MVZ 69414–69416); Mayten
(MVZ 13333). IDAHO, Owyhee Co.: T10S, R5W, Sec.
22/27 (BSU 1352–1359); T4S, R3W, Sec. 26, 1500 m (BSU
1285–1286, 1288–1290, 1292, 1315–1317, 1319–1320,
[Volume 62
1323–1324, 1326; CIMNH 639, 790–798, 801); T9S, R5W,
Sec. 17 (BSU 1293–1294). NEVADA, Churchill Co.: 16
mi E Stillwater (USNM 93403–93404). Clark Co.:
Charleston Mts. (KU 75703; USNM 26888/34304, 205558);
Charleston Peak (USNM 26253/33659, 26268/33674, 26891/
34307, 26894/34310, 26895/34311, 26899/34315, 26922/
34338, 208980). Elko Co.: 0.5 mi W Debbs Cr., Pilot Peak
(MVZ 68468–68473, 68475–68479*, 68480–68482). Lander
Co.: Reese River, lat. 39 (USNM 93408). Lincoln Co.:
0.25 mi W UT–NV boundary+ (MVZ 59565); 3.5 mi N
Eagle Valley (MVZ 48770); SW base Groom Baldy, T5S,
R55/56E (MVZ 48753); E slope Irish Mts. (MVZ 48755,
48760, 48763); Meadow Valley (MVZ 48765); Panaca
(USNM 28409/40513); 11 mi E Panaca (MVZ 48767, 48769);
2 mi S Pioche (MVZ 59562–59563); 2 mi SE Pioche (MVZ
59561, 59564). Lyon Co.: 2 mi SW Pine Grove (MVZ
64408, 64412, 64428); 3.5 mi SW Pine Grove (MVZ
64410). Nye Co.: Grapevine Mts. (USNM 28696/40800).
Pershing Co.: S slope Granite Peak (MVZ 74174). Washoe
Co.: 3 mi E Reno (MVZ 71072). White Pine Co.: Cheery
Creek (MVZ 46226); Cleve Creek, T16N, R65E, NE1/4
(MVZ 46224); 8 mi N Lund (MVZ 53152–53159); Overland Pass, E slope Ruby Mts. (KU 47146); W side Ruby
Lake (KU 47143–47145). OREGON, Crook Co.: at mouth
Bear Creek (MVZ 87730–87732); mouth Bear Creek on
Crooked River (MVZ 87727–87728); 13.5 mi S, 2.5 mi W
Prineville (OSUFW 1635); 12 mi S, 2.5 mi E Prineville
(UW 20029–20031). Deschutes Co.: 14 mi S, 10 mi E Bend
(OSUFW 2753); Crooked River, 20 mi SE Prineville (USNM
78660*); 2 mi N, 4 mi E Redmond (OSUFW 2778); 3 mi
N, 1 mi W Redmond (OSUFW 2232); 2 mi S, 1 mi E Redmond (OSUFW 2652); 3 mi N, 3 mi E Sisters (OSUFW
1508); 3 mi N, 4 mi E Sisters (OSUFW 2207); 9 mi N, 6.5
mi E Sisters (OSUFW 2651); 10 mi N, 4 mi W Sisters
(OSUFW 2231); 2 mi S, 5 mi E Sisters (OSUFW 2887);
0.25 mi N, 4 mi W Terrebonne (OSUFW 5508); 1 mi N, 1
mi W Terrebonne (OSUFW 3084); 1 mi N, 3 mi W Terrebonne (OSUFW 3035); 1 mi S, 3 mi W Terrebonne
(OSUFW 2989); 2 mi S, 8 mi W Terrebonne (OSUFW
2187); 3 mi W Terrebonne (OSUFW 2888, 2911, 3016,
3072, 3248, 3477, 3523, 4185–4192, 5154, 7070, 7896,
7971, 7982, 8095); 4 mi W Terrebonne (OSUFW 3432,
3862–3864, 5451). Harney Co.: 9 mi NE Riley (BSU
1304–1306); T24S, R25E, N1/2 Sec. 3 (KU 145222); T30/
31S, R35E, Sec. 35/2 (KU 145215–145217, 145218). Jackson Co.: Brownsboro (USNM 262546); 7 mi S, 15 mi W
Medford (OSUFW 3479); T38S, R2E, Sec. 22 (PSU 2717).
Jefferson Co.: 10 mi S Madras (PSU 1256); 10 mi W
Metolius (OSUFW 5144–5145, 5197). Josephine Co.: 13 mi
SW Galice (USNM 203581–203584, 203587); Grants Pass
(USNM 17276/24205, 32135/43947, 32137/43949, 203653);
2 mi N, 1 mi W Grants Pass (OSUFW 2478, 2487, PSU
2468); T39S, R7W, Sec. 34 (PSU 2716, 2718–2719). Klamath
Co.: 0.7 mi N, 0.7 mi E Keno (PSU 2721). Lake Co.: Oatman Flat, 8 mi NW Silver Lake (PSU 1888); Oatman Flat,
7 mi N Silver Lake (PSU 882); Oatman Flat, 7 mi N, 1 mi
W Silver Lake (PSU 680). UTAH, Millard Co.: Wah Wah
Mts. (USNM 356962–356963). Washington Co.: 11 mi N
St. George (USNM 327174, 327181); Mt. Meadows (USNM
167535–167537, 167542); Pine Valley (USNM 166769,
166776).
2002]
465
APPENDIX 2. Hue, value, and chroma with Munsell Color (1975) names for colors found in Peromyscus
truei in the northern Great Basin and environs.
10YR2/1
10YR6/6
10YR6/8
10YR7/3
10YR7/4
10YR7/6
10YR7/8
10YR8/1
10YR8/3
10YR8/4
10YR8/6
Black
Brownish yellow
Brownish yellow
Very pale brown
Very pale brown
Yellow
Yellow
White
Very pale brown
Very pale brown
Yellow
7.5YR3/2
7.5YR3/4
7.5YR6/6
7.5YR7/6
7.5YR7/8
7.5YR8/4
7.5YR8/6
7.5YR2/0
Dark brown
Dark brown
Reddish yellow
Reddish yellow
Reddish yellow
Pink
Reddish yellow
Black
5YR3/4
Dark reddish brown
A HETEROTROPHIC DESERT STREAM?
THE ROLE OF SEDIMENT STABILITY
Urs Uehlinger1, Markus Naegeli2,3, and Stuart G. Fisher2
ABSTRACT.—In autumn 1998 stream metabolism was measured in the Hassayampa River, Arizona, a Sonoran Desert
stream, using single-station diel oxygen curves and an oxygen mass balance model. Oxygen consumption rates of parafluvial and channel sediments were determined with respiration chambers. Bedload of channel sediments (sand) prevented significant primary production by benthic autotrophs, despite favorable nutrient, light, and temperature conditions. Ecosystem respiration was relatively low (1.50 g O2 m–2d–1) and presumably fueled by production in the riparian
zone and riverine marshes. Respiration rates in the parafluvial zone and in channel sediments ranged from 0.6 to 1.4 g
O2 m–3 sediment h–1. Sediment organic matter (ash-free dry mass) was 4.0 ± 1.8 kg m–3 sediment and did not significantly differ between the channel and the parafluvial zone. Results indicate that heterotrophic processes may dominate
the metabolism of desert stream segments over extended periods of time if unstable sandy bed sediments prevail.
Key words: desert stream, stream metabolism, diel oxygen curve, sediment respiration, disturbance, sediment stability.
Environmental conditions in streams of warm
arid or semiarid climates favor high primary
production rates that may exceed ecosystem
respiration (Minshall 1978, Cushing and Wolf
1984, Grimm 1987, Lamberti and Steinman
1997, Sinsabaugh 1997). Respiration in streams
mainly occurs in hyporheic and parafluvial
sediments (Grimm and Fisher 1984, Mulholland et al. 1997, Naegeli and Uehlinger 1997).
Jones et al. (1995) found that in Sycamore
Creek, a warm desert stream, benthic production fueled more than 80% of the hyporheic
respiration through leaching dissolved material. Input of allochthonous organic matter is of
minor importance if a riparian zone is narrow
and upland vegetation is sparse as in arid environments (Cushing 1997, Schade and Fisher
1997). However, river corridors may include
floodplain marshes, shrubs, and trees (e.g.,
Prach et al. 1996, Stromberg et al. 1997) that
provide organic matter through aerial deposition of plant litter, fine root production or release of organic matter by roots, or processing
of particulate organic matter buried during
floods (Robertson et al. 1999).
The stability of bed sediments is an important determinant of the biologically mediated
energy flow through lotic ecosystems. Bed-
moving spates damage benthic primary producers but have only a minor influence on the
heterotrophic community of the hyporheic
zone (Grimm and Fisher 1984, Naegeli and Uehlinger 1997). These disturbances shift stream
metabolism toward heterotrophy (Fisher et al.
1982, Uehlinger and Naegeli 1998, Uehlinger
2000). Flash floods in desert streams decimate
primary producers, but recovery can be rapid
due to favorable light and temperature conditions and stable substrata after flood recession
(Fisher et al. 1982, Grimm and Fisher 1989).
This study focuses on the metabolism of a
warm desert stream, where light availability,
temperature, and nutrient concentrations are
ample to support high primary production rates.
However, bed sediments consisted of sand,
the superficial layer of which was in continuous motion even at low flow. We hypothesized
that the metabolism of this desert stream would
be dominated by respiration because of this
persistent bedload. Further, we expected that
metabolism rates in the wetted channel would
be smaller than in the parafluvial zone because
low in-stream primary production was thought
to result in low organic matter supply compared to inputs from the riparian vegetation
and riverine marshes to the parafluvial zone.
1Department of Limnology, Swiss Federal Institute for Environmental Science and Technology (EAWAG), CH-8600 Dübendorf, Switzerland.
2Department of Biology, Arizona State University, Tempe, AZ 85287-1501, USA.
3Present address: XPERTEAM Management Consultants AG, Glattalstrasse 501, CH-8153 Ruemlang, Switzerland.
466
HETEROTROPHY OF A DESERT STREAM
2002]
METHODS
Study Reach
The Hassayampa River drains a catchment
of about 2000 km2 into the Gila River (central
Arizona). Perennial, intermittent, and ephemeral
reaches characterize the river. The study was
conducted downstream of Wickenburg (Maricopa County, AZ, USA), where a shallow bedrock layer causes perennial surface flow for
about 8 km. Sand dominates channel sediments
in the bedrock-confined reach (Stromberg et al.
1993), which is an upwelling zone of alluvialand basin-filled groundwater ( Jenkins 1989).
The study reach is 1 km downstream of The
Nature Conservancy’s Hassayampa River Preserve, close to the end of the perennial river
section. Elevation is about 610 masl, and channel slope averages 0.006 (m/m). The reach is
relatively homogenous with respect to channel
structure (pool-riffle-run frequency), depth,
slope, and riparian vegetation. At the Hassayampa River Preserve, mean annual discharge
averages 0.1 m3s–1 ( Jenkins 1989). Effluent
from a municipal wastewater treatment plant
upstream of the study area makes up 0.01 to
0.02 m3s–1 of the flow. Large floods (Qmax >
500 m3s–1), which change channel area and instream riparian habitat structure, occur predominantly in winter (Stromberg et al. 1997).
Emergent macrophytes, such as Typha domingensis Pers., is occasionally dominant within
the river channel and abundant in adjacent
marshlands (Wolden et al. 1994). Baccharis salicifolia [Ruíz et Pavón] Pers. occurs along stream
margins; more distal riparian vegetation is dominated by cottonwood (Populus fremonti Wats.)
and willow (Salix goodnigii Ball). Floodplain
width ranges from 100 to 200 m. Adjacent uplands consist of Sonoran Desert scrub vegetation (Stromberg et al. 1993). The study reach is
unreplicated; therefore, only tentative conclusions about ecosystem metabolism and sediment
respiration of desert streams may be drawn.
Physics and Chemistry
Channel width and depth were measured
along a 260-m reach. In the wetted channel
and the parafluvial zone of a 75-m sub-reach,
we installed 15 permanent piezometers with
sampling depths of 10 cm (6), 30 cm (6), and
80 cm (3) for sampling interstitial water. A
small side channel longitudinally divided the
parafluvial zone.
467
Interstitial water temperatures were measured using 2 temperature loggers (StowAway,
Onset Corporation, North Falmouth, MA, USA)
that were installed 20 cm below the sedimentwater interface in the wetted channel and 20
cm below the water table in the parafluvial
zone. Surface water temperatures were continuously recorded with a combination temperature-oxygen probe (see below). We measured
discharge with the slug injection method
(Gordon et al. 1992), using NaCl as tracer and
a conductivity meter (WTW LF 340, Wissenschaftlich-Technische Werkstätten GmbH, Weilheim, Germany) as detector. Discharge records
of the Hassayampa River at the Morristown
gage (about 6 km downstream of the study site)
were recorded by the USGS (http://waterdata.
usgs.gov/nwis-w/AZ/).
Surface and interstitial water samples for
chemical analyses were collected in triplicate
on 28 October and 1 November 1998 (surface
water was also sampled on 27 October 1998).
Samples were stored on ice, filtered in the
laboratory (Whatman GF/F glass fiber filters),
and analyzed for nitrate (NO3) and ammonium
(NH4) nitrogen, soluble reactive phosphorus
(SRP), and dissolved organic carbon (DOC).
Analyses were performed as described by
Holmes et al. (1998). Dissolved oxygen (O2) and
temperature of subsurface water were measured in the field with an oxygen meter (ATI
Orion model 830).
Metabolism
We measured sediment respiration as
changes in O2 concentration over time in
sealed Plexiglas cores (4.5-cm diameter, 32 cm
long), filled with sediments from 10 or 30 cm
depth and water from the respective depths,
and sealed with rubber stoppers (after Jones et
al. 1995). Cores were incubated in situ for 2 to
3 hours. Oxygen concentrations and temperatures were measured with an oxygen meter
(ATI Orion model 830). After incubation, sediments were analyzed for particulate organic
matter as ash-free dry mass (loss on ignition)
and grain-size distribution. Sampling sites
were located in the main channel and in the
parafluvial zone.
To assess gross primary production and ecosystem respiration in the channel, oxygen concentrations and temperatures were continuously measured at the end of the study reach
468
from 27 October to 1 November 1998 with
Orbisphere equipment (probe 2115 with O2meter 2607 of Orbisphere, Geneva, Switzerland). Temperature and O2 signals were averaged for 5 minutes and stored (datalogger LI1000, LI-COR Inc., Lincoln, NE, USA). We
used an oxygen mass-balance model to calculate gross primary production GPP (g
O2m–2d–1) and ecosystem respiration ER (g
O2m–2d–1). A detailed description of the
model is given by Uehlinger et al. (2000).
Model simulations and parameter estimations
were performed with the computer program
AQUASIM (version 2.0; Reichert, 1994, 1995).
Respiration per volume r (g O2m–3d–1) was
parameterized as:
ER
r = – ____
z
(1)
where ER is the respiration rate per area (g
O2m–2d–1) and z the mean water depth (m).
Gross primary production was described as a
linear function of incident light (I):
GPP = pIz
(2)
where p is the slope of the linear P–I curve (g
O2m–1d–1W–1) and I is incident light intensity
(Wm–2). Linear or almost linear relationships
between ecosystem primary production and
light intensitiy are usually obtained with open
system methods (e.g., Duffer and Dorris 1966,
Hornberger et al. 1976, Uehlinger 1993, Uehlinger et al. 2000). We described temperature
dependence of ER and GPP with the following
Arrhenius equation:
X(T)=X(20ºC)βx(T–20)
(3)
where the subscript X is p or r and βX is a constant ( βX > 1; Bowie et al. 1985). Saturation
concentrations of O2 at the study site were
calculated using water temperature and the
barometric pressure recorded at Wickenburg
Airport (about 6 km from the study site). We
determined the reaeration coefficient Ks with
propane as tracer gas (Genereux and Hemond
1992). Temperature dependence of Ks was
described according to Elmore and West
(1961). Estimates of the parameters r, p, and
βX were based on the minimization of the sum
of squares (SS):
[Volume 62
n
SS = Σ (O2(ti )–O2meas,i )2
i=1
(4)
where O2meas,i is the O2 concentration measured at the end of the reach at the time ti,
O2(ti ) is the O2 concentration at the end of the
reach at the time ti is calculated with the
model, and n is the number of observations.
To judge the identifiability for model parameters, we calculated the collinearity index (γ ),
which is a measure for the degree of approximate linear dependence of sensibility functions (Brun et al. 2001). Critical values of γ are
in the range of 5 to 20. To evaluate the relative
importance of ER, GPP, and the influence of
temperature on both processes, we performed
model runs by activating and deactivating primary production, respiration, and temperature
dependence of both processes (i.e., by setting
β = 1 or p or r = 0).
To assess differences in sediment respiration rates and sediment organic matter, we
used 2-way ANOVA. Prior to the analysis, respiration data were transformed (log(x + 1)).
Temperature dependence of sediment respiration was evaluated using linear regression
analysis. Differences or regressions were considered significant when P < 0.05.
RESULTS
Discharge measured on 3 occasions averaged 0.094 ± 0.007 m3s–1. At this flow rate,
width and depth of the wetted channel averaged 6.6 m and 0.034 m, respectively, and
mean current velocity was 0.41 ms–1. Between
0900 and 1200 hours, the main channel stage
decreased by about 1 cm (presumably due to
diel transpiration cycles of the riparian vegetation). As a consequence, the upper part of the
small side channel fell dry each day during the
investigation. A transient flow increase due to
a rainstorm during the night of 31 October/
1 November 1998 (flood marks indicated a
stage increase by 3–5 cm) scoured some of the
algal patches located along the margins of the
wetted channel.
Main channel surface water temperature
varied between 12.4°C and 26.7°C (Table 1,
Fig. 1). The high diel temperature variation of
the interstitial water in the sediments of the
wetted channel (∆T = 9.5°C) indicated a substantial water exchange between sediment and
surface water. Temperature in the parafluvial
2002]
469
TABLE 1. Temperature of surface water and interstitial water (°C) in the Hassayamapa River. Temperatures were
recorded in 10-minute intervals.
Period
Minimum
Maximum
Average
Surface water
Sediment below
the main channela
Saturated sediments of
the parafluvial zoneb
27–31 Oct
12.4
26.7
17.4
27–30 Oct
14.1
23.6
17.6
30 Oct–2 Nov
13.0
15.3
14.3
a25 cm below the sediment water interface
b25 cm below the water table
zone averaged only 14.3°C, and diel variations
were small (∆T = 2.3°C). This pattern suggests minor water exchange between surface
water and parafluvial sediments.
Relatively high concentrations of nitrate
and SRP characterize the surface and interstitial water of the main channel (Table 2). Vertical concentration gradients of the interstitial
water were small or absent, and even at 80 cm
depth O2 was surprisingly high. In the parafluvial zone, O2 and nitrate concentrations were
lower than in the channel sediments, and SRP
increased with depth. We found higher DOC
concentrations in parafluvial sediments than in
channel sediments. Ammonium concentrations
were relatively high and uniform at all sites.
Differences in grain-size distribution between the channel and the parafluvial zone
were small (Fig. 2). Apart from a narrow zone
along the channel margins, surface bed sediments (D90 about 2 mm) were in continuous
motion (based on average channel characteristics such as width, depth, slope, and D90, the
discharge threshold for initiation of sediment
transport, was estimated to be 0.078 m3s–1;
Günther 1971). Sediment movement occurred
between the beginning of October 1998 (1st
visit to the site) and the end of the study in
mid-November 1998, and also when we returned to the reach in February 1999. This
bedload restricted habitats suitable for algae
to small patches along the channel margins;
we observed that more than 95% of the wetted
channel area was free of algal patches visible
by eye. Site (main channel, parafluvial zone)
and depth had no significant effect on sediment organic matter, which averaged 4.0 ± 1.8
kg m–3 sediment (Table 3).
Site and depth significantly influenced sediment respiration. Average respiration rates
decreased by about 40% from the parafluvial
zone to the main channel and by 30% from 10
Fig. 1. Top: Surface water temperature (bold line) in the
main channel of the Hassayampa River and global radiation (fine line) at Wickenburg airport. Bottom: Dissolved
oxygen concentration in the surface water of the main
channel.
to 30 cm depths (Table 3). During incubation,
average temperatures in the sediment tubes
ranged from 15.8°C to 24.5°C in the channel
and from 12.5°C to 22.7°C in the parafluvial
zone. However, the regression between the
respiration rates and temperature was significant only for parafluvial sediments (R2 =
0.198).
Oxygen concentrations in the surface water
were always below saturation concentration
and showed distinct diel variation (∆O2 = 2.3 ±
3 mg O2 L–1) with daily minimum in the early
afternoon (between 1300 and 1500 hours; Fig.
1). The small spate of 30 October resulted in a
distinct but transient increase in O2. The reaeration coefficient Ks (20°C) for O2 determined with the gas tracer method was 89 ± 18
d–1. Table 4 summarizes the results of a series of
calculations with models of different complexity
(number of active processes). The value of the
collinearity index (γ < 5) points out that the 4
470
[Volume 62
TABLE 2. Concentrations of oxygen, major nutrients, and DOC on 28 October and 1 November 1998 between 1100 and
1400 hours (channel surface water was also sampled on 27 October 1998).
Main channel
Surface water
Sediments (depth 10 cma)
Parafluvial zone
Sediments (depth 10 cmb)
O2
(mg L–1)
NH4-N
(µg L–1)
NO3-N
(µg L–1)
SRP
(µg L–1)
DOC
(mg L–1)
7.7 ± 0.4
6.8 ± 1.1
7.1 ± 0.7
7.3 ± 0.4
28 ± 15
20 ± 11
26 ± 23
26 ± 19
230 ± 15
278 ± 32
296 ± 20
302 ± 9
53 ± 1
48 ± 7
42 ± 14
51 ± <1
1.67 ± 0.12
1.61 ± 0.12
1.53 ± 0.07
1.52 ± 0.05
3.3 ± 3.6
2.8 ± 2.1
1.0 ± 0.3
23 ± 9
31 ± 14
30 ± 16
165 ± 139
288 ± 343
171 ± 164
58 ± 20
63 ± 15
73 ± 16
2.71 ± 1.58
2.30 ± 0.88
1.83 ± 0.39
aBelow the sediment-water interface
bBelow the water table
was the most parsimonious fit and led to good
agreement between calculated and measured
oxygen data. Estimates of daily gross primary
production and ecosystem respiration based
on fits 5, 6, and 7 are given in Table 5. We
assume that the values provided by fits 5 and
6 most probably delimited the range of gross
primary production and ecosystem respiration
during the investigation.
DISCUSSION
Fig. 2. Grain-size distribution of main channel sediments (10 and 30 cm below the bed surface) and parafluvial sediments (10 and 30 cm below the water table) in the
Hassayampa River.
model parameters (ER, p, βr, βp) can be identified. The sum of squares (SS) of the basic
model, which considered only gas exchange,
was 145.9. Respiration had the largest influence
on SS through all levels of model complexity
and indicates the importance of this process
for oxygen balance. In contrast, primary production had no (fits 1, 3, 4) or only a small
effect (fits 6, 7). Temperature dependence of
gross primary production and ecosystem respiration reduced SS to 17.0. However, we did
not consider this model in the estimation of
metabolism rates because βp became unrealistically high (a value βp = 1.2311 means an 8fold increase in gross primary production if
temperature rises from 10°C to 20°C). Fit 5
As expected, heterotrophic processes dominated the metabolism of this desert stream.
The permanent motion of the uppermost bed
sediments in most parts of the wetted channel
prevented substantial periphyton accrual and
thus any substantial primary production. Apart
from sediment stability, environmental conditions were favorable for the growth of benthic
algae; the river was open canopied, the water
was clear, and concentrations of major nutrient were high. Algal patches were restricted to
areas with stable substrata such as channel
margins, backwater areas, and parafluvial sediments inundated for only a few hours a day.
However, the contribution of these stable areas
to the wetted channel area was small (< 5%).
The metabolism of the Hassayampa River
was heterotrophic during this study; estimates
of P/R ranged from 0 to 0.10 depending on the
model applied (fits 5 and 6) to calculate metabolism rates. Ecosystem respiration rates were
relatively small (1.33–1.50 g O2 m–2d–1) compared to rates reported from other desert
streams (3.6–6.5 O2 m–2d–1; Grimm 1987, Cushing and Wolf 1984). Algae can provide significant quantities of dissolved and particulate
2002]
471
TABLE 3. Sediment respiration rates and sediment organic matter (mean and standard deviation) in the Hassayampa
River measured on 29 and 31 October, and 10 November 1998. N = number of samples.
Location
Depth
(cm)
Respiration rate
(g O2 m–3 sediment h–1)
Organic matter
(kg m–3 sediment)
N
10a
30a
10b
30b
0.96 ± 0.11
0.55 ± 0.31
1.39 ± 0.82
0.78 ± 0.31
3.8 ± 1.8
4.4 ± 1.9
4.2 ± 1.9
4.4 ± 1.2
22
9
21
6
Main channel
Parafluvial zone
aBelow the sediment-water interface
bBelow the water table
TABLE 4. Evaluation of the processes influencing the oxygen balance in the Hassayampa River. Reaeration, Ks(20°C),
was measured (see text). The other parameters were fitted to the models defined by equations 1, 2, and 3. Standard
error estimates are in parentheses; empty cells mark the parameters not included in a fit.
Number of
parameters
fitted
Ks(20°C)
(d–1)
—
1
2
0
1
1
89
89
89
3
4
5
2
2
2
89
89
89
6
3
89
7
4
89
Fit no.
ER
(g O2 m–2d–1)
P
(g O2 W–1d–1)
βr
βp
2.90 10–7 a
1.290
(0.021)
1.295a
1.476
(0.022)
1.707
(0.038)
1.958
(0.049)
2.77.10–7 a
0.00a
9.04 10–4
(1.27 10–4)
9.72 10–4
(1.30 10–4)
1.00001a
1.0502
(0.0034)
1.0698
(0.0046)
1.1087
(0.0060)
1.2311
(0.0216)
SS
γ
145.9
145.9
25.8
—
1.00
1.00
145.9
25.8
20.3
4.04
1.82
1.53
19.0
3.19
17.0
4.47
aStandard error could not be estimated.
organic matter supporting respiration in the
hyporheic zone (Jones et al. 1995). We suggest
that the lack of a substantial algal community
may account for the low ecosystem respiration
rates, and we hypothesize that organic matter
released by macrophytes of the riverine marshes
and the riparian vegetation, in addition to organic matter buried during floods, may have
fueled sediment respiration. Organic matter
from riverine marshlands and the riparian vegetation bordering the investigated parafluvial
zone may have accounted for the differences
in respiration rates between parafluvial and
channel sediments; the differences in DOC
concentrations between the parafluvial zone
and channel sediments may reflect different
organic matter sources. Sediment organic matter in the Hassayampa River averaged 4.0 kg
m–3 sediment in the uppermost 30 cm, which
is in the range of values reported from Sycamore Creek (2.8–5.7 kg m–3 sediment in the
top 15 cm; Valett et al. 1990). This material is
presumably of suboptimal quality, but it is an
energy reservoir that may increase the resistance of stream metabolism to short-term
environmental fluctuations. Sediment organic
matter was homogeneously distributed in the
uppermost 30 cm of the sediments (channel
and saturated parafluvial zone), but respiration
rates significantly decreased from 10 to 30 cm
depth. This may reflect the increasing refractory nature of the sediment organic matter
with depth.
One prediction of the river continuum concept is that changes in the relative importance
of primary production and respiration for
energy flow through lotic ecosystems can be
attributed to changes in stream size and riparian vegetation (Vannote et al. 1980); for example, environmental settings in arid or semiarid
regions were found to favor high rates of primary production exceeding ecosystem respiration (Minshall 1978, Lamberti and Steinman
1997, Sinsabaugh 1997). However, this study
and recent investigations indicate that the stability of bed sediments has to be considered in
472
TABLE 5. Gross primary production, ecosystem respiration, and P/R (average and standard deviation) in the Hassayampa River, 28–31 October 1998. Daily rates (g O2
m–2d–1) were calculated with 3 models of different complexity (fits 5, 6, and 7; Table 4).
Model (fit)
5
6
7
Gross primary
production
Ecosystem
respiration
P/R
0.00
0.15 ± 0.03
0.29 ± 0.10
1.33 ± 0.05
1.50 ± 0.07
1.65 ± 0.13
0.00
0.10 ± 0.01
0.17 ± 0.05
predictions on the relative importance of stream
metabolism (Uehlinger and Naegeli 1998, Uehlinger 2000). During high flow, bed sediments
may become unstable, which mainly affects
benthic algae and primary production (Fisher
et al. 1982, Uehlinger and Naegeli 1998). But
the effects of such events are usually transient
because spates are limited in time and primary production recovers more or less rapidly
depending on environmental conditions (Fisher
et al. 1982, Uehlinger and Naegeli 1998, Uehlinger 2000). At the Morristown gage (about 6
km downstream of our study site), spates were
recorded on 29 March 1998 (Qmax = 21 m3s–1)
and on 12 August 1998 (Qmax ≈ 2 m3s–1). During the spate-free period between August 1998
and the beginning of our measurements, permanently moving bed sediments apparently
prevented periphyton accrual in most parts of
the channel; i.e., sediment instability seems to
account for a persistent dominance of heterotrophic processes in the metabolism of this
warm desert stream. Streams with sandy, unstable sediments can be found throughout the
semiarid Southwest of North America, though
we do not know the extent to which they comprise a significant proportion of southwestern
river habitat. Conditions of high bedload may
be restricted to certain segments or transient
(e.g., during the wet season). In the Hassayampa River, for example, the study reach represented a particular successional stage of a
desert river corridor. Large floods dramatically
change prevailing substrata and the spatial
extent of the wetted channel, parafluvial zones,
riverine marshes, floodplain forests, and shrublands within the river corridor (Stromberg et
al. 1997). Such flood-induced alterations in the
configuration of the river corridor may affect
stream metabolism.
Studies of stream metabolism in the arid
West of North America indicate that desert
[Volume 62
streams are autotrophic ecosystems (Minshall
1978, Cushing and Wolf 1984, Grimm 1987).
The results of this investigation suggest that
heterotrophic processes may dominate the
metabolism of desert stream segments over
extended periods of time if unstable sandy bed
sediments prevail. However, a test of this hypothesis needs reliable estimates of annual
metabolism rates. Such estimates require measurements at least during one annual cycle
and with a temporal resolution that accounts
for the hydrological disturbance regime of these
systems.
ACKNOWLEDGEMENTS
We thank Cathy Kochert for analyzing chemical parameters, Jennifer Zachary for POM
and grain-size distribution analysis, and the
stream group at ASU for their valuable support. We appreciate the critical comments of
Nancy Grimm, James Ward, and 2 anonymous
reviewers.
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Accepted 6 November 2001
A SEED CHALCID (EURYTOMA SQUAMOSA BUGBEE) PARASITIZES
BUCKBRUSH (CEANOTHUS FENDLERI GRAY) SEEDS
IN A PONDEROSA PINE FOREST OF ARIZONA
David W. Huffman1
ABSTRACT.—Predispersal seed parasitism rates were quantified for buckbrush (Ceanothus fendleri Gray) in 3 Arizona
ponderosa pine (Pinus ponderosa Laws.) forest units that had been thinned for ecological restoration objectives. The
chalcidoid wasp Eurytoma squamosa Bugbee (Eurytomidae) was responsible for 35% of total seed loss in a single year.
These findings represent an expansion of the known range and host list for E. squamosa and increase our understanding
of factors that may constrain regeneration of C. fendleri in Southwest ponderosa pine forests.
Key words: predispersal seed predation, range expansion, Hymenoptera, Eurytoma squamosa, Ceanothus fendleri,
ecological restoration, plant-insect interactions, parasitism.
In ponderosa pine (Pinus ponderosa Laws.)
forests of the Southwest, ecological restoration
treatments have been initiated that include
thinning dense stands of young trees and applying low-intensity fire (Covington and Moore
1994, Covington et al. 1997, Moore et al. 1999).
Although reestablishment of ecological function is theorized to flow from overstory structural manipulation and the reintroduction of
surface fire, ecological interactions, particularly between arthropods and understory plant
hosts, have been minimally researched.
In 1999 I began a study to examine various
factors affecting regeneration of Ceanothus
fendleri Gray in areas undergoing ecological
restoration treatments. Ceanothus fendleri is a
shrub common in the understory of ponderosa
pine forests in the Southwest. It is a preferred
browse of mule deer (Odocoileus hemionus
hemionus Rafinesque; Urness et al. 1975), a
nitrogen fixer (Story 1974), and a provider
of structural heterogeneity in predominantly
grassy and herbaceous understory communities. Resprouting of C. fendleri after disturbances such as fire is common (Pearson et al.
1972, Ffolliott et al. 1977, Vose and White
1991). Its seeds are forcibly ejected from dehiscing capsules and likely remain in forest
floor seed banks for years until stimulated by
heat from fire to germinate (Kearney and Peebles 1951, Reed 1974, Krishnan 1989). Large
ungulate use of understory forage can increase
after forest thinning (Patton 1974), and herbivory on C. fendleri can be intense and constrain flower and fruit production (Huffman
and Moore in preparation). Other factors that
likely affect C. fendleri regeneration and vary
with forest condition include seed predation
and parasitism.
Prior to this study, rates of seed parasitism
and predation had not been reported for C.
fendleri, although nearly complete predispersal
consumption of seed crops by invertebrates
has been reported for overstory ponderosa
pine (Blake et al. 1985). The objectives of my
research were to (1) collect preliminary data
on predispersal seed parasitism for Ceanothus
fendleri in areas thinned for ecological restoration objectives and (2) identify common insect
species infesting C. fendleri seeds. Information
concerning processes that affect ovule and
seed fate can help resource managers predict
rates of ecological recovery in communities
undergoing restoration treatments.
METHODS
The study was conducted from 1999 to
2000 on the Fort Valley Experimental Forest
(latitude 35°16′N, longitude 111°41′W) in
Coconino County approximately 10 km northwest of Flagstaff, Arizona. The site is approximately 2300 m above mean sea level and has
flat to gently rolling topography with slopes
1Ecological Restoration Institute, Northern Arizona University, Flagstaff, AZ 86011-5017.
474
2002]
SEED CHALCID PARASITIZES BUCKBRUSH
generally less than 20%. Annual precipitation
averages 52 cm, of which approximately half
falls as snow in late winter. Soils are moderately well drained and classified as Brolliar
stony clay loam (Meurisse 1971) developed
from tertiary basalt parent material.
Overstory vegetation is nearly pure ponderosa pine with scattered occurrence of
Gambel oak (Quercus gambelii Nutt.). Common understory species include bunchgrasses
Festuca arizonica Vasey, Muhlenbergia montana
(Nutt.) A.S. Hitchc., Elymus elymoides (Raf.)
Swezey, and Blepharoneuron tricholepis (Torr.)
Nash; forbs Lupinus spp., Erigergon spp., Erigonium spp., Achillea millifolium L., and Antennaria spp.; and woody shrubs Ceanothus fendleri and Rosa woodsii Lindl.
Large mammalian herbivores present on
the site are mule deer and elk (Cervus elaphus
Linnaeus). Livestock were excluded from the
study site.
Overstory trees were thinned in February
1999, which reduced tree density in three 15-ha
experimental forest restoration units to approximately 111–120 trees per hectare. In May
1999, I established 60 plots (4 m2 in size) centered on Ceanothus fendleri shrubs in each of
the 3 restoration units (180 shrub plots total).
Ceanothus fendleri plants on these plots produced no fruit in 1999. Therefore, I collected
seeds from C. fendleri shrubs growing on
microsites adjacent to the restoration units and
monitored parasite emergence from these seeds
until June 2001. Adult parasites emerging were
captured and immediately preserved in 70%
isopropyl alcohol. Specimens were sent to Dr.
Robert Zuparko at the California Academy of
Science (CAS), San Francisco, California, USA,
and to the USDA Systematic Entomology Laboratory (SEL; specimens identified by E. Eric
Grissell, Research Entomologist), Bethesda,
Maryland, USA, for identification.
Fruit developed on 11 total C. fendleri plots
in July 2000. To capture seeds as fruits dehisced,
I installed seed traps constructed from bridal
veil material (mesh size <2 mm) tied around
fruiting stems. Twenty-three traps were installed
on 1 to 5 stems per plot and each enclosed 1 to
14 fruits.
Seeds collected from traps were separated
from plant debris and counted in the laboratory. Seeds were classified as “developed” or
“undeveloped.” Developed seeds were approximately 2 mm in diameter, had smooth, full
475
seed coats, and were glossy brown in appearance (Fig. 1a). In contrast, undeveloped seeds
were typically smaller than 2 mm and flattened; additionally, they had wrinkled, yellowish seed coats (Fig. 1b). Seeds were examined
under a dissecting scope (10–20X) for parasite
emergence holes or other signs of infestation
and then dissected to determine embryo condition and presence of parasite larvae/pupae.
RESULTS
Parasite specimens collected as they emerged
from 1999 seeds were all from a single chalcidoid wasp species, Eurytoma squamosa Bugbee (Eurytomidae; Figs. 1c–d). Although emergence was not specifically studied, I observed
adult wasps emerging up to 20 months after
the 1999 seed collection.
A total of 144 seeds were recovered in 2000
from traps installed on C. fendleri fruiting
stems. Each trap yielded 1 to 24 seeds. Fifty
percent of the seeds captured were undeveloped (Table 1), and in these seeds no parasite
larvae or signs of infestation were found.
Parasitism was responsible for 35% of the
total seed loss and 71% of the loss of developed seeds (Table 1). No clear trends were observed for rate of parasitism and number of
seeds per trap or seeds per plot. Parasite emergence holes in developed seeds were approximately 0.5 mm in diameter (Fig. 1e). Embryonic
tissue of seeds with emergence holes was completely consumed, leaving the seeds hollow.
Seeds housing parasite larvae showed no
external signs of infestation. Emergence hole
appearance (i.e., size and shape) was consistent between seeds trapped in 2000 and those
collected in 1999. Likewise, characteristics of
larvae within dissected seeds of both collections were consistent. We found no evidence
of other parasitic species associated with C.
fendleri seeds.
The majority of developed seeds that had
not been parasitized were filled with apparently healthy embryos (Fig. 1f, Table 1). However, approximately 3% of these showed no signs
of parasitism but were hollow nonetheless.
DISCUSSION
These results represent an expansion of the
known range and host list for Eurytoma squamosa. Prior to this study, this wasp species was
not known to occur in Arizona or on Ceanothus
476
[Volume 62
Fig. 1. Images (10–20X) showing (a) normally developed seeds of Ceanothus fendleri, (b) undeveloped seeds, (c) adult
parasite Eurytoma squamosa, (d) emerging seed parasite, (e) characteristic emergence holes made by seed parasite, and
(f) cross-section of developed seed with healthy embryo.
fendleri. Occurrence of the species has been
reported in Idaho, Washington, and California, USA (Bugbee 1967). In these states its
hosts are other species of Ceanothus including
C. divaricatus, C. thyraiflorus, C. cordulatus,
C. velutinus, and C. sanguineus. For C. sanguineus in Idaho, E. squamosa and 2 other
phytophagous insects were responsible for an
average loss of 9–27% of total seeds in fruits
over a 3-year period (Furniss et al. 1978).
Another Eurytoma species (E. greggii Bugbee)
has been reported to infest >80% of the seeds
of Ceanothus greggii Gray in chaparral ecosystems of Arizona (Bugbee 1971). Seed chalcids
are well-known parasites of seeds of commercial tree species such as ponderosa pine in
SEED CHALCID PARASITIZES BUCKBRUSH
2002]
477
TABLE 1. Fate of seeds collected from Ceanothus fendleri plants in ponderosa pine forests of northern Arizona.
Developed
_____________________________________________
a
Parasitized
____________________
Seeds
Undeveloped
Total
Number
Percentage of developed
seeds
Emerged
37
Larval
14
Filled
19
Hollow
2
72
144
51.4
19.4
26.4
2.7
—
100
PERCENTAGE OF TOTAL
25.7
9.7
13.2
1.3
50
100
aParasites (Eurytoma squamosa) either had emerged prior to seed collection or were found within seeds as larvae or pupae.
northern Arizona (e.g., Blake et al. 1985), but
little research has quantified parasitism on
understory plants.
With the data presented here, long-term
effects of seed parasitism on Ceanothus fendleri regeneration are difficult to determine.
Traits such as innate dormancy suggest that C.
fendleri utilizes a seed bank strategy similar to
congeneric species (Quick 1935, Reed 1974,
Conard et al. 1985, Krishnan 1989). However,
resprouting after disturbance is also common
(Ffolliott et al. 1977, Vose and White 1991).
Thus, although my data suggest that E. squamosa constrained replenishment of seeds in
soil seed banks, temporal patterns of seed parasitism and the relative importance of sprouting versus seedling recruitment for C. fendleri
persistence are not yet clear.
Presettlement characteristics of C. fendleri–
E. squamosa interactions are not known. Plausibly, frequent surface fires common in ponderosa pine forests prior to Anglo and Hispanic
settlement of northern Arizona (ca 1876; Fulé
et al. 1997) could have functioned to control
larval populations of E. squamosa in dispersed
seeds. Fires heating C. fendleri seeds in soil
seed banks may have caused wasp mortality
while stimulating seed germination. Thus, future
research could address questions related to
fire or heat effects on parasite and seed survival and seed germination. Studies illuminating presettlement conditions with respect to
plant–insect interactions could help ecologists
and land managers expand their understanding of ecosystem dynamics and, in turn, make
informed decisions concerning forest restoration and management.
ACKNOWLEDGMENTS
I thank Dr. Margaret M. Moore and Amy
Waltz for insightful comments on an earlier
draft of this manuscript and W. Walker Chancellor for field assistance. This project was
funded by USFS Rocky Mountain Research
Station–Northern Arizona University School
of Forestry, Research Joint Venture Agreement
RMRS-99167-RJVA.
LITERATURE CITED
BLAKE, E.A., M.R. WAGNER, AND T.W. KOERBER. 1985.
Insects destructive to ponderosa pine cone crops in
northern Arizona. Conifer Tree Seed in the Inland
Mountain West Symposium, 5–6 August 1985, Missoula, MT.
BUGBEE, R.E. 1967. Revision of chalcid wasps of genus
Eurytoma in America north of Mexico. Proceedings
of the United States National Museum 118:433–552.
______. 1971. A new species of Arizona Eurytoma phytophagous in Ceanothus greggii seeds. Journal of the
Kansas Entomological Society 44:111–112.
CONARD, S.G., A.E. JARAMILLO, K. CROMACK, JR., AND S.
ROSE. 1985. The role of the genus Ceanothus in
western forest ecosystems. USDA Forest Service
General Technical Report PNW-182.
COVINGTON, W.W., AND M.M. MOORE. 1994. Southwestern
ponderosa forest structure: changes since Euro-American settlement. Journal of Forestry 92(1):39–47.
COVINGTON, W.W., P.Z. FULÉ, M.M. MOORE, S.C. HART,
T.E. KOLB, J.N. MAST, S.S. SACKETT, AND M.R. WAGNER. 1997. Restoring ecosystem health in ponderosa
pine forests of the Southwest. Journal of Forestry
95(4):23–29.
FFOLLIOTT, P.F., W.P. CLARY, AND F.R LARSON. 1977. Effects
of a prescribed fire in an Arizona ponderosa pine
forest. USDA Forest Service Research Note RM-336.
FULÉ, P.Z., W.W. COVINGTON, AND M.M. MOORE. 1997.
Determining reference conditions for ecosystem
management of southwestern ponderosa pine forests.
Ecological Applications 7:895–908.
FURNISS, M.M., T.A. LEEGE, AND R.J. NASKALI. 1978. Insects
that reduce redstem Ceanothus seed production in
Idaho. In: Proceedings of the First International
Rangeland Congress.
KEARNEY, T.H., AND R.H. PEEBLES. 1951. Arizona flora.
University of California Press, Los Angeles.
KRISHNAN, S. 1989. Propagation of four native drought tolerant shrubs—Ceanothus spp. and Sherpherdia spp.
Master’s thesis, Colorado State University, Fort
Collins.
478
MEURISSE, R.T. 1971. Soil report on the San Francisco
Peaks area, Elden and Flagstaff Ranger Districts,
Coconino National Forests. USDA Forest Service
(unpublished).
MOORE, M.M., W.W. COVINGTON, AND P.Z. FULÉ. 1999.
Reference conditions and ecological restoration: a
southwestern ponderosa pine perspective. Ecological Applications 9:1266–1277.
PATTON, D.R. 1974. Patch cutting increases deer and elk
use of a pine forest in Arizona. Journal of Forestry
72:764–766.
PEARSON, H.A., J.R. DAVIS, AND G.H. SCHUBERT. 1972.
Effects of wildfire on timber and forage production in
Arizona. Journal of Range Management 25:250–253.
QUICK, C.R. 1935. Notes on the germination of Ceanothus
seeds. Madroño 3:345–346.
REED, M.J. 1974. Ceanothus L. ceanothus. In: C.S. Schopmeyer, technical coordinator, Seeds of woody plants
in the United States. USDA Handbook 450.
[Volume 62
STORY, M.T. 1974. Nitrogen fixation by Ceanothus fendleri
and Lupinus argenteus as a function of parent material and vegetal cover. Master’s thesis, University of
Arizona, Tucson.
URNESS, P.J., D.J. NEFF, AND R.K. WATKINS. 1975. Nutritive value of mule deer forages on ponderosa pine
summer range in Arizona. USDA Forest Service
Research Note RM-304, Rocky Mountain Forest and
Range Experiment Station, Fort Collins, CO.
VOSE, J.M., AND A.S. WHITE. 1991. Biomass response
mechanisms of understory species the first year after
prescribed burning in an Arizona ponderosa-pine
community. Forest Ecology and Management 40:
175–187.
Received 7 February 2001
NEST SITE CHARACTERISTICS AND REPRODUCTIVE SUCCESS
OF THE WESTERN TANAGER (PIRANGA LUDOVICIANA)
ON THE COLORADO FRONT RANGE
Karen N. Fischer1, John W. Prather1,2, and Alexander Cruz1
ABSTRACT.—From 1999 through 2001 we located and monitored Western Tanager (Piranga ludoviciana) nests in public open-space properties in Boulder County, Colorado. Fifty-four of 58 nests were located in ponderosa pine and the
remainder in Douglas-fir. Nests were generally placed near the midpoint of branches in areas of high canopy cover
(>50%) in the middle section of nest trees. Nest height varied as a function of nest tree height, and nests were oriented
randomly in relation to trunks of nest trees. Tanager nesting success varied annually, with estimates using the Mayfield
method ranging from 11.3% in 2000 to 75.3% in 2001. At least 8 nests were predated, and predation was the primary
cause of nest failure. Parasitism by Brown-headed Cowbirds (Molothrus ater) occurred in 7 of 17 (41%) nests found during egg-laying or incubation. Clutch size averaged 3.8 in 10 unparasitized nests, but only 2.4 in 8 parasitized nests.
Brood parasitism dramatically reduced the number of tanager fledglings produced per nest.
Key words: Western Tanager, Piranga ludoviciana, breeding biology, nest site selection, Colorado.
The Western Tanager (Piranga ludoviciana),
a neotropical migrant, is widespread throughout western portions of the United States and
Canada. Breeding occurs in open, mixed coniferous-deciduous forests from the southeastern
tip of Alaska to the Trans-Pecos region of Texas
(Hudon 1999) and east as far as western South
Dakota (Peterson 1995). In Colorado the Western Tanager occurs primarily from mid-May
until mid-September in montane portions of
the state (Andrews and Righter 1992, Versaw
1998).
Western Tanager nests are difficult to locate
and monitor since they are often high and well
hidden in conifers. Few studies have examined ecological factors that influence reproductive success in the Western Tanager, and
most of these are anecdotal or unpublished. To
the best of our knowledge, a study by Hudon
(1999) in Alberta (n = 7) and Goguen and
Mathews (1998) in New Mexico (n = 39) are
the only ones that have examined in detail the
breeding biology and nest site characteristics
of this species. In addition, Project Tanager,
conducted by the Cornell Laboratory of Ornithology, examined the effects of habitat fragmentation on tanager species breeding in
North America (Rosenberg et al. 1999), but it
has limited information on breeding biology or
microhabitat characteristics of nest sites ( Jim
Lowe personal communication). From 1998
through 2001 we collected data on the breeding biology of neotropical migratory birds on
open-space properties in Boulder County, Colorado. We present here information on the nest
site selection and reproductive success of the
Western Tanager.
STUDY AREAS
Nests were located on open-space properties maintained by the city and county of
Boulder at elevations of 1600–1900 m in the
foothills west of Boulder, Colorado (40°0′N,
105°16′W). On these properties we searched
for nests every 2–3 days on 10 plots ranging
from 6 to 24 ha in size. These plots are dominated by ponderosa pine (Pinus ponderosa)
woodland and savannah with a mixture of
Douglas-fir (Pseudotsuga menziesii) at higher
elevations. Riparian corridors border some
plots, although we located no nests in riparian
vegetation. Neither did we locate nests in residential areas, though once again such areas
bordered on some plots. All of our plots were
within a few kilometers of the city of Boulder
and were subject to varying degrees of human
disturbance.
1Environmental, Population, and Organismic Biology Department, University of Colorado, Boulder, CO 80309-0334.
2Corresponding author. Present address: Center for Environmental Science and Education, Northern Arizona University, Flagstaff, AZ 86011-5694.
479
480
METHODS
We monitored nests at intervals of 2–4 days
(normally every 3 days) following standard
nest-monitoring protocols (Ralph et al. 1993),
until they were no longer active. Attempts
were made to limit nest failure due to factors
associated with nest monitoring (Martin and
Geupel 1993). We determined nest contents
by direct observation or by using a 6-m mirror
pole whenever possible, but some nests were
too high for contents to be monitored safely.
For nests that we were unable to directly
determine the contents, we relied on adult behavior and nest condition to ascertain whether
the nest was active, predated, or abandoned. A
search for fledglings was conducted in the
vicinity of every nest in which chicks were
presumed to have fledged. All nests that were
confirmed as being active, either by monitoring the contents or by observing adults sitting
on and/or visiting the nest with food, were
included in this study.
After nests were no longer active, we measured habitat characteristics at each site using
standardized protocols ( James and Shugart
1970, Martin and Roper 1988). For each nest
we measured nest tree height, nest tree diameter (dbh), canopy cover over the nest, distance of nest to trunk and to tip of supporting
branch, and height of the lowest living branch
on the nest tree. All distances were measured
with a measuring tape whenever possible.
However, it was usually necessary to measure
heights of tall trees and higher nests using a
Suunto PM-5/360 PC clinometer. Canopy cover
was measured by averaging 4 measurements
taken with a Lemmon model-A convex spherical densiometer at a distance of 1 m from the
nest in the 4 cardinal directions (Lemmon
1957). Additionally, we measured slope and
aspect of the terrain around the nest site using
a compass and the clinometer. Finally, we documented the location of each nest using a
Garmin GPS-12 global positioning system.
We calculated nest success following the
method proposed by Mayfield (1975), incorporating modifications suggested by Manolis et
al. (2000). For nests of unknown fate, we used
the last day the nest was observed to be active
as the last active date; for nests of known fate,
we used the midpoint between the last active
day and the day the nest was observed to be
empty or destroyed. We used estimates of the
[Volume 62
egg-laying/incubation period (13 days) and nestling period (12 days) from a subsample of our
own nests from which we could obtain this
data (see below).
Shapiro-Wilks W-tests were used to determine whether Western Tanager nests were
normally distributed in their placement in
relation to measured habitat variables. Linear
regression and/or Student’s t tests also were
used to search for patterns in nest placement
in relation to measured habitat variables. All
aforementioned tests were carried out using
JMP statistical software (SAS Institute 1995).
In addition, we used a Rayleigh test (Zar 1999)
to determine if there was a significant directional component to nest orientation in relation to the trunk of the nest tree. Means and
standard deviations are provided for numerical data whenever applicable.
RESULTS
We located 17 nests in both 1999 and 2000
and 24 nests in 2001, for a total of 58 nests.
Half of these nests came from 2 plots on which
we located 75% or more of the active tanager
nests in each season; the remainder were located
in less rigorously searched areas. Using nests
with known dates of laying and fledging, and
those for which dates could be extrapolated
from available data, we determined that the
breeding season extended from 28 May through
29 July. Active nests were observed from 2 June
through 29 July. The peak of the breeding season (at least 50% of nests active) occurred between 6 June and 1 July. We have no evidence
of 2nd broods.
We found 54 of 58 nests in ponderosa pine
and the remaining 4 in Douglas-fir. In general,
tanager nests were well hidden along midpoints
of branches in the middle portion of nest trees.
Canopy cover was high at all nest sites, averaging 71%, and never being less than 31%.
This distribution was slightly skewed from
normal, with more nests at lower canopy coverage (W = 0.95, P = 0.055). Nests were placed
an average of 63% of the distance out toward
the tip of the branch in a distribution that was
skewed toward the end of the branch (W =
0.95, P = 0.054). Nests were distributed normally in relation to the height of the nest tree
(W = 0.97, P = 0.378), with the mean of the
distribution at 54% of the height of the tree.
Tree height was a good predictor of nest height
2002]
WESTERN TANAGER REPRODUCTIVE SUCCESS
(F = 27.96, R2 = 0.34, P < 0.001). We found
no significant patterns of nest placement in
relation to the other habitat variables that we
measured. Nest orientation in relation to trunk
of the nest tree was not significantly different
from uniform (Rayleigh’s R = 10.84, z = 1.99,
P = 0.16), although there was a slight bias in
number of nests oriented toward the south and
east. Twenty-nine nests (50%) were oriented
between 80° and 200°.
The egg-laying and incubation period lasted
13.0 ± 2 days (range 11–15, n = 3). Nestlings
remained in the nest for 12.2 ± 1.5 days after
the initial egg hatched (range 11–14, n = 4).
One additional nest that fledged young in only
8 days may have fledged only a cowbird. A
few late nests may have been 2nd nesting
attempts due to a previous failure.
Mean clutch size for nests not parasitized
by Brown-headed Cowbirds (Molothrus ater)
was 3.8 ± 0.4 (n = 10). Eight of 18 (44%) nests
found during egg-laying or incubation were
known to be parasitized by cowbirds. These
nests contained an average of 2.4 ± 0.9 tanager
eggs and 1.1 ± 0.4 cowbird eggs (n = 8). Of 38
nests with known outcome, 28 (74%) successfully fledged at least 1 young (Table 1). Mean
number of fledglings produced by all nests
with known outcome and for which actual
numbers of fledglings could be determined
was 1.0 ± 1.3 (n = 28). This number is less
than the number of nests with known outcome
since we could not accurately determine how
many young fledged from 10 successful nests.
Mean number of fledglings produced by successful nests that were not parasitized was 2.7
± 0.7 (n = 15), while parasitized nests fledged
only 0.7 ± 1.2 tanager young (n = 3). Mayfield
nest success was calculated to be 51.8% for all
years combined, but it varied dramatically
between years (Table 1). Mayfield nest success
was lower than apparent nest success due to
the large number of nests located after hatching of the eggs.
DISCUSSION
In Colorado the breeding season for Western Tanagers appears to begin in late May and
end in early August. We found tanager nests
with eggs as early as 2 June, with an estimated
1st date of laying of 28 May. Active nests were
present on our study sites through at least 29
481
July. Records of active nests from the Colorado Breeding Bird Atlas range from 1 June
through 7 August (Versaw 1998). Initiation of
the breeding season occurs between mid-May
and mid-June in most areas (Hudon 1999).
We found only a few patterns in the placement of nests in relation to habitat characteristics. Tanagers did not appear to preferentially
orient their nests in relation to the trunk of the
nest tree. There was a slight directional component to nest placement (mean orientation
132°), but this was not significantly different
from uniform. In general, tanager nests were
located on large branches near the middle of
nest trees. All but 1 nest were placed 30–90%
of the distance from the trunk of the tree to
the tip of the branch, in locations with high
overhead canopy. Unlike other studies, which
have generally reported tanagers as nesting very
near tips of branches (Hudon 1999), we rarely
found nests in such sites. We believe this may
be because high canopy cover is important to
Western Tanagers. In ponderosa pines, which
often have a distinctly conical shape, nests
placed at the tips of branches would normally
be very exposed. Therefore, nests may be placed
somewhat closer to the trunk of ponderosa
pines than they would be in other trees. High
canopy cover (77 ± 12%) was also reported over
tanager nests in New Mexico (Hudon 1999).
In Colorado, as in other areas (Hudon 1999),
Western Tanagers appear to lay 4 eggs normally. We have data on incubation and nestling periods for very few nests since the majority of nests were located after young had
hatched. However, these data also support
conclusions from other studies. Our nests followed through incubation had eggs hatch in
11–13 days, as compared to other studies that
report 13–14 days (Versaw 1998, Hudon 1999).
Additionally, the mean nestling period for
nests in our study was 12.2 days, which is close
to the mean of 11.3 days reported by Hudon
(1999). Nesting success was high in 1999 and
2001 but low in 2000 due to a high rate of predation (Table 1). Nests normally fledged fewer
young than eggs laid.
Because so few nests have been located
during egg-laying and incubation, few studies
have reported rates or effects of cowbird parasitism on breeding Western Tanagers. Prior to
this study there were only a few records of
cowbird parasitism on tanager nests in Colorado (Chace and Cruz 1996, Versaw 1998).
482
[Volume 62
TABLE 1. Clutch size and reproductive success of Western Tanagers breeding around Boulder, Colorado, 1999–2001.
Year
1999
2000
2001
Total
No. nests
No. successful
No. predated
No. abandoned
No. unknown
No. parasitizeda
17
7 (41%)
1 (6%)
1 (6%)
8 (47%)
1 (17%)
17
7 (41%)
5 (29%)
0
5 (29%)
2 (40%)
24
14 (58%)
1 (4%)
1 (4%)
8 (33%)
5 (71%)
58
28 (48%)
7 (12%)
2 (05%)
21 (36%)
8 (44%)
3.8 (n = 5)
4.0 (n = 3)
3.5 (n = 2)
3.8 (n = 10)
97
69.9 ± 15.1%
89
11.3 ± 24.5%
189
76.5 ± 10.3%
375
51.8 ± 15.8%
Clutch size
Exposure days
Mayfield success
aOf 18 nests located before the eggs hatched (6 in 1999, 5 in 2000, 7 in 2001).
However, in New Mexico cowbirds parasitized
33 of 39 nests (85%) over the course of a 4-year
study (Goguen and Mathews 1998). In contrast,
Chace et al. (2000) reported 1 of 9 nests parasitized (11%) in Arizona, and parasitism was
recorded in only 2 of 39 nests (5%) in British
Colombia (Hudon 1999). In our study we found
parasitism in 8 of 18 nests (44%) located before
the eggs hatched (Table 1). Parasitism varied
considerably between years; 5 of 7 nests found
with eggs were parasitized in 2001, while only
3 of 11 nests found with eggs were parasitized
in 1999–2000 (Table 1). Three additional parasitized nests were found after egg-hatching in
2001, while only 1 parasitized nest was found
after egg-hatching in 1999–2000. Additional
nests may have been parasitized, but we were
unable to differentiate between cowbird and
tanager young in many cases because of the
difficulty of directly observing these nests.
Cowbirds had a strong negative effect on
tanager breeding success. Parasitized nests held
an average of 1.2 fewer tanager eggs than unparasitized nests, suggesting cowbirds removed
tanager eggs from the nests. In addition, we
could confirm that tanager young fledged from
parasitized nests in only 2 cases. In one of these
cases, the cowbird egg failed to hatch, and the
nest fledged 3 tanager young. In the other, 2
tanager nestlings survived to fledge along with
1 cowbird nestling. In at least 3 cases, parasitized nests fledged only cowbird young, with
the tanager young either dying or disappearing from the nest. As in our study, parasitized
nests in New Mexico contained fewer eggs
(2.4) than unparasitized nests (4.0) and fledged
lower numbers of young (0.9 fledglings per
nest) than unparasitized nests (2.5 fledglings
per nest) (Goguen and Mathews 1998).
ACKNOWLEDGMENTS
Funding for this study was obtained from
the City of Boulder Parks and Recreation
Department, the Boulder County Parks and
Open Space Department, and the Undergraduate Research Opportunities program at the
University of Colorado. We thank Jameson
Chace, John Walsh, Heather Swanson, and all
of our field assistants who took data on nests
and nest sites.
LITERATURE CITED
ANDREWS, R., AND R. RIGHTER. 1992. Colorado birds: a
reference to their distribution and habitat. Denver
Museum of Natural History, Denver, CO.
CHACE, J.F., AND A. CRUZ. 1996. Knowledge of the Colorado host relations of the parasitic Brown-headed
Cowbird (Molothrus ater). Journal of the Colorado
Field Ornithologists 30:67–81.
CHACE, J.F., S.T. MCKINNEY, AND A. CRUZ. 2000. Nest-site
characteristics and nesting success of the Greater
Pewee in Arizona. Southwestern Naturalist 45:
169–175.
GOGUEN, C.B., AND N.E. MATHEWS. 1998. Songbird species
composition and nesting success in grazed and ungrazed pinyon-juniper woodlands. Journal of Wildlife
Management 62:474–484.
HUDON, J. 1999. Western Tanager (Piranga ludoviciana).
In: A. Poole and F. Gill, editors, The birds of North
America. Volume 432. American Ornithologists’
Union, Washington, DC, and The National Academy
of Science, Philadelphia, PA.
JAMES, F.C., AND H.H. SHUGART, JR. 1970. A quantitative
method of habitat description. Audubon Field Notes
24:727–736.
LEMMON, P.E. 1957. A new instrument for measuring forest
overstory density. Journal of Forestry 55:667–668.
MANOLIS, J.C., D.E. ANDERSEN, AND F.J. CUTHBERT. 2000.
Uncertain nest fates in songbird studies and variation in Mayfield estimation. Auk 117:615–626.
MARTIN, T.E., AND G.R. GEUPEL. 1993. Nest-monitoring
plots: methods for locating nests and monitoring
success. Journal of Field Ornithology 64:507–519.
2002]
WESTERN TANAGER REPRODUCTIVE SUCCESS
MARTIN, T.E., AND J.J. ROPER. 1988. Nest predation and
nest site selection of a western population of the
Hermit Thrush. Condor 90:51–57.
MAYFIELD, H.F. 1975. Suggestions for calculating nest
success. Wilson Bulletin 87:456–461.
PETERSON, R.A. 1995. The South Dakota breeding bird
atlas. South Dakota Ornithological Union, Aberdeen.
RALPH, C.J., G.R. GEUPEL, P. PYLE, T.E. MARTIN, AND D.F.
DESANTE. 1993. Handbook of field methods for monitoring landbirds. General Technical Report PSWGTR-144, Pacific Southwest Research Station, U.S.
Forest Service, Albany, CA.
ROSENBERG, K., J. LOWE, AND A. DHONDT. 1999. Effects
of forest fragmentation on breeding tanagers: a
483
continental perspective. Conservation Biology 13:
568–583.
SAS INSTITUTE. 1995. JMP users guide. Version 3.1. SAS
Institute, Cary, NC.
VERSAW, A. 1998. Western Tanager. Pages 442–443 in H.
Kingery, editor, The Colorado breeding bird atlas.
Colorado Bird Atlas Partnership, Denver, CO.
ZAR, J.H. 1999. Biostatistical analysis. 4th edition. PrenticeHall Inc., Englewood Cliffs, NJ.
Received 7 February 2001
Accepted 4 December 2001
CAPNIA CARYI, AN INTERESTING NEW SPECIES OF
WINTER STONEFLY FROM THE AMERICAN SOUTHWEST
(PLECOPTERA: CAPNIIDAE)
R.W. Baumann1 and G.Z. Jacobi2
ABSTRACT.—Capnia caryi, a new species of Nearctic Capniidae, is described from adults collected from high-elevation locations in the Southern Rocky Mountains of southern New Mexico and Arizona. Males are distinguished by their
rounded club-shaped epiproct and sclerotized knobs on abdominal terga 8 and 9. Females possess a darkly sclerotized
subgenital plate that covers most of tergum 8 and is produced posteriorly as a pair of broadly rounded lobes.
Key words: Plecoptera, stonefly, Capniidae, Capnia caryi, American Southwest.
Very dry years have characterized the American Southwest in the late 1900s and early part
of the 21st century. The reduced amount of
snowfall during these years has enabled collectors to gain access to streams at high elevations that were otherwise usually inaccessible.
In 1999 the mountains in the Mogollon Rim
Complex of the Gila Wilderness Area received
only 5% of the normal winter snowpack (P.
Stewart, U.S. Forest Service, personal communication).
Winter stoneflies have been the object of
increased collecting efforts since the formation of the western chapter of the Winter Stonefly Club in the 1980s as part of a revision of
the genus Capnia (Nelson and Baumann 1989).
Previous studies of winter stoneflies in New
Mexico include Jacobi and Baumann (1983),
Jacobi and Cary (1986, 1996). Because this interest has continued in many areas of the West,
with the collection of new material one can
easily compare specimens with the named
species. When these different specimens from
the Southwest were found in the winters of
1999 and 2001 at high-elevation locations in
Arizona and New Mexico, it was determined
that they represented a previously undescribed
Capnia species.
Capnia caryi, new species
MALE.—Wings brachypterous. Length of
forewings 3.5–4.0 mm; length of body 5.0–6.0
mm. Body and appendages dark brown; wings
slightly fumose. Eighth abdominal tergum with
small, rounded medial knob, located just before
posterior margin (Figs. 1, 2). Ninth tergum bearing large, darkly sclerotized, wedge-shaped
hump on posterior margin; apex wide, thin, and
broadly rounded (Figs. 1, 2). Tenth tergum
darkly sclerotized except for round, median,
light area where epiproct rests (Fig. 2). Area
under base of epiproct also light and membranous (Figs. 1, 2). Epiproct recurved, extending
forward to middle of tergum 9, very narrow at
base, tapering abruptly to widest expanse near
apex, producing overall rounded club shape,
especially in lateral view, apex pointed and
bearing tiny, blunt hook (Fig. 1).
FEMALE.—Wings macropterous. Length of
forewings 5.0–6.0 mm; length of body 6.0–7.0
mm. Coloration similar to male, with broad,
membranous dorsal stripe extending to posterior
margin of tergum 8. Subgenital plate restricted
to sternum 8; entire segment darkly sclerotized;
anterior margin with V-shaped indentation
medially; posterior margin very darkly sclerotized, expanded into broadly rounded, bilobed
margin, with median indented area (Fig. 3).
DIAGNOSIS.—Capnia caryi does not fit easily into any of the species groups proposed by
Nelson and Baumann (1989). It is most similar
to species in the Mariposa group because the
large dorsal hump of the male is on tergum 9.
However, even though it looks similar to Capnia giulianii Nelson and Baumann in lateral
view, the hump is not divided in C. caryi as in
1Department of Integrative Biology and Monte L. Bean Life Science Museum, Brigham Young University, Provo, UT 84602.
22314 Calle Colibri, Sante Fe, NM 87505, and New Mexico Highlands University (retired), Las Vegas, NM 87710.
484
NEW SPECIES OF WINTER STONEFLY
2002]
485
1
2
3
Capnia caryi: Fig. 1. Adult male terminalia lateral, epiproct. Fig. 2. Adult male terminalia dorsal, epiproct. Fig. 3. Adult
female terminalia ventral, subgenital plate.
486
all 3 species of the Mariposa group. Thus, the
presence of a large, undivided hump on tergum 9 of the male, along with a small medial
knob on tergum 8, serves to separate this species
from all other known Capnia males. In addition, the female subgenital plate is distinctive
in that it has a broadly rounded posterior lobe
on each side of the midline. Otherwise, the
presence of a large, darkly sclerotized subgenital plate could place it in any of several species
groups.
TYPES.—Holotype male, allotype female,
and 7 male and 3 female paratypes, NEW MEXICO, Catron Co., Upper Iron Creek, Forest
Trail 151, 2650 m, 32°22.67′N, 108°34.09′W,
22 February 1999, G.Z. Jacobi and S.J. Cary.
PARATYPES: ARIZONA, Apache Co., Mamie
Creek, Forest Road 275, 2400 m, Escudilla
Mountain, southeast of Springerville, 19 March
2001, G.Z. Jacobi and S.J. Cary, 2 males. Holotype and allotype deposited at the United States
National Museum, Washington, D.C.; paratypes
at Monte L. Bean Life Science Museum, Brigham Young University, and personal collection
of G.Z. Jacobi.
ETYMOLOGY.—We are pleased to name this
species after our friend and colleague Steve
Cary. He was present when all type specimens
were collected and has been a major participant in surveys of the stonefly fauna of New
Mexico.
NOTES.—The 2 creeks in which this new
species was collected are located near the border between southern Arizona and New Mexico. Iron Creek is in New Mexico and is a
small tributary of the Middle Fork of the Gila
River, southeast of Reserve in the Gila National
Forest (Julyan 1998). Mamie Creek lies in Arizona at the base of Escudilla Peak, southeast
of Springerville, and is part of the Little Colorado River drainage in the Apache National
Forest. In Iron Creek, Capnia confusa Claassen
[Volume 62
was emerging, and nymphs of Skwala americana (Klapalek), Nemouridae, and Chloroperlidae were collected. In addition to C. caryi, a
male of Capnia decepta (Banks) was collected
at the same time in Mamie Creek. Stream flow
measurements by the New Mexico Environmental Department in 1996 showed Iron
Creek to be low in flow in mid-July (0.1 m3),
with a similarly estimated flow in February
1999. Surface stream flow ceased 100 m upstream of the collection location in February
1999. The substrate consisted of scattered
boulders and a mixture of cobble with gravels;
gradient was 3%. The water was clear (<2
NTU) and cool (<10°C), with low amounts of
dissolved materials (total phosphorus <0.01
mg ⋅ L–1 and total CaCo3 alkalinity of <20 mg
⋅ L–1) and low non-filterable residues (<3.0 mg
⋅ L–1).
ACKNOWLEDGMENTS
Nicole Cox and Randy Baker prepared the
illustrations.
LITERATURE CITED
JACOBI, G.Z., AND R.W. BAUMANN. 1983. Winter stoneflies
(Plecoptera) of New Mexico. Great Basin Naturalist
43:585–591.
JACOBI, G.Z., AND S.J. CARY. 1986. New records of winter
stoneflies (Plecoptera) from southwestern New Mexico, with notes on habitat preferences and zoogeographical origins. Southwestern Naturalist 31:503–510.
______. 1996. Winter stoneflies in seasonal habitats in New
Mexico, USA. Journal of the North American Benthological Society 15:690–699.
JULYAN, R. 1998. The place names of New Mexico. University of New Mexico Press, Albuquerque. 385 pp.
NELSON, C.R., AND R.W. BAUMANN. 1989. Systematics and
distribution of the winter stonefly genus Capnia (Plecoptera: Capniidae) in North America. Great Basin
Received 12 April 2002
DISTRIBUTION OF NEUROTRICHUS GIBBSII IN CALIFORNIA
WITH A RANGE EXTENSION IN THE SIERRA NEVADA
Leslie N. Carraway1, B.J. Verts1, and J.W. Goertz2
Key words: shrew-mole, Neurotrichus gibbsii, distribution, range extension, California.
Published distribution maps for the shrewmole (Neurotrichus gibbsii) in California depict
widely variable occupied ranges (Figs. 1A–C).
In our review of published information on the
species (Carraway and Verts 1991), we chose
to use a modification of Hall’s (1981:67, Map
39) distribution map because it was based on
published locality records. It showed no records
for the Sierra Nevada. Regrettably, we overlooked the statement by Jameson and Peeters
(1988:107) that the species “is found in the
northern Sierra Nevada south at least to Plumas
County.”
On 15 June 1999, one of us (JWG) found a
dead specimen about 6 miles S, 4 miles E of
Nevada City, Nevada County, California. The
specimen, preserved in alcohol, is deposited
in the Museum of Vertebrate Zoology, University of California, Berkeley (MVZ 196311).
The locality extends the range approximately
203 km (127 miles) SSE from previous published records at Carberry Ranch (about 9
miles E of the town of Montgomery Creek),
Shasta County; about 189 km (118 miles) ESE
of South Yolla Bolly Mountain, Tehama County;
and about 155 km (97 miles) E of Mill Creek,
Lake County (Williams 1975, Hall 1981). The
locality is approximately 114 km (71 miles)
south of the northern border of Plumas County
in the Sierra Nevada.
The extensive distances between published
locality records and the site of the new Nevada
County record, combined with the widely
variable ranges depicted in published maps
(Figs. 1A–C), caused us to produce a specimen-based distribution map for the species in
California (Fig. 1D). We plotted localities (n =
185) for specimens (n = 466) obtained from
curators of 15 museum collections (Appendix).
Three additional localities from which 4 specimens were collected were not plotted because
of inadequate or contradictory information.
Because all unplotted localities occurred in
counties (Humboldt and Marin) containing
other collection sites, we do not believe their
deletion was significant. Symbols for 39 localities were obscured by those nearby (Fig. 1D).
The distribution of N. gibbsii in California
is considerably more widespread than depicted
in previous publications (Fig. 1). Although our
specimen remains the southernmost record for
the species in the Sierra Nevada, other localities in Plumas and Nevada counties confirm
the occurrence of the species in that mountain
range (Fig. 1D). We postulate that specimens
from these localities and others outside previously depicted ranges are not the result of recent
invasion of new territory by the species, but
represent collections in previously unsampled
areas or are samples obtained from disjunct
and scattered residual populations.
ACKNOWLEDGMENTS
We thank curators of mammal collections at
the National Museum of Natural History
(USNM); American Museum of Natural History (AMNH); Museum of Vertebrate Zoology,
University of California, Berkeley (MVZ); Field
Museum (FMNH); Museum of Wildlife and
Fisheries Biology, University of California, Davis
(MWFB); Natural History Museum of Los
Angeles County (LACM); Carnegie Museum
of Natural History (CM); Occidental College,
Moore Laboratory of Zoology (MLZ); Santa
Barbara Museum of Natural History (SBMNH);
San Diego Natural History Museum (SDNHM);
Humboldt State University (HSU); University
1Department of Fisheries and Wildlife, Nash 104, Oregon State University, Corvallis, OR 97331-3803.
21402 Caddo St., Ruston, LA 71270-5220.
487
488
[Volume 62
Fig. 1. Distribution of the shrew-mole, Neurotrichus gibbsii, in California as indicated by A, Ingles (1965); B, Hall
(1981); and C, Jameson and Peeters (1988); D, 185 localities at which 466 museum specimens of shrew-moles were collected (see Appendix). Three localities were not mapped; symbols for 39 localities are obscured by those nearby. Open
symbol indicates new southernmost record in the Sierra Nevada (MVZ 196311).
2002]
NOTES
of California, Los Angeles, Dickey Collection
(UCLA); California Academy of Sciences (CAS);
University of Kansas, Natural History Museum
(KU); and California State University, Stanislaus (CSCS) for providing locality data for
specimens in their care. We also thank the following curators of collections for responding
to our query: California State University, Long
Beach (CSULB); California State University,
Chico (CSUC); California State University,
Northridge (CSUN); and California State Polytechnic University, Pomona (CSPUP). J.L. Patton commented on an earlier draft of the manuscript.
LITERATURE CITED
CARRAWAY, L.N., AND B.J. VERTS. 1991. Neurotrichus gibbsii. Mammalian Species 387:1–7.
HALL, E.R. 1981. The mammals of North America. 2nd
edition. John Wiley & Sons, New York 1:1–600 + 90.
INGLES, L.G. 1965. Mammals of the Pacific states: California, Oregon, and Washington. Stanford University
Press, Stanford, CA.
JAMESON, E.W., JR., AND H.J. PEETERS. 1988. California
mammals. University of California Press, Berkeley.
WILLIAMS, D.F. 1975. Distribution records for shrew-moles,
Neurotrichus gibbsii. Murrelet 56:2–3.
Accepted 16 August 2001
APPENDIX. Specimen localities for Neurothrichus
gibbsii in California. Localities followed by an
asterisk (*) were not plotted on Figure 1D.
U.S.A.—California, Butte Co.: 1 mi W Forbestown (1
MVZ). Del Norte Co.: Crescent City (2 USNM); Requa (1
SBMNH, 1 CAS); Smith River, 0.25 mi W Douglas Park
(1 MVZ). Glenn Co.: Brittan Ranch, Mendocino National
Forest, 39°23′45.9″N, 122°39′29.2″W (1 CAS); Masterson
Camp (1 MVZ); Telephone Campground, 1.5 mi NNW
Blackbutte (1 MVZ). Humboldt Co.: 7.2 km N, 1.6 km E
Salyer (1 MVZ); Ammon Creek, 6.9 km S Salyer (2 HSU);
Arboritos, 15.3 km S, 1.9 km W Salyer (1 MVZ); Arcata (1
MVZ, 5 HSU); 1 mi E Arcata (1 LACM); Big Lagoon (3
MVZ); Big Rock, 5.5 km S, 3.9 km W Salyer (1 MVZ); 2
mi N, 4 mi E Bridgeville (1 HSU); Brush Mountain, 3 mi
W Salyer (1 MWFB); Brush Mountain, 0.5 km S, 7.1 km
W Salyer (1 MVZ); Carlotta (2 MVZ, 1 SDNHM, 1 HSU,
12 UCLA, 1 KU, 3 CAS); 1 mi E Carlotta (4 UCLA);
Cedar Creek, 1.6 km N, 8 km E Willow Creek (2 MVZ);
Clam Beach (3 HSU); Clam Beach County Park (1 HSU);
Cuddeback (9 MVZ); Despenadero, 1.6 km S, 6.4 km W
Salyer (1 MVZ); Eureka (6 FMNH, 1 MVZ, 1 HSU); 4 mi
N Eureka (1 HSU); 4 mi E Fortuna, T3N, R1W, sec. 34
[sic]* (1 HSU); Halcon, 5.6 km S, 2.9 km E Salyer (1
MVZ); Hatchery Inlet Creek (1 HSU); Hoopa Valley (1
USNM); Humboldt State University (2 HSU); 1 mi NE
489
Humboldt State University (1 HSU); Kneeland (1 HSU);
Maple Creek, 1 mi N jct Mad River (2 MVZ); Orick (3
SDNHM); 3 mi N Orick (1 USNM); 6 mi S, 6 mi E Orick
(1 LACM); Six Rivers National Forest* (1 MWFB);
Trinidad (1 USNM, 4 MVZ, 2 SDNHM, 1 HSU, 2 UCLA,
2 CAS); 3 mi N Trinidad (2 MWFB); 4 mi N Trinidad (3
MWFB); 1.5 mi N, 2 mi W Willow Creek (1 HSU); 3 mi N
Willow Creek (5 MVZ, 1 HSU); 4.5 mi N, 1.5 mi E Willow
Creek (3 MVZ); 1 mi N, 14 mi W Willow Creek (1 HSU);
2 mi S, 12 mi E Willow Creek, T6N, R3E, sec. 8 and 17
[sic]* (2 HSU). Lake Co.: Mill Creek, 4000 ft (1 CSCS).
Marin Co.: Audubon Canyon Ranch (1 CAS); Abalone
Point, 3 mi NW Bolinas (1 MVZ); 3 mi N Bolinas (1 MVZ);
1 mi W Bolinas (1 MVZ); Bolinas Bay (1 MWFB); W end
Elk Valley [37.84187N, 122.550963W] (32 MVZ); Fort
Baker (2 CAS); Camp Taylor, 2 mi WNW Lagunitas (1
MVZ); Fairfax (1 CAS); Canyon near Fairfax (1 MVZ); W
Portal of Tunnel, Fort Baker (1 MVZ); West Portal area,
Fort Barry Military Reservation (17 MVZ); Fort Cronkhite
(1 MVZ); Inverness (1 USNM, 1 MVZ, 1 CAS); 1 mi S
Inverness (1 MVZ); 1 mi SE Inverness (2 MVZ); 2 mi
WNW Inverness (19 MVZ); 3 mi W Inverness (5 MVZ); 4
mi NW Inverness (2 SDNHM); 5 mi W Inverness (1
MVZ); Lagunitas (2 MVZ); Lagunitas Canyon (1 MVZ);
Marin Headlands, Golden Gate National Recreational
Area (1 CAS); Mill Valley (7 CAS); Muir Beach (3 CAS);
Muir Woods (1 MVZ); Murphy’s Ranch, 3.5 mi W Point
Reyes Hill (1 MVZ); Nicasio (2 USNM, 1 AMNH, 5 CAS);
3 mi W Novato (1 CAS); 5 mi NNE Point Reyes Lighthouse (8 MVZ); Point Reyes National Seashore (1 CAS);
Point Reyes, Umentore [= Limentour] Beach (1
SBMNH); 6 mi SSE Tomales Bluff (2 CAS); San
Geronoms [= San Geronimo] (1 FMNH, 1 MVZ); San
Rafael, 82 Southern Heights Blvd (1 MVZ); Sausalito (1
CAS); 1 mi N Stinson Beach (1 CAS); Mount Tamalpais (1
CAS). Mendocino Co.: 0.5 mi E Albion (4 AMNH); 0.5 mi
NE Albion (1 CSCS); Point Arena (4 CM, 4 MVZ); 6 mi
SE Point Arena (1 CM); 8 mi SE Boonville (1 HSU); Brandon Gulch, Jackson State Park (1 MVZ); 5 mi N, 3.5 mi W
Branscomb (1 HSU); 6 mi N, 0.5 mi W Branscomb (1
HSU); 7 mi N, 1.5 mi E Branscomb (1 HSU); 8 mi N, 3 mi
W Branscomb (1 HSU); Gualala (1 CM, 1 MVZ); Gualala
River, near Gualala (1 CM); 2 mi N. Gualala (1 MVZ); Laytonville (1 MVZ); Mendocino City (1 MVZ); 4.3 mi E
Navarro (2 MVZ); Russian Gulch (1 MVZ); Russian Gulch
State Park (16 MVZ); Van Damme Beach State Park (1
CAS); 5 mi NW Yorkville (1 MVZ, 6 CAS). Napa Co.:
Angwin [T8N, R5W] (1 MVZ); Bothe State Park, Napa
Valley (1 MVZ); Napa Valley, 1 mi W Napa, Portrich Rd (1
MVZ). Nevada Co.: 8 mi NE Nevada City (1 MWFB); 6
miles S, 4 miles E, Nevada City (1 MVZ). Plumas Co.:
Buck Ranch, 39°55′N, 121°20′W (1 UCLA); Butt Creek, 8
mi S Chester (1 MVZ); Quincy (3 MWFB); 2 mi E Quincy
(2 MWFB, 1 MVZ); Rich Gulch, 8 mi N, 11 mi W Quincy
(5 MVZ); Sierra Valley (1 CAS). San Benito Co.: Fremont
Peak (1 USNM); Gabilan Mountains, 3.5 mi W San Juan
Bautista (1 MVZ). San Francisco Co.: San Francisco, near
SW corner Olympic Club, Lakeside (1 MVZ); San Francisco, S end Lake Merced (3 CAS). San Mateo Co.:
Colma (1 CAS); 0.9 mi NE Colma (1 CAS); Crystal
Springs (1 CAS); Palo Alto (1 FMNH); Portola (1 UCLA);
2 mi WNW Portola (3 MVZ); Portola Valley (2 CAS); Tunitas (8 SDNHM); Woodside (1 CAS); 2 mi N Woodside
[Filoli] (1 MVZ). Santa Cruz Co.: 5 mi S Aptos (10
USNM); 2 mi N Ben Lomond (1 CAS); Bear Creek, 2 mi
N Boulder Creek [T9S, R2W] (2 MVZ); Big Basin, E Waddell Creek (1 MVZ); Greenly Farm, 4 mi from Boulder
490
Creek (1 CAS); Santa Cruz (5 USNM, 2 MVZ, 1
SBMNH); 7 mi N Santa Cruz (1 CAS); Santa Cruz Mountains between Santa Cruz and Los Gatos (1 CAS); 7 mi W
Stanford University (2 CAS); Valencia Park Ranch [T11 S,
R1E] (1 MVZ); Waddell Creek, 1 mi N, 3 mi E Ano Neuvo
Point (1 MVZ); Waddell Creek, 0.5 mi N, 4 mi E Ano
Neuvo Point (1 MVZ); Waddell Creek [T9S, R4W] (1
MVZ). Shasta Co.: Carberry Ranch, between Mt. Shasta
and Mt. Lassen [about 9 miles E town of Montgomery
Creek] (3 USNM); 12 mi S McCloud River (1 MWFB);
Tower House (2 MVZ). Siskiyou Co.: Beswick (1 USNM);
Clear Creek, 3 mi W Klamath River (1 MVZ); Davis
Creek, 2 km N, 0.5 km E Somes Bar (1 MVZ); 8 mi SE
Somes Bar (1 MVZ); Mt. Shasta, Upper Mud Creek (3
USNM); Mt. Shasta, Wagon Camp (1 USNM); Mt. Shasta
City (2 UCLA); 0.5 mi SW Mt. Shasta City (1 MVZ);
Salmon Mountains, near Etna Mills (1 USNM); Ukonom
Station, 1 km N, 1.6 km E Somes Bar (2 MVZ). Sonoma
Co.: 7 mi W Cazadero (1 MVZ); Duncan’s Mills (1 MVZ);
Freestone (1 MVZ); Gualala, S side Gualala River (8
MVZ); 1 mi W Guerneville (5 MVZ); Salmon Creek, 2 mi
[Volume 62
N Bogeda Bay (3 LACM); Salmon Creek, 2 mi NW
Bogeda Bay (6 LACM); Salmon Creek, 3 mi N Bogeda
Bay (1 LACM); Sonoma (1 CAS). Tehama Co.: 2 mi W
Black Butte (2 MVZ); Lyman, 4 mi NW Lyonsville (1
MVZ); 2 mi E Mineral (1 MVZ); 2 mi S South Yolla Bolly
Mountain (1 MVZ). Trinity Co.: Beartooth Mountain, 4.8
km N, 10 km E Burnt Ranch (6 MVZ); 1.6 km N, 6.9 km E
Burnt Ranch (1 HSU); Cedar Basin, 1 mi SW North Yolla
Bolly Mountain (1 AMNH); Dyer Creek, 0.8 km S, 9 km E
Salyer (1 MVZ); Eastman Gulch [40°45′22″N, 122°46′35″W]
(1 SDNHM); Hennessy Peak, 3.2 km W Burnt Ranch (3
MVZ); Dulce, 3.5 km S, 5.5 km E Big Bar (1 MVZ); 4.5
km W, 7.2 km N Big Bar (1 HSU); Hennessy Ridge, 1.5
km E Salyer (1 MVZ); 4.8 km E Salyer (2 HSU); 1.5 mi N
Mad River Bridge, South Fork Mountain (3 MVZ); 3 mi N
Mad River Bridge, South Fork Mountain (1 MVZ);
Nogaar’s Ranch, South Fork Trinity River [40.415129N,
123.408238W] (1 MVZ); South Fork, 6 km S, 1.6 km E
Salyer (1 MVZ); Swede Creek, 7.4 km N, 8 km W Big Bar
(4 MVZ). Yolo Co.: Copay Valley (1 MWFB).
HELMINTHS OF THE PLAINS SPADEFOOT, SPEA BOMBIFRONS,
THE WESTERN SPADEFOOT, SPEA HAMMONDII,
AND THE GREAT BASIN SPADEFOOT, SPEA INTERMONTANA
(PELOBATIDAE)
Stephen R. Goldberg1 and Charles R. Bursey2
Key words: Spea bombifrons, Spea hammondii, Spea intermontana, helminths, Trematoda, Cestoda, Nematoda.
The plains spadefoot, Spea bombifrons (Cope,
1863), occurs from southern Alberta, Saskatchewan, and Manitoba to eastern Arizona and
northeastern Texas south to Chihuahua, Mexico;
the western spadefoot, Spea hammondii (Baird,
1859), occurs from the Great Valley of California and Coast Ranges south of San Francisco
Bay, California, into northwestern Baja California, mainly below 910 m; the Great Basin
spadefoot, Spea intermontana (Cope, 1883),
ranges from southern British Columbia through
the Great Basin to northwestern Arizona (Stebbins 1985). The 3 species are allopatric throughout their ranges. Taxonomy is according to
Crother (2000): Spea = Scaphiopus in part.
There are 2 reports of helminths (Rodgers
1941, Brooks 1976) for S. bombifrons; but, to
our knowledge, there are no reports of helminths for S. hammondii or S. intermontana,
although the biology of S. intermontana has
been summarized (Hall 1998). The purpose of
this paper is to add to the helminth list of S.
bombifrons and to provide the initial account
of helminths for S. hammondii and S. intermontana.
Thirty-five adult specimens (19 female, 16
male) of Spea bombifrons collected 1953–
1962, 31 adult specimens (9 female, 22 male)
of S. hammondii collected 1938–1975, and 34
adult specimens (11 female, 23 male) of S.
intermontana collected 1937–1964 were borrowed from museum collections. All S. bombifrons were from Arizona (snout-vent length,
SVL = 46 mm ± 3 s, range = 38–53 mm). All
S. hammondii were from California (SVL = 46
mm ± 7 s, range = 31–58 mm). Twelve S.
intermontana were from Arizona (SVL = 57
mm ± 2 s, range = 54–61 mm), 8 from Nevada
(SVL = 50 mm ± 3 s, range = 47–55 mm), and
14 from Utah (SVL = 57 mm ± 6 s, range =
44–67 mm). Museum accession numbers and
counties of collection are given in the Appendix. For each toad the body cavity was opened
and the lungs, esophagus, stomach, small intestine, large intestine, bladder, and body cavity
were searched for helminths. Each nematode
was placed in a drop of glycerol on a glass
slide, allowed to clear, and then identified.
Cestodes and trematodes were stained in hematoxylin, dehydrated in a graded series of ethanol, cleared in xylene, and mounted in balsam
for identification. Representative samples were
deposited in the United States National Parasite
Collection, Beltsville, Maryland (Appendix).
Five (14%) of 35 Spea bombifrons were found
to harbor helminths: 1 male with immature individuals of the trematode Polystoma nearcticum (Paul, 1935); 1 male with gravid individuals of the nematode Aplectana incerta Caballero, 1949; 1 female and 1 male with gravid
individuals of the nematode Aplectana itzocanensis Bravo Hollis, 1943; and 1 female with
1 larva of Physaloptera sp. (Nematoda). Infection rates are too low for comparative (female,
male) statistical analyses; helminth numbers,
site of infection, prevalence (percentage of
infected toads), mean intensity (mean number
of helminths per infected toad) ± 1 s, and
range (low to high number of helminths per
infected toads) are presented in Table 1. Spea
bombifrons represents a new host record for
Polystoma nearcticum, Aplectana incerta, and
A. itzocanensis.
1Department of Biology, Whittier College, Whittier, CA 90608.
2Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, PA 16146.
491
492
[Volume 62
TABLE 1. Number of helminths, prevalence, mean intensity ± 1 s, range, and infection site for 35 Spea bombifrons
from Arizona.
Helminth
Site of infection
TREMATODA
Polystoma nearcticum
lung
NEMATODA
Aplectana incerta
Large intestine
Aplectana itzocanensis
Large intestine
Physaloptera sp.
Stomach
Number
helminths
Prevalence
(%)
Mean
intensity
Range
3
3
3.0
0
29
6
14.5 ± 19.0
1–28
1
3
1.0
0
1
3
1.0
0
Five of 31 S. hammondii were found to harbor gravid females of Aplectana incerta (large
intestine of 3 male and 2 female toads, prevalence = 16%, mean intensity = 3.6 ± 1.7 s,
range 1–5). Spea hammondii is a new host
record for Aplectana incerta.
Sixteen (47%) of 34 S. intermontana were
found to harbor helminths: 3 females and 11
males with gravid individuals of the trematode
Polystoma nearcticum; 2 males with gravid individuals of the cestode Distoichometra bufonis Dickey, 1921; 6 males with gravid individuals of Aplectana incerta; 1 male with 5 larvae
of an unidentified species of an acuariid nematode; and 1 male with 2 larvae of Physaloptera
sp. There was no statistical difference for infection by Polystoma nearcticum in female and
male toads (χ2 = 0.57, 1 df, P > 0.05); infection rates by the other species of helminths are
too low for comparative statistical analyses. We
report the presence of Aplectana incerta only
in male hosts because only male toads comprised the Nevada subsample. Helminth numbers, site of infection, prevalence, mean ± 1 s,
and range by location are presented in Table 2.
Spea intermontana represents a new host record
for Polystoma nearcticum, Distoichometra bufonis, and Aplectana incerta.
The monogenean Polystoma nearcticum was
originally described from Hyla versicolor collected in New England and H. cinerea from
Florida (Paul 1935). Examination of the life
cycle has revealed the presence of a rapidly
maturing brachial form (22 days) and a more
slowly maturing (3 years) bladder form (Paul
1938). Only brachial forms were found in S.
bombifrons; both brachial and bladder forms
were found in S. intermontana. It should be
noted that Rodgers (1941) described the mono-
genean Neodiplorchis scaphiopodis (= Diplorchis scaphiopodis) from S. bombifrons collected in Oklahoma and that Brooks (1976)
reported it from the same host collected in
Nebraska. In the original descriptions, immature forms of these 2 species were reported to
have different numbers of hooks between the
anterior suckers of the opisthohaptor: N. scaphiopodis with 6, P. nearcticum with 8. Because
immature trematodes collected from S. bombifrons in this study possessed 8 hooks between
the anterior suckers, we have assigned them
to P. nearcticum.
Distoichometra bufonis was originally described from Bufo terrestris collected in Georgia (Dickey 1921). Distoichometra kozloffi was
placed in synonymy with D. bufonis by Jones
(1987); thus, only a single species of Distoichometra is recognized for North America.
This species is a common cestode of North
American anurans and, in addition to B. terrestris, has been reported from Bufo americanus,
B. boreas, B. cognatus, B. debilis, B. microscaphus, B. punctatus, B. retiformis, B. woodhousii, Pseudacris regilla, Pternohyla fodiens,
Scaphiopus couchii, S. holbrookii, and Spea multiplicata (Brandt 1936, Odlaug 1954, Douglas
1958, Koller and Gaudin 1977, Goldberg and
Bursey 1991a, 1991b, Goldberg et al. 1996a,
1996b, 1999). Although D. bufonis was found
only in Nevada in this study, it has been reported from Arizona and Utah (Parry and
Grundmann 1965, Goldberg and Bursey 1991a).
Aplectana incerta was originally described
from Bufo marinus collected in Mexico (Caballero 1949) and has been reported from toads
of Arizona and New Mexico, namely, Bufo debilis, B. microscaphus, B. retiformis, B. woodhousii, Gastrophryne olivacea, Scaphiopus
Helminth
Site of infection
0
5.0
7
5
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
0
8
1
1.0
1–3
2.0 ± 1.4
14
2–44
—
—
—
—
90
50
22.5 ± 23.7
4
—
—
—
—
0
4.0
13
4
—
—
—
—
2–18
10.5 ± 7.3
29
33
3.3 ± 2.1
1–5
13
75
2.2 ± 1.5
1–5
42
(n = 14)
(n = 8)
(n = 12)
13
Trematoda
Polystoma nearcticum
Lung and bladder
Cestoda
Distoichometra bufonis
Small intestine
Nematoda
Aplectana incerta
Small and large intestines
Physaloptera sp. (larvae)
Stomach
Acuariidea gen. sp. (larvae)
In cysts on stomach wall
Nevada
____________________________________
Number Prevalence Mean
helminths
(%)
intensity Range
Utah
____________________________________
helminths
(%)
intensity Range
NOTES
Arizona
____________________________________
helminths
(%)
intensity Range
TABLE 2. Number of helminths, prevalence, mean intensity ± 1 s, range, and infection site for 34 Spea intermontana from Arizona, Nevada and Utah.
2002]
493
couchii, and Spea multiplicata (Goldberg et al.
1998). This study extends the range of A.
incerta, previously considered a middle-American species (see Goldberg et al. 1998), into
the Great Basin of western North America as
well as California.
Aplectana itzocanenesis was originally described from Spea multiplicata (= Scaphiopus
multiplicatus) from Puebla, Mexico, by Bravo
Hollis (1943) and was also found in S. multiplicata from New Mexico by Goldberg et al.
(1995). It has been reported from the following toads of Arizona and New Mexico: Bufo
alvarius, B. cognatus, B. debilis, B. microscaphus, B. punctatus, B. retiformis, B. woodhousii,
Gastrophryne olivacea, Scaphiopus couchii,
and Spea multiplicata (Goldberg et al. 1998).
Additional examinations of Great Basin anurans will be required to determine if, like A.
incerta, the range of A. itzocanensis extends
north of Arizona and New Mexico.
Third stage larvae of Physaloptera sp. (but
not adults) are known from a variety of amphibians and reptiles (see Goldberg et al. 1993).
Members of the Physalopteridae require insect
intermediate hosts (Anderson 2000). They enter
amphibians or reptiles in insect prey, no further
development occurs, and they subsequently
pass from the body with feces. Species of the
Acuariidae are bird parasites requiring arthropod intermediate hosts (Anderson 2000). They
also enter with prey and then migrate to tissue
where they are found in cysts; no further
development occurs.
Four species of Spea occur in North America: S. bombifrons, S. hammondii, S. intermontana, and S. multiplicata (Crother 2000). Prior
to this study, Rodgers (1941) and Brooks (1976)
reported the trematode Neodiplorchis scaphiopodis (= Diplorchis scaphiopodis) from S. bombifrons collected in Oklahoma and Nebraska,
respectively; Lamothe-Argumedo (1973)
found N. scaphiopodis in S. multiplicata from
Mexico; and Goldberg et al. (1995) reported
Distoichometra bufonis, Aplectana incerta, and
A. itzocanensis from S. multiplicata collected
in New Mexico. Although these helminths are
generalist parasites (occurring in 2 or more host
species) of anurans, they have been encountered infrequently in species of Spea. However, these helminths are now known to occur
in at least 2 species of Spea: Neodiplorchis
scaphiopodis in S. bombifrons and S. multiplicata; Polystoma nearcticum in S. bombifrons
494
and S. intermontana; Distoichometra bufonis
in S. intermontana and S. multiplicata;
Aplectana incerta in S. bombifrons, S. hammondii, S. intermontana, and S. multiplicata;
Aplectana itzocanensis in S. bombifrons and S.
multiplicata. Further examination of species of
Spea as well as Scaphiopus will be required
before the number of shared helminth species
within the genus and the family Pelobatidae
can be known.
Spea bombifrons were loaned to us by
Charles H. Lowe (University of Arizona).
David B. Wake (Museum of Vertebrate Zoology, University of California, Berkeley), Bradford D. Hollingsworth (San Diego Natural
History Museum), and David A. Kizirian (Natural History Museum of Los Angeles County)
loaned the S. hammondii. Michael E. Douglas
(formerly of Arizona State University), Jack W.
Sites, Jr. (Brigham Young University), David
A. Kizirian, and Charles H. Lowe allowed us
to examine S. intermontana.
LITERATURE CITED
ANDERSON, R.C. 2000. Nematode parasites of vertebrates:
their development and transmission. 2nd edition.
CABI Publishing, Wallingford, Oxon, U.K. 650 pp.
BRANDT, B.B. 1936. Parasites of certain North Carolina
Salientia. Ecological Monographs 6:491–532.
BRAVO HOLLIS, M. 1943. Dos neuvos nemátodos parásitos
de anuros del sur de Puebla. Anales del Instituto de
Biología, Universidad Nacional Autónoma de México 14:69–78.
BROOKS, D.R. 1976. Parasites of amphibians of the Great
Plains. Part 2. Platyhelminths of amphibians in
Nebraska. Bulletin of the University of Nebraska
State Museum 10:65–92.
CABALLERO, C.E. 1949. Estudios helmintológicos de la
región oncocercosa de México y de la República de
Guatemala. Nematoda. 5a parte. Anales del Instituto
de Biología, Universidad Nacional Autónoma de
México 20:279–292.
CROTHER, B.I. 2000. Scientific and standard English names
of amphibians and reptiles of North America north
of Mexico, with comments regarding confidence in
our understanding. Society for the Study of Amphibians and Reptiles, Herpetological Circular 29. 82 pp.
DICKEY, L.B. 1921. A new amphibian cestode. Journal of
Parasitology 7:129–137.
DOUGLAS, L.T. 1958. The taxonomy of nematotaeniid cestodes. Journal of Parasitology 44:261–273.
GOLDBERG, S.R., AND C.R. BURSEY. 1991a. Helminths of
three toads, Bufo alvarius, Bufo cognatus (Bufonidae), and Scaphiopus couchii (Pelobatidae), from
southern Arizona. Journal of the Helminthological
Society of Washington 58:142–146.
______. 1991b. Helminths of the red-spotted toad, Bufo
punctatus (Anura: Bufonidae), from southern Arizona.
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GOLDBERG, S.R., C.R. BURSEY, AND H. CHEAM. 1998. Nematodes of the great plains narrow-mouthed toad, Gastrophryne olivacea (Microhylidae), from southern
Arizona. Journal of the Helminthological Society of
GOLDBERG, S.R., C.R. BURSEY, AND G. GALINDO. 1999.
Helminths of the lowland burrowing treefrog, Ptyernohyla fodiens (Hylidae), from southern Arizona.
Great Basin Naturalist 59:195–197.
GOLDBERG, S.R., C.R. BURSEY, K.B. MALMOS, B.K. SULLIVAN, AND H. CHEAM. 1996a. Helminths of the southwestern toad, Bufo microscaphus, Woodhouse’s toad,
Bufo woodhousii (Bufonidae), and their hybrids from
central Arizona. Great Basin Naturalist 56:369–374.
GOLDBERG, S.R., C.R. BURSEY, B.K. SULLIVAN, AND Q.A.
TRUONG. 1996b. Helminths of the Sonoran green
toad, Bufo retiformis (Bufonidae), from southern Arizona. Journal of the Helminthological Society of
GOLDBERG, S.R., C.R. BURSEY, AND I. RAMOS. 1995. The
component parasite community of three sympatric
toad species, Bufo cognatus, Bufo debilis (Bufonidae), and Spea multiplicata (Pelobatidae) from New
Mexico. Journal of the Helminthological Society of
GOLDBERG, S.R., C.R. BURSEY, AND R. TAWIL. 1993. Gastrointestinal helminths of the western brush lizard, Urosaurus graciosus graciosus (Phrynosomatidae). Bulletin of the Southern California Academy of Sciences
92:43–51.
HALL, J.A. 1998. Scaphiopus intermontanus Cope. Great
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and Reptiles 650.1–650.17.
JONES, M.K. 1987. A taxonomic revision of the Nematotaeniidae Lühe, 1910 (Cestoda: Cyclophyllidea). Systematic Parasitology 10:165–245.
KOLLER, R.L., AND A.J. GAUDIN. 1977. An analysis of
helminth infections in Bufo boreas (Amphibia:
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southern California. Southwestern Naturalist 21:
503–509.
LAMOTHE-ARGUMEDO, R. 1973. Monogéneos de los anfibios
de México V. Descripción de la larva de Neodiplorchis scaphiopi (Rodgers, 1941) Yamaguti, 1963 (Monogénea: Polystomatidae). Anales del Instituto de Biología, Universidad Nacional Autónoma de México
44:9–14.
ODLAUG, T.O. 1954. Parasites of some Ohio Amphibia.
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PARRY, J.E., AND A.W. GRUNDMANN. 1965. Species composition and distribution of the parasites of some common amphibians of Iron and Washington counties,
Utah. Proceedings of the Utah Academy of Arts and
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PAUL, A.A. 1935. Polystoma integerrimum nearcticum n.
subsp. from the urinary bladder, genital ducts, kidneys and gills of Hyla versicolor Le Conte. Journal of
Parasitology 21:442.
______. 1938. Life history studies of North American freshwater polystomes. Journal of Parasitology 24:489–510.
RODGERS, L.O. 1941. Diplorchis scaphiopi, a new polystomatid monogenean fluke from the spadefoot toad.
Journal of Parasitology 27:153–157.
2002]
NOTES
STEBBINS, R.C. 1985. A field guide to western reptiles and
amphibians. Houghton Mifflin Company, Boston,
MA. 336 pp.
Received 3 December 2000
Accepted 16 August 2001
APPENDIX
LOCALITIES AND MUSEUM NUMBERS
FOR SPECIMENS EXAMINED
Spea bombifrons: University of Arizona (UAZ) ARIZONA (Cochise County) 7378, 7379, 7381, 7386, 7388,
7389, 7391, 7392, 7364–7366, 7368, 7395, 7399–7408,
7410–7412, 7414–7418, 7420, 7421, 7423, 7425.
Spea hammondii: Natural History Museum of Los
Angeles County (LACM) CALIFORNIA (Madera County)
147873; Museum of Vertebrate Zoology (MVZ) (Madera
County) 25963, 31874, 54046–54049, 54202, 54203,
55519, 56734, 60284–60291, 60983, 76003; San Diego
495
Natural History Museum (SDSNH) (San Diego County)
55335–55337, 55518–55524.
Spea intermontana: Arizona State University (ASU)
NEVADA (Lincoln County) 21290, 21291, 21293, 21297–
21299, 21301, 21302; Brigham Young University (BYU)
UTAH (Garfield County) 1970, 1973, 1977, 2772, (Carbon
County) 2058, 2800, (Daggett County) 14180, (Tooele
County) 14789, 14791; Natural History Museum of Los
Angeles County (LACM) UTAH (Carbon County) 90959–
90963; University of Arizona (UAZ) ARIZONA (Mohave
County) 14668, 14670, 14672, 14674, 14676, 14679, 14680,
14683–14685, 14688, 14690.
*****
Accession numbers for helminths in the U.S. National
Parasite Collection (USNPC):
Spea bombifrons: Polystoma nearcticum (91240); Aplectana
incerta (91241); Aplectana itzocanensis (91242); Physaloptera sp. (91243). Spea hammondii: Aplectana incerta
(92015). Spea intermontana: Polystoma nearcticum (90903,
90906); Distoichometra bufonis (90904); Aplectana incerta
(90905, 90907); Physaloptera sp. (90902); acuariid larvae
(90908).
EXTIRPATION OF BAILEY’S POCKET MOUSE,
CHAETODIPUS BAILEYI FORNICATUS (HETEROMYIDAE: MAMMALIA),
FROM ISLA MONTSERRAT, BAJA CALIFORNIA SUR, MEXICO
Sergio Ticul Alvarez-Castañeda1 and Patricia Cortés-Calva1
Key words: Rodentia, Chaetodipus baileyi, islands, extinction, Mexico.
Seven taxa of rodents endemic to northwestern Mexico recently have been reported
as extirpated: Peromyscus maniculatus cineritius, P. pembertoni, Dipodomys gravipes, Oryzomys couesi peninsularis, Neotoma anthonyi,
N. bunkeri, and N. martinensis (Lawlor 1983,
Mellink 1992, Smith et al. 1993, Alvarez-Castañeda 1994, Alvarez-Castañeda and CortésCalva 1996). Extirpation of these rodents may
be a consequence of human activity and the
introduction of nonnative species, primarily
cats (Felis silvestris) that prey on the endemic
rodents and Mus musculus and Rattus rattus
that may compete with native species for resources (Alvarez-Castañeda 1997). We report
the possible extirpation of an endemic rodent,
Bailey’s pocket mouse (Chaetodipus baileyi
fornicatus), from Isla Montserrat.
Isla Montserrat, located 13 km east of the
Baja California peninsula, has an area of 19.4
km2 (Nieto-Garibay 1999). The island has many
small mountains and canyons; the soil is poor
and in some areas very stony. Dominant plants
on Isla Montserrat include golondrina (Euphorbia magdalenae), pitaya agría (Stenocereus gummosus), matacora ( Jatropha cuneata), cholla
(Opuntia cholla), dipúa o medesá (Cercidium
microphyllum), and palo fierro (Olneya tesota;
León De La Luz and Pérez Navarro 1997).
Chaetodipus baileyi has 7 subspecies, 2 of
which inhabit islands in the Sea of Cortez: C.
b. fornicatus on Isla Montserrat and C. b. insularis on Isla Tiburon. Three other subspecies
occur on the Baja peninsula and 2 in the state
of Sonora (Patton and Alvarez-Castañeda 1999).
The Bailey’s pocket mouse of Montserrat island
(C. b. fornicatus) was described by Burt (1932)
on the basis of 12 specimens. Ecological data
were not provided in the original description.
This taxon is considered rare by Mexico (NOM059-Ecol 1994) but is not listed by CITES
(1989).
The last 2 specimens of C. b. fornicatus from
Montserrat island (10, collected 21 May 1975)
are housed in the mammal collection of the
Instituto de Biología of the Universidad Nacional Autónoma de México. Subsequent surveys
of the islands did not yield additional specimens, but specimens of Burt’s deer mouse,
Peromyscus caniceps, were collected on 20 May
1975 (31), 10 August 1986 (2), 16 January 1987
(5), 11 May 1987 (3), 24 October 1995 (3), 18
May 1997 (3), and 27 April 1998 (1).
Four surveys of rodents in various areas of
Montserrat island were conducted over a period
of 5 years by the Centro de Investigaciones
Biológicas del Noroeste S. C. and the Universidad Autónoma of México. The surveys
attempted to avoid seasonal and annual variations in capture success by trapping in different years and months: August 1993 (600 trapnights on the southwestern part of the island);
November 1995 (950 trap-nights, western; 700
trap-nights, northwestern); May 1997 (479 trapnights, northeastern; 300 trap-nights, eastern);
and April 1998 (120 trap-nights, northern end
of the island), for a total of 3149 trap-nights.
Sherman and Museum Special traps were used
in all surveys. Transects of 40 traps, with 10-m
spacing between traps, were set in a variety of
habitats, avoiding cliffs and favoring scrub areas
likely preferred by the mice.
The proportion of P. caniceps collected in
every survey was about 0.2% (Alvarez-Castañeda et al. 1998), but no C. baileyi fornicatus
were trapped. No Mus or Rattus were collected.
1Centro de Investigaciones Biológicas del Noroeste, Mar Bermejo, 195 Playa Palo Santa Rita, La Paz, Baja California Sur, 23090 México.
496
2002]
NOTES
497
cial support to STA-C was provided by the
Consejo Nacional de Ciencia y Tecnología
(CONACyT J28319N).
LITERATURE CITED
–––
Fig. 1. Location of Montserrat island, off the eastern
coast of Baja California peninsula. Heavy black borders
around the island represent cliffs.
Feces of feral cats were found on the island in
all years, but these did not contain Chaetodipus hairs or bones. Cat feces were observed in
all parts of the island visited during a botanical
survey of the island in 1998 (Leon De la Luz
personal communication). It is not known when
cats were introduced to the island. Isla Montserrat currently has a fishermen’s camp, but
human activity otherwise is very low.
It is now more than 25 years since the last
C. b. fornicatus were trapped, and there is no
subsequent evidence of this rodent occurring
on the island. The most plausible explanation
for the apparent extirpation of Chaetodipus
baileyi fornicatus is predation by domestic cats.
We thank the Mexican Navy for transport
to Montserrat island; D. Hafner, M. Hafner, B.
Riddle, and Cheryl Patten for reviewing the
manuscript draft; and F.A. Cervantes for use of
the Colección Nacional de Mamíferos. Finan-
ALVAREZ-CASTAÑEDA, S.T. 1994. Current status of the rice
rat Oryzomys couesi peninsularis. Southwestern Naturalist 39:99–100.
______. 1997. Diversidad y conservación de mamíferos terrestres en el estado de Baja California Sur, México.
Ph.D. thesis, Universidad Nacional Autónoma de
México. 221 pp.
ALVAREZ-CASTAÑEDA, S.T., AND P. CORTÉS -CALVA. 1996.
Anthropogenic extinction of the endemic deer mouse,
Peromyscus maniculatus cineritius, on San Roque
island, Baja California Sur, Mexico. Southwestern
ALVAREZ-CASTAÑEDA, S.T., P. CORTÉS-CALVA, AND C. GÓMEZMACHORRO. 1998. Peromyscus caniceps. Mammalian
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LEÓN DE LA LUZ, J.L., AND J.J. PÉREZ NAVARRO. 1997.
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islands, Baja California, México. Baja California
Botanical Symposium, San Diego Natural History
Museum.
MELLINK, B.E. 1992. Status de los Heterómidos y Cricétidos endémicos del Estado de Baja California. Contribuciones Académica, Serie Ecológica, Centro Investigaciones Científicas y Educativas Superiores de
Ensenada, 1–10.
NIETO -GARIBAY, A. 1999. Características generales del
Noroeste de México. Pages 10–25 in S.T. AlvarezCastañeda and J.L. Patton, editors, Mamíferos del
Noroeste de México. Centro de Investigaciones Biológicas del Noroeste, S. C., La Paz, México.
NOM-059-ECOL. 1994. Norma Oficial Mexicana, que
determina las especies y subespecies de flora y fauna
silvestre terrestre y acuática en peligro de extinción,
amenazadas, raras y sujetas a protección especial, y
establece especificaciones para su protección. 16
mayo.
PATTON, J.L., AND S.T. ALVAREZ-CASTAÑEDA. 1999. Family
Heteromyidae. Pages 351–444 in S.T. Alvarez-Castañeda and J.L. Patton, editors, Mamíferos del Noroeste de México. Centro de Investigaciones Biológicas del Noroeste, S. C., La Paz, México.
SMITH, F.A., B.T. BESTELMEYER, J. BIARDI, AND M. STRONG.
1993. Anthropogenic extinction of the endemic woodrat, Neotoma bunkeri Burt. Biodiversity Letters 1:
149–155.
Received 7 August 2000
REMOVAL OF RHODODENDRON MACROPHYLLUM PETALS
BY CAMPONOTUS MODOC
Michael D. Weiser1
Key words: Formicidae, nectivory, pollination, pollinator.
Most research on the relationship between
ants and plants has focused on mutualistic interactions in which plants benefit from the presence of ants (Hölldobler and Wilson 1990).
While there are examples of ants removing
petals as a food source (Cerdá et al. 1992, Hölldobler and Wilson 1990), I know of no report
of ants removing petals specifically to access
floral nectaries. There are few reports in the
literature of ants collecting floral nectar (Tobin
1994), but ants will readily accept floral nectar
when it is accessible (Schubart and Anderson
1978, Guerrant and Fiedler 1981, Haber et al.
1981). Herein I report observations of Camponotus modoc removing petals from Rhododendron macrophyllum flowers, a behavior that
may have impacts on R. macrophyllum pollination biology.
Camponotus modoc (Hymenoptera: Formicidae: Formicinae), a common carpenter ant of
the Pacific Northwest, nests at the base of live
trees, as well as in dead logs in old-growth
forests. In clear-cuts they often nest in and
under stumps. Rhododendron macrophyllum
(Ericaceae) is a common shrub of moist to dry,
coniferous or mixed forests ranging from British
Columbia south to California. In the Western
Cascade Range of Oregon, R. macrophyllum
flowers in May and June. Flowers have large
pink blossoms that are collected in racemes of
20 or more flowers. Individual flowers last a
few days, and then the entire corolla (individual petals are fused to each other) wilts, turns
brown, and falls off the flower. Rhododendron
macrophyllum is bee pollinated (Halverson
1986), but many other insects and hummingbirds visit Rhododendron flowers for nectar
(Pojar 1975).
I made all observations reported here in
H.J. Andrews Experimental Forest, Willamette
National Forest, in the Western Cascade Range
of Oregon. My initial observation, in full sun
at 1400 hours on 2 June 1997, was of R. macrophyllum in flower in a clear-cut experimental
plot at ~650 m. Several C. modoc workers were
clipping petals off apparently normal R. macrophyllum flowers. The workers, typically a single individual to a flower, were cutting around
the base of the petals with their mandibles,
clipping off the entire corolla intact, which then
fell to the ground. The remaining flower had
very little or no petal remaining (i.e., all pinkcolored portions of the flower were removed.)
The ants remained on the flower head after
removing the corolla and appeared to collect
nectar from the base of the flower. They placed
their mouthparts on the base of the flower and
their gasters appeared distended, indicating
that they had taken up liquid from the plant.
None of the flowers I observed being clipped
had any serious damage to their petals (e.g.,
browning or wilting) before cutting by the ant.
I observed this behavior on 6 other R. macrophyllum in the immediate area. Two R. macrophyllum in the area had C. modoc on them,
but I did not observe petal removal. I saw this
behavior over several days at this single site,
but not on R. macrophyllum at other localities
within H.J. Andrews. I also observed at least
10 species of flying insects, including Hymenoptera, Lepidoptera, and Diptera, landing on
intact flowers. The only flying insects observed
landing on flowers lacking petals were asilid
flies that used them as perches.
While speculative, I suggest that C. modoc
workers removed the petals to access the floral
nectar of R. macrophyllum and that removing
1University of Oklahoma, Department of Zoology, 730 Van Vleet Oval, Room 314, Norman, OK 73019-0235. Present address: University of Arizona,
Department of Ecology and Evolutionary Biology, 1041 E. Lowell Street, Tucson, AZ 85721.
498
2002]
NOTES
petals allowed the ants to monopolize the nectar source. It is possible that the petals limited
C. modoc access to the inside of the corolla.
Two smaller species of ants (Tapinoma sessile
and Formica sp.) were observed within intact
flowers. Several small C. modoc were observed
within corollas of unclipped flowers, but many
more were observed on the outer surface of
the flowers, which may indicate that petals
limit C. modoc access to the floral nectaries.
Given that removal of the corolla was almost
complete, pollinators, which use the showy
pink petals as a cue to find floral nectar, would
be less likely to visit flowers lacking these cues.
As only C. modoc and asilid flies were seen
visiting clipped flowers, the nectaries were, in
effect, monopolized by the ants.
To differentiate between the hypotheses
that flower clipping behavior was for access or
monopolization (or both), one could manipulate petals of R. macrophyllum in the presence
of ants and observe their behavior. For example, if a portion of the corolla large enough for
the ants to pass through were removed, then the
access hypothesis predicts that they would not
continue clipping petals. The monopolization
hypothesis predicts complete petal removal.
I observed more than 10 flowers being cut
from a single plant in 2 hours. Estimating 150–
200 flowers on the plant, one can easily extrapolate the potential impact of this behavior on
the pollination biology of R. macrophyllum.
Ants are not effective pollinators (Hölldobler
and Wilson 1990), and removal of petals should
reduce the number of potential pollination
events by reducing blooming time for individual flowers. Petal removal also may affect the
499
energy budget of R. macrophyllum, as non-pollinating nectar-feeders such as hummingbirds
would not access flowers lacking petals. As the
flowers clipped by ants look superficially similar to flowers that have naturally dehisced
their corolla, this behavior has implications for
field studies of R. macrophyllum, specifically
in the examination of seed set and pollination
success.
LITERATURE CITED
CERDÁ, X., J. RETANA, S. CARPINTERO, AND S. CROS. 1992.
Petals as the main resource collected by the ant,
Cataglyphis floricola (Hymenoptera: Formicidae).
Sociobiology 20:315–319.
GUERRANT, E.O., JR., AND P.G. FIEDLER. 1981. Flower
defenses against nectar-pilferage by ants. Biotropica
13(suppl. 2): 25–33.
HABER, W.A., G.W. FRANKIE, H.G. BAKER, I. BAKER, AND
S. KOPTUR. 1981. Ants like flower nectar. Biotropica
13:211–214.
HALVERSON, N.M. 1986. Major indicator shrubs and herbs
on national forests of western Oregon and southwestern Washington. R6-TM-229. U.S. Department
of Agriculture, Forest Service, Pacific Northwest
Region, Portland, OR.
HÖLLDOBLER, B., AND E.O. WILSON. 1990. The ants. Belknap Press of Harvard University Press, Cambridge,
MA.
POJAR, J. 1975. Hummingbird flowers of British Columbia. Syesis 8:25–28.
SCHUBART, H.O.R., AND A.B. ANDERSON. 1978. Why don’t
ants visit flowers? A reply to D.H. Janzen. Biotropica
10:310–311.
TOBIN, J.E. 1994. Ants as primary consumers: diet and
abundance in the Formicidae. Pages 278–309 in J.H.
Hunt and C. Napela, editors, Nourishment and evolution in insect societies. Westview Press, Boulder,
CO.
Received 20 November 2000
Accepted 17 October 2001
Western North American Naturalist 62(4), © 2002, p. 500
BOOK REVIEW
Birds of the Lahontan Valley: a Guide to
Nevada’s Wetland Oasis. Graham Chisholm and Larry A. Neel. University of
Nevada Press, Reno and Las Vegas. 2002.
$21.95, paperback; 256 pages + 60 illustrations, 5 maps, 2 appendixes. ISBN
0-87417-479-1.
Beyond J. Linsdale’s early publication of
Nevada’s avifauna, The Birds of Nevada (1936),
and then the much latter Birds of the Great
Basin by F. Ryser (1985) that included Nevada,
and finally Birds of Nevada by R.J. Alcorn
(1988), most other works dealing with Nevada’s
birds tend to be somewhat regional in nature.
So it is with this book. But, indeed, it is an
excellent review and summary of bird records
that have occurred in this wonderful wetlands
in far western Nevada’s Lahontan Valley.
The geography encompassed by the book
covers that region surrounding Reno and Carson City, thence eastward a bit beyond Fallon
and Stillwater. It is primarily a book on wetlands, including such locations as Walker, Pyramid, Winnemucca, Humboldt, and Carson lakes
plus the Stillwater Marshes, Carson Sinks, and
Lahontan Reservoir. These are the remnants,
totaling about 10,000 acres, of the 5.5-millionacre Pleistocene Lake Lahontan that covered
the region and that by 10,000 years ago had all
but disappeared. The authors detail a historical record of the region together with a discussion of contemporary impacts by post-European settlement—all interesting reading. They
introduce us to topics such as early Paiute
peoples there, irrigation projects that impact(ed)
wetlands, problems from mining and agriculture contamination, naturalists and ornithologists that have plied the region, and a brief
discussion of a much larger geographic region
called the Great Basin and her wetlands.
Not each bird is illustrated, but pen-andink drawings by Mimi Hoppe Wolf are liberally scattered throughout the book—and they
are delightful. Most are generally accurate and
add a pleasant touch to the reading. Overall,
the bulk of the text, pages 33–186, is an annotated account of each species from the region.
The authors have used specimen records, literature, and contemporary sight records to
round out each account.
The final section consists of birding sites
within the Lahontan Valley, including 4 maps.
This is followed by an appendix giving a yearby-year individual count, between 1986 and
1999, of colony-nesting wading birds; 1 ibis, 2
heron species, 3 egret species, and a cormorant species. Next come 2 extensive tables
giving shorebird and larid counts, by year,
during essentially the same time period. Total
counts for 26 species varied from 112,103
individuals in 1990 to as few as 10,026 in 1993.
Presumably the same methods and same degree
of search effort occurred both years.
To me, these tables are somewhat confusing,
however. Both tables give the same species, by
the same year, and from the same source (U.S.
Fish and Wildlife and Nevada Division of Wildlife) based on spring and fall counts, but Table
2 produces different numbers from Table 3.
Perhaps Table 2 is spring and Table 3 autumn
counts; but, if so, the differentiation is certainly not clear to me. Perhaps it is a problem of
table legends.
Regardless, there is a wealth of information
for birders. Anyone interested in wetlands and
wetland birds could benefit by having this book.
I recommend it, not so much for the birds, but
for all the other added information and discussion it provides at a time when all wetlands
are threatened by human encroachment.
500
Clayton M. White
Department of Zoology
Brigham Young University
Provo, UT 84602
BOOK REVIEW
Trilobite Eyewitness to Evolution. Richard
Fortey. Vintage Books of Random House,
Inc., New York. 2001. $14.00, paperback;
284 pages. ISBN 0-375-70621-6.
Why do we like what we like? How do we
choose our life’s work? What is interesting?
Richard Fortey, a museum scientist and world
authority on trilobites, explains some answers
to these questions as he looks at diverse issues
of the biological world through the eyes of
these magnificent animals. He teaches us much
about these extinct arthropods that were once
highly diverse. Fortey discusses as well the
lives and personalities of scientists with their
passions, quirks, and foibles. He considers how
he began studying these creatures and became
thoroughly involved with understanding their
lives, without having access to any living members of the group. Fortey’s humorous and engaging style brings these long-dead animals to
virtual life. This book is full of comparisons to
classical and popular literature, culture, and
everyday human experience:
When my children were young they used to
play a game with sea shells. Holding a large
whelk to one ear they contrived to “hear” the
sea: the distant crashing of waves on the shore,
or the insistent whistling of a gentle sea
breeze. Later they understood that the conch
merely amplified the murmuring of the air
around them, but they never forgot the leap
of imagination that joined shell to sea.
Palaeontology is all about listening to what
fossil shells have to say. We have to pay attention to shells . . . (p. 27).
He also introduces us to the anatomy and
purported physiology of these animals using a
few, but not too many, technical terms. He
clearly speaks as a scientist and yet doesn’t
wallow in the mystique of too many latinized
words. One clearly understands that when he
refers to Paradoxides, Agnostus, Phacops, and
Ogygiocarella, he is not trying to impress us
with his Latin training, which is proper
indeed, but rather is speaking fondly of old
friends from bygone years. We learn together
of the effort of many scientists to understand
trilobite shells, legs, crystal eyes, and pygidia.
We learn of Ken Towe working on trilobite
vision at the Smithsonian Institution:
. . . his office had a view across the grand
avenue to the FBI building. . . . Using the
trilobite lens as a substitute camera lens Ken
photographed the FBI building—not perfectly, but recognizable. What more curious
tribute to J. Edgar Hoover than to have his
workplace photographed through the eyes of
an ancient fossil (p. 106)!
The magnificent soft-bodied fossils of the
Burgess Shale are visited in a discussion of the
purported “Cambrian explosion” of animal
diversification that might have occurred there.
He properly credits Charles Doolittle Walcott,
Harry Whittington, Derek Briggs, Simon Conway Morris, and others for the detailed work
they did in these strata, as well as the popularization made by Stephen J. Gould. He makes
no bones, however, that, based on his collaborations with Briggs, this “explosion” was an
orderly one in which no special interpretations of phylogeny were required to produce a
tree summarizing relationships. Still, more
explanation is needed:
So now we are left with a paradox. There is a
tree of descent which helps us understand
the history of our characters before their
spectacular appearance on stage—but of this
earlier history there is no evidence. Even the
traces left by animals, their scratches and
burrows, are rare before the latest Precambrian. Where can these animals be? . . . My
favoured theory is that the earlier branches
in the tree were tiny animals, which were not
easily preserved as fossils (pp. 138–139).
The book has scattered diagrams, figures,
and many photographs of trilobites, their
parts, and lore, but the book’s small size and
rough ivory paper detract ever so slightly from
the beautiful content they attempt to convey.
501
502
Readers interested more in artistic photographs of these creatures would be better off
obtaining Riccardo Levi-Setti’s 1975 or 1993
book Trilobites. Still, the photos in Fortey’s
book are usable and impressive.
He continues with a concise, yet informative, treatment of the need for taxonomists,
systematists, and museums. The next chapter
deals with the life history stages of individual
species of trilobites and the scientists that
revealed them to us. Additional chapters follow a discussion of their decline and disappearance from the fossil record near the end
of the Permian. They left with a whimper
rather than a bang, just prior to the largest
extinction event the world has ever seen.
In discussing “Possible Worlds,” Dr. Fortey
traces his (and others’) efforts to reconstruct
the history of continents and seas for the 300
or so million years that were witnessed by
trilobites during their tenure on earth. He
then attempts a prediction at what else these
creatures might teach us of themselves and
life in general. Could we better approach key
[Volume 62
issues of contemporary biology and environmentalism from a novel viewpoint? Could the
trilobites reveal what they “learned” during
their long history? Near the end of the book
he waxes justifiably poetic:
Will we be able to see the mysteries of this
time of innovation explained, as once Walcott
surveyed the mysteries of the trilobite limb?
These are not Gradgrind facts, “dull realities”:
they are wings for flights of the imagination.
I wish I could live long enough to know, and
even if I did, I should never cry “enough” (p.
263).
Oh, that all could be so excited about some
aspect of their life! I am a better person for
having heard the passion and viewpoints of
Richard Fortey in this book.
C. Riley Nelson
Department of Integrative Biology
Provo, UT 84602
Western
North American
Naturalist
INDEX
Volume 62—2002
INDEX
Volume 62—2002
AUTHOR INDEX
Agenbroad, Larry D., 129
Allphin, Loreen, 423
Alvarez-Castañeda, Sergio Ticul, 127, 496
Ames, Abbe D., 240
Applegate, Roger D., 227
Giordano, Mark R., 341
Goad Henry, Susanna, 124
Goertz, J.W., 487
Goldberg, Stephen R., 160, 243, 491
Gul, Bilquees, 101
Bartolome, James W., 73
Bateman, Terry A., 316
Baumann, R.W., 484
Beaty, Barry J., 120
Belthoff, James R., 112
Betancourt, Julio L., 348, 405
Bock, Carl E., 370
Bolling, John D., 88
Bosworth III, William R, 451
Boudell, Jere A., 14
Bourassa, Jean B., 240
Breitbarth, Jessica H., 437
Bryan, Scott D., 197
Bursey, Charles R., 160, 243, 491
Busse, M.D., 141
Butcher, Laurence R., 365
Hall, Linnea S., 370
Harris, Charles E., 151
Heath, James E., 437
Heath, Maxine S., 437
Holechek, Jerry, 300
Huffman, David W., 474
Calisher, Charles H., 120
Carraway, Leslie N., 458, 487
Cartron, Jean-Luc E., 249
Christoferson, Laurel L., 370
Clark, William H., 230
Cortés-Calva, Patricia, 496
Crane, Kenneth A., 240
Cruz, Alexander, 479
Douglass, John F., 253
Dunbar, Mike R., 341
Ingersoll, Thomas E., 124
Inouye, Richard S., 360
Jackson, Stephen T., 405
Jacobi, G.Z., 484
Jass, Christopher N., 129
Jehl, Jr., Joseph R., 335
Jensen, Mark E., 257
Johansen, Jeffrey R., 14
Johansson, Carl, 335
Jones, Cheri A., 120
Karp, Catherine A., 106
Kato, Thomas T., 151
Kaushal, Sujay S., 246
Kelly, Brian T., 151
Khan, M. Ajmal, 101
King, R. Andrew, 112
Kippenhan, Michael G., 381
Kolb, Thomas E., 266
Kondratieff, Boris C., 59, 385
Koopman, Marni E., 151
Evans, Howard E., 206
Fehmi, Jeffrey S., 73
Felix, Todd A., 240
Fischer, Karen N., 479
Fisher, Stuart G., 466
Fleury, Scott A., 365
Frey, Jennifer K., 120
Gaddis, Stanley E., 240
Galt, Dee, 300
Galuszka, Donna M., 266
Leatherman, David A., 206
Leavitt, Steven W., 348
Leberg, Paul L., 32
Lechleitner, Richard A., 385
Lester, Gary T., 230
Lindstrom, Robert F., 44
Link, Steven O., 14
Linz, George M., 39
López González, Carlos A., 218
Lutz, R. Scott, 227
Lyford, Mark E., 405
504
2002]
INDEX
Martin, John, 370
McCann, Geraldine R., 240
Mead, Jim I., 129
Meadows, Dwayne W., 377
Medlyn, David A., 210
Meik, Jesse M., 234
Meretsky, Vicky J., 307
Miller, Brian J., 218
Molinar, Francisco, 300
Molles, Jr., Manuel C., 249
Morrison, Amy D., 129
Morrison, Michael L., 370
Mueller, Gordon, 106
Mulcahy, Daniel G., 234
Myers, Marilyn J., 1
Naegeli, Markus, 466
Naumann, Tamara, 414
Navo, Kirk W., 124
Nelson, C. Riley, 383, 501
Nonacs, Peter, 188
North, Eric G., 307
Nowak, Robert S., 327
O’Farrell, Thomas P., 151
Oliver, George V., 451
Olson, Richard A., 414
Pedicino, Lisa C., 348
Perryman, Barry L., 414
Petersburg, Stephen, 414
Petersen, Brett E., 240
Pierce, John R., 257
Prather, John W., 479
Ramaley, Robert F., 44
Reed, J. Michael, 365
Resh, Vincent H., 1
Rhodes, Howard A., 59
Rich, Terrell D., 288
Riegel, G.M., 141
Rissler, Peter H., 82
Robinson, Anthony T., 197
Root, J. Jeffrey, 120
505
Rosenberg, Daniel K., 280
Rychert, Robert C., 223
Sanborn, Allen F., 437
Scoppettone, G. Gary, 82
Setser, Kirk, 234
Shumake, Stephen A., 240
Smith, James F., 316
Smith, Stanley D., 327
Sterner, Ray T., 240
Stevens, Lawrence E., 307
Stockwell, Craig A., 32
Stromberg, Juliet C., 170
Strong, Thomas R., 370
Sturm, Ken K., 280
Svejcar, T.J., 141
Sweetser, Michael G., 197
Thomas, Milton, 300
Tidwell, William D., 210
Titus, Jonathan H., 327
Titus, Priscilla J., 327
Twedt, Daniel J., 39
Uehlinger, Urs, 466
Valenzuela, Michael, 25
Van de Water, Peter K., 348
Verts, B.J., 458, 487
Walker, Lawrence R., 88
Walsh, John J., 246
Weber, Darrell J., 101
Weiser, Michael D., 498
Weiss Bizzoco, Richard L., 44
White, Clayton M., 254, 255, 500
Wilkinson, Jack A., 253
Williams, Christopher K., 227
Wilson, Paul, 25
Windham, Michael D., 423
York, Melissa M., 280
Zoellick, Bruce W., 151
Zuellig, Robert E., 59
KEY WORD INDEX
Taxa described as new to science appear in boldface type.
Acanthocephala, 160
Acer negundo, 266
acid, 44
adaptation
local, 32
thermal, 437
adaptive management, 32
age structure, 170
allelopathy, 141
Alnus oblongifolia, 266
ambersnail, 307
American Southwest, 484
Amphipoda, 230
amphipods, 1
ants, 188
Anura, 160
aquatic macrophyte vegetation,
257
aquatic plant communities, 257
Araucarioxylon, 210
arbuscular mycorrhizae, 327
Arizona, 243, 266, 370
northern, 348
Artemisia tridentata, 14
506
artificial burrows, 112
Athene cunicularia, 112, 280
atmospheric carbon dioxide,
348
coyote, 341
Crenichthys, 82
cryptobiotic crusts, 223
cryptogamic crusts, 14
Baja California Sur, [Mexico],
127
banding, 124
behavior, 365
mobbing, 253
benthic, 59
birds, 246
Black Hills, [South Dakota], 129
body weight, 188
breeding
biology, 479
birds, 370
Brewer’s Sparrow, 288
Brian Head mountainsnail, 451
Bromus tectorum, 14
brush control, 300
Bufo
kelloggi, 160
mazatlanensis, 160
bullsnake, 240
Burrowing Owl, 112, 280
burrows, 327
dendrochronology, 266
desert
springs, 1
stream, 466
diel oxygen curve, 466
diet, 249, 280
Dipogon, 206
discrimination, isotopic, 348
distribution(s), 120, 197, 206,
234, 487
disturbance, 88, 466
diversity, community, 414
caddisflies, 1
California, 73, 280, 487
Imperial Valley, 280
Mono Lake, 335
San Joaquin Valley, 151
Canis latrans, 253, 341
Capnia caryi, 484
Capniidae, 484
carbon isotopes, 348
cattle, 300
caves, 124
Ceanothus fendleri, 474
central Wyoming, 405
Cestoda, 160, 491
Chaetodipus baileyi, 496
character-count method, 25
chlorophyll, 223
cicadas, 437
Cicindela, 381
climate, 266
Coleoptera, 381
Colinus virginianus, 227
Colorado, 59, 120, 479
North Fork Cache la Poudre
River, 59
Plateau, 307
River, 82
Colubridae, 243
communication, 437
community diversity, 414
contamination, 44
Corophium, 230
[Volume 62
habitat
associations, 234
fragmentation, 377
use, 365
halophytes, 101
health, 341
helminths, 160, 243, 491
hematology, 341
herbarium specimens, 348
herbicides, 300
herpetofauna, 234
heterozygosity, 423
home range, 151
hot spring, 44
howling survey, 341
hybridization, 25
Hymenoptera, 206, 474
Eared Grebe, 335
ecological
classification, 257
restoration, 474
ecophysiology, 348
Empetrichthyidae, 82
endemic species, 1
Erigeron, 423
Eriogonum shockleyi, 316
ethephon, 101
Eurytoma squamosa, 474
extinction, 496
Idaho, 288
southwestern, 112
Imperial Valley, [California],
280
inbreeding, 32
individual variation, 288
introduced species, 230
islands, 496
isotopes, 246
isotopic discrimination, 348
ISSR, 316
fire recovery, 223
fleabane, 423
flooding, 227
floods, 170
foliar nutrients, 141
food habits, 218
food-niche breadth, 280
foraging, 188
forest regeneration, 141
Formica, 188
Formicidae, 498
fossil wood, 210
Fraxinus velutina, 266
fusicoccin, 101
jaguar, 218
Juglans major, 266
juniper woodlands, 405
gastropods, 451
gene flow, 423
genetic bottleneck, 32
genetic variability, 423
geographic variation, 288, 458
gibberellic acid, 101
Goodeidae, 82
granivores, 246
grassland management, 73
grazing, 300
Great Basin, 234
northern, 458
Great Salt Lake, [Utah], 335
Gulf of California, [Mexico],
249
kinetin, 101
kit fox, 127
Larrea tridentata, 88
late Holocene, 405
Late Pleistocene, 129
leaves, 348
Lepidomeda vittata, 197
Leptodactylus melanonotus,
160
Little Colorado spinedace, 197
local adaptation, 32
longevity, 124
macroinvertebrate, 59
Mammoth Site, [South Dakota],
129
mass, 39
Mesembrioxylon, 210
Mexican vole, 120
Mexico, 160, 496
Baja California Sur, 127
Gulf of California, 249
microbe, 44
microhabitat use, 365
microorganism, 44
2002]
Microtus
californicus, 73
mexicanus, 120
mogollonensis, 120
migration, 335
mobbing behavior, 253
Mogollon vole, 120
Mojave Desert, 327
mollusks, 129, 451
molt, 39
monitoring, 335
Mono Lake, [California], 335
morphology, 39
Morrison Formation, [Wyoming],
210
mortality, 227
Mount Rainier National Park,
[Washington], 385
movement(s), 151, 197, 377
Mugil cephalus, 249
mycorrhizal inoculum potential,
327
Myotis, 124
native grasses, 73
natural invasions, 405
nectivory, 498
Nelson-Wittenberg Site, [South
Dakota], 129
Nematoda, 160, 491
nest-site selection, 112, 479
Neurotrichus gibbsii, 487
Nevada, 381
River Mountains, 327
White River, 82
Snake Range, 234
Nevada Test Site, 327
North Dakota, 39
North Fork Cache la Poudre
River, [Colorado], 59
northern Arizona, 348
northern Great Basin, 458
Northern Bobwhite, 227
Odocoileus hemionus, 253
oil-field development, 151
Okanagana
striatipes, 437
utahensis, 437
open-hole index, 240
Oregon, 341
Oreohelix
parawanensis, 451
peripherica wasatchensis,
377
Osprey, 249
outbreeding, 32
Oxyloma, 307
INDEX
Pachymedusa dacnicolor, 160
paleoenvironments, 129
Pandion haliaetus, 249
Panthera onca, 218
parasitism, 474
Parowan Mountains, [Utah],
451
pelage color, 458
Penstemon, 25
Peromyscus truei, 458
physiology, 341
Pinus jeffreyi, 141
piñon mice, 458
Piranga ludovicana, 479
plant community comparison,
73
plant-insect interactions, 474
Platanus wrightii, 170, 266
Plecoptera, 1, 375, 484
pocket gopher, 240
Podiceps nigricollis, 335
Pogonomyrmex, 188
point frame, 360
pollination, 498
pollinator, 498
Polygonaceae, 316
Pompilidae, 206
population(s), 316, 335
estimate, 82
trends, 370
Populus fremontii, 266
predispersal seed predation,
474
predation, 197, 240
predator attacked by prey, 253
prescribed fire, 414
prey, 206, 218
radio-tracking, 240
ramet, 170
Rana
forreri, 160
magnaocularis, 160
range expansion, 474
range extension, 487
rangeland, 300
rapid evolution, 32
rare plants, 316
razorback sucker, 106
recovery, 59
regeneration, 266
reintroduction, 32
revegetation, 88
riparian, 266, 370
habitat, 170
River Mountains, [Nevada],
327
road avoidance, 377
Rodentia, 496
507
sagebrush, 288
steppe, 360
Salicornia rubra, 101
Salix, 266
salmonids, 197
sampling effort, 360
San Juan River inflow, 106
San Joaquin kit fox, 151
San Joaquin Valley, [California],
151
seasonality, 327
sediment, 59
respiration, 466
stability, 466
seed
banks, 14
dormancy, 101
seedling
establishment, 170
growth, 141
shrew-mole, 487
shrub-steppe, 14
Smilisca baudini, 160
snail, 377
Snake Range, [Nevada], 234
Snake River, 230
Snake River Birds of Prey Conservation Area, 112
Snake River Plain, 223, 288
soil
chemical, 141
microbial biomass, 141
recovery, 88
song, 437
dialects, 288
South Dakota
Black Hills, 129
Mammoth Site, 129
Nelson-Wittenberg Site, 129
southwestern Idaho, 112
southwestern USA, 348
spatial organization, 151
Spea
bombifrons, 491
intermontana, 491
species
endemic, 1
richness, 414
spiders, 206
Spizella breweri, 288
springfish, 82
stonefly(ies), 385, 484
stream
flow, 266
metabolism, 466
Strongylura, 249
succession, 88, 141
Succineidae, 307
sulphur, 44
508
swarming, 124
synecological study, 257
taxonomy, 307
temperature, 437
Thamnophis cyrtopsis, 243
thermal adaptation, 437
thiourea, 101
tiger beetles, 381
tree ring, 266
Trematoda, 160, 491
trend analysis, 39
trophic level, 246
Tylosurus, 249
Upper Jurassic, 210
Utah, 377, 451
Great Salt Lake, 335
Parowan Mountains, 451
variety, 316
vegetation
classification, 257
cover, 414
history, 405
vegetative cover, 360
Vulpes
macrotis mutica, 151
velox, 127
warm springs, 82
Washington, 385
Mount Rainier National
Park, 385
[Volume 62
Western Tanager, 479
White River, [Nevada], 82
woodpecker, 365
woodrat middens, 405
Wyethia mollis, 141
Wyoming, 210
central, 405
Morrison Formation, 210
Yellowstone National Park,
44
Xanthocephalus xanthocephalus,
39
Xyrauchen texanus, 106
Yellow-headed Blackbird, 39
Yellowstone National Park,
[Wyoming], 44
TABLE OF CONTENTS
Volume 62
No. 1—January 2002
Articles
Trichoptera and other macroinvertebrates in springs of the Great Basin: species composition, richness,
and distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marilyn J. Myers and Vincent H. Resh
1
Effect of soil microtopography on seed bank distribution in the shrub-steppe . . . . Jere A. Boudell,
Steven O. Link, and Jeffrey R. Johansen
14
Three naturally occurring Penstemon hybrids . . . . . . . . . . . . . . Paul Wilson and Michael Valenzuela
25
Ecological genetics and the translocation of native fishes: emerging experimental approaches . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Craig A. Stockwell and Paul L. Leberg
32
Morphometric changes in Yellow-headed Blackbirds during summer in central North Dakota . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Daniel J. Twedt and George M. Linz
39
Invisible invasion: potential contamination of Yellowstone hot springs by human activity . . . . . . . . .
. . . . . . . . . . . . . . . . . . Robert F. Lindstrom, Robert F. Ramaley, and Richard L. Weiss Bizzoco
44
Benthos recovery after an episodic sediment release into a Colorado Rocky Mountain river . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . Robert E. Zuellig, Boris C. Kondratieff, and Howard A. Rhodes
59
Species richness and California voles in an annual and a perennial grassland . . . . . Jeffrey S. Fehmi
and James W. Bartolome
73
Status of the Preston White River springfish (Crenichthys baileyi albivallis) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. Gary Scoppettone and Peter H. Rissler
82
Fertile island development around perennial shrubs across a Mojave Desert chronosequence . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . John D. Bolling and Lawrence R. Walker
88
Improving seed germination of Salicornia rubra (Chenopodiaceae) under saline conditions using
germination-regulating chemicals . . . . . . M. Ajmal Khan, Bilquees Gul, and Darrell J. Weber
101
2002]
INDEX
Razorback sucker movements and habitat use in the San Juan River inflow, Lake Powell, Utah,
1995–1997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Catherine A. Karp and Gordon Mueller
Nest-site characteristics of Burrowing Owls (Athene cunicularia) in the Snake River Birds of Prey
National Conservation Area, Idaho, and applications to artificial burrow installation . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . James R. Belthoff and R. Andrew King
509
106
112
Notes
New records of the Mogollon vole, Microtus mogollonensis (Mearns 1890), in southwestern Colorado
. . . . . Jennifer K. Frey, J. Jeffrey Root, Cheri A. Jones, Charles H. Calisher, and Barry J. Beaty
120
Observations of swarming by bats and band recoveries in Colorado . . . . . . . . . . . . . . Kirk W. Navo,
Susanna Goad Henry, and Thomas E. Ingersoll
124
Noteworthy record of the kit fox (Mammalia: Canidae: Vulpes velox macrotis) in Vizcaino Desert,
Baja California Sur, México . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sergio Ticul Alvarez-Castañeda
127
No. 2—April 2002
Articles
Late Pleistocene mollusks from the southern Black Hills, South Dakota . . . . . Christopher N. Jass,
Jim I. Mead, Amy D. Morrison, and Larry D. Agenbroad
129
Does the presence of Wyethia mollis affect growth of Pinus jeffreyi seedlings? . . . . . . G.M. Riegel,
T.J. Svejcar, and M.D. Busse
141
Movements and home ranges of San Joaquin kit foxes (Vulpes macrotis mutica) relative to oil-field
development . . . . . . . . . . . . . . . . . . . . . . . Bruce W. Zoellick, Charles E. Harris, Brian T. Kelly,
Thomas P. O’Farrell, Thomas T. Kato, and Marni E. Koopman
151
Helminth parasites of seven anuran species from northwestern Mexico . . . . . Stephen R. Goldberg
and Charles R. Bursey
160
Flood flows and population dynamics of Arizona sycamore (Platanus wrightii) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Juliet C. Stromberg
170
Patterns of energy allocation within foragers of Formica planipilis and Pogonomyrmex salinus . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Peter Nonacs
188
Movement, distribution, and predation: Lepidomeda vittata and nonnative salmonids in eastern
Arizona . . . . . . . . . . . . . . . . . . Michael G. Sweetser, Scott D. Bryan, and Anthony T. Robinson
197
New records and range extensions of species of Dipogon (Hymenoptera, Pompilidae) in Colorado . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Howard E. Evans and David A. Leatherman
206
Mesembrioxylon obscurum, a new combination for Araucarioxylon? obscurum Knowlton, from the
Upper Jurassic Morrison Formation, Wyoming . . . . David A. Medlyn and William D. Tidwell
210
Do jaguars (Panthera onca) depend on large prey? . . . . . . . . . . . . . . . . . . Carlos A. López González
and Brian J. Miller
218
Assessment of cryptobiotic crust recovery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Robert C. Rychert
223
Notes
The effect of flooding on Northern Bobwhites . . . . . . . . . . . . . . . . . . . . . . . . . . Roger D. Applegate,
Christopher K. Williams, and R. Scott Lutz
227
Occurrence of Corophium spinicorne Stimpson, 1857 (Amphipoda: Corophiidae) in Idaho, USA . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gary T. Lester and William H. Clark
230
510
[Volume 62
Herpetofauna of the southern Snake Range of Nevada and surrounding valleys . . . . . . Kirk Setser,
Jesse M. Meik, and Daniel G. Mulcahy
234
Movements of a bullsnake (Pituophis catenifer) following predation of a radio-collared northern
pocket gopher (Thomomys talpoides) . . . . . . . . . . . . . . . . . . . Ray T. Sterner, Brett E. Petersen,
Stephen A. Shumake, Stanley E. Gaddis, Jean B. Bourassa, Todd A. Felix,
Geraldine R. McCann, Kenneth A. Crane, and Abbe D. Ames
240
Gastrointestinal helminths of the blackneck garter snake, Thamnophis cyrtopsis (Colubridae) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stephen R. Goldberg and Charles R. Bursey
243
Comparison of nitrogen isotope ratios in feathers from seven species of Colorado breeding birds . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sujay S. Kaushal and John J. Walsh
246
Osprey diet along the eastern side of the Gulf of California, Mexico . . . . . . . . Jean-Luc E. Cartron
and Manuel C. Molles, Jr.
249
Mule deer group kills coyote . . . . . . . . . . . . . . . . . . . . . . . . Jack A. Wilkinson and John F. Douglass
253
Book Reviews
Bird Hand Book
Victor Schrager and A.S. Byatt . . . . . . . . . . . . . . . . . . . . . . . . . Clayton M. White
The World of the Hummingbird
Robert Burton . . . . . . . . . . . . . . . . . . . . . . . . . Clayton M. White
254
255
No. 3—July 2002
Articles
A classification of aquatic plant communities within the Northern Rocky Mountains . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . John R. Pierce and Mark E. Jensen
257
Tree growth and regeneration response to climate and stream flow in a species-rich southwestern
riparian forest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Donna M. Galuszka and Thomas E. Kolb
266
Diet and food-niche breadth of Burrowing Owls (Athene cunicularia) in the Imperial Valley,
California . . . . . . . . . . . . . . . . . . . . . Melissa M. York, Daniel K. Rosenberg, and Ken K. Sturm
280
The short song of Brewer’s Sparrow: individual and geographic variation in southern Idaho . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Terrell D. Rich
288
Soil depth effects on Chihuahuan Desert vegetation . . . . . . . . . Francisco Molinar, Jerry Holechek,
Dee Galt, and Milton Thomas
300
Kanab ambersnail and other terrestrial snails in south central Utah . . . . . . . . . . Vicky J. Meretsky,
Eric G. North, and Lawrence E. Stevens
307
Genetic differentiation of rare and common varieties of Eriogonum shockleyi (Polygonaceae) in
Idaho using ISSR variability . . . . . . . . . . . . . . . . . . . . . . . James F. Smith and Terry A. Bateman
316
Arbuscular mycorrhizae of Mojave Desert plants . . . . . . . . . . . Jonathan H. Titus, Priscilla J. Titus,
Robert S. Nowak, and Stanley D. Smith
327
Autumnal migration of Eared Grebes (Podiceps nigricollis) through southwestern Wyoming: a key
to assessing the size of the North American population . . . . . . . . . . . . . . . . . Joseph R. Jehl, Jr.,
and Carl Johansson
335
Abundance and condition indices of coyotes on Hart Mountain National Antelope Refuge, Oregon
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mike R. Dunbar and Mark R. Giordano
Historical variations in δ13Cleaf of herbarium specimens in the southwestern U.S. . . . . . . . . . . . . .
. . . . . . . . Lisa C. Pedicino, Steven W. Leavitt, Julio L. Betancourt, and Peter K. Van de Water
348
Sampling effort and vegetative cover estimates in sagebrush steppe . . . . . . . . . . Richard S. Inouye
360
341
2002]
INDEX
Orientation and vertical distribution of Red-naped Sapsucker (Sphyrapicus nuchalis) nest cavities . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Laurence R. Butcher, Scott A. Fleury, and J. Michael Reed
511
365
Notes
Bird populations in riparian areas of southeastern Arizona in 1985–86 and 1994–95 . . . . . . . . . . . . .
. . . . . . . . . . . . . . Linnea S. Hall, Michael L. Morrison, Laurel L. Christoferson, John Martin,
Carl E. Bock, and Thomas R. Strong
370
The effect of roads and trails on movement of the Ogden Rocky Mountain snail (Oreohelix peripherica wasatchensis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dwayne W. Meadows
377
A new state record of Cicindela nigrocoerulea nigrocoerulea LeConte (Coleoptera: Cicindelidae) in
Nevada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Michael G. Kippenhan
381
Book Review
The Geology of the Parks, Monuments, and Wildlands of Southern Utah Robert Fillmore . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. Riley Nelson
383
No. 4—October 2002
Articles
Stoneflies (Plecoptera) of Mount Rainier National Park, Washington . . . . . . . . . . . B.C. Kondratieff
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . and Richard A. Lechleitner
385
A 4000-year record of woodland vegetation from Wind River Canyon, central Wyoming . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . Stephen T. Jackson, Mark E. Lyford, and Julio L. Betancourt
405
Vegetation response to prescribed fire in Dinosaur National Monument . . . . . . Barry L. Perryman,
Richard A. Olson, Stephen Petersburg, and Tamara Naumann
414
Morphological and genetic variation among populations of the rare Kachina daisy (Erigeron kachinensis) from southeastern Utah . . . . . . . . . . . . . . . . . . . . Loreen Allphin and Michael D. Windham
423
Temperature responses and habitat sharing in two sympatric species of Okanagana (Homoptera:
Cicadoidea) . . . Allen F. Sanborn, Jessica H. Breitbarth, James E. Heath, and Maxine S. Heath
437
Oreohelices of Utah, II. Extant status of the Brian Head mountainsnail, Oreohelix parawanensis
Gregg, 1941 (Stylommatophora: Oreohelicidae) . . . . George V. Oliver and William R. Bosworth III
451
Geographic variation in pelage color of piñon mice (Peromyscus truei) in the northern Great Basin
and environs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leslie N. Carraway and B.J. Verts
458
A heterotrophic desert stream? The role of sediment stability . . . . . . . . . . . . . . . . . . Urs Uehlinger,
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Markus Naegeli, and Stuart G. Fisher
466
A seed chalcid (Eurytoma squamosa Bugbee) parasitizes buckbrush (Ceanothus fendleri Gray)
seeds in a ponderosa pine forest of Arizona . . . . . . . . . . . . . . . . . . . . . . . . . . David W. Huffman
474
Nest site characteristics and reproductive success of the Western Tanager (Piranga ludoviciana) on
the Colorado Front Range . . . . . . . . . . Karen H. Fischer, John W. Prather, and Alexander Cruz
479
Capnia caryi, an interesting new species of winter stonefly from the American Southwest
(Plecoptera: Capniidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R.W. Baumann and G.Z. Jacobi
484
Notes
Distribution of Neurotrichus gibbsii in California with a range extension in the Sierra Nevada . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leslie N. Carraway, B.J. Verts, and J.W. Goertz
487
Helminths of the plains spadefoot, Spea bombifrons, the western spadefoot, Spea hammondii, and
the Great Basin spadefoot, Spea intermontana (Pelobatidae) . . . . . . . . . . Stephen R. Goldberg
and Charles R. Bursey
491
512
[Volume 62
Extirpation of Bailey’s pocket mouse, Chaetopidus baileyi fornicatus (Heteromyidae: Mammalia),
from Isla Montserrat, Baja California Sur, Mexico . . . . . . . . . . Sergio Ticul Alvarez-Castañeda
and Patricia Cortés-Calva
496
Removal of Rhododendron macrophyllum petals by Camponotus modoc . . . . . . . Michael D. Weiser
498
Book Reviews
Birds of the Lahontan Valley: a Guide to Nevada’s Wetland Oasis
Graham Chisholm and Larry
A. Neel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clayton M. White
500
Trilobite Eyewitness to Evolution
Richard Fortey . . . . . . . . . . . . . . . . . . . . . . . . . C. Riley Nelson
501
Index to Volume 62 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
503
Introducing . . .
Monographs of the
VOLUME 1
_____________
The monograph comprises the following 3 articles:
Mammals of the Grand Staircase–Escalante National Monument
A literature & museum survey
Jerran T. Flinders, Duke S. Rogers, Jackee L. Webber, Harry A. Barber
________________________________
Stoneflies (Plecoptera) of southern Utah with an updated
checklist of Utah species
Ronald G. Call, Richard W. Baumann
________________________________
Annotated checklist of the millipeds of California
(Arthropoda: Diplopoda)
Rowland M. Shelley
________________________________
Standard price $20
$16 per copy with this offer
This monograph is in the final stages of publication. A complimentary copy
will be sent to each subscriber upon completion of printing. Additional copies
can be purchased at the $16 pre-publication price. Please make checks
payable to Western North American Naturalist and send to M.L. Bean Life Science Museum, Brigham Young University, Provo UT 84602.

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