Great Basin Naturalist

Volume 49 Number 2 Article 5

4-30-1989

Species composition, emergence, and habitat preferences of Trichoptera of the Sagehen Creek Basin, , USA

Nancy A. Erman University of California, Berkeley

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Recommended Citation Erman, Nancy A. (1989) "Species composition, emergence, and habitat preferences of Trichoptera of the Sagehen Creek Basin, California, USA," Great Basin Naturalist: Vol. 49 : No. 2 , Article 5. Available at: https://scholarsarchive.byu.edu/gbn/vol49/iss2/5

This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. SPECIES COMPOSITION, EMERGENCE, AND HABITAT PREFERENCES OF TRICHOPTERA OF THE SAGEHEN CREEK BASIN, CALIFORNIA, USA

Nancy A. Erman 1

Abstract. —An extensive study of larvae and adult Trichoptera of the Sagehen Creek basin. Sierra Nevada, California, USA, revealed 77 species representing 14 families and 41 genera. Twenty-six species were restricted to small water bodies (spring sources, seeps, spring streams, temporary ponds, and intermittent streams); 27 were restricted to Sagehen Creek, a second-order stream, and the mouths of two large spring streams. Similarity between species caught at black lights and those in emergence traps was 43%. There were two major peaks in adult emergence by species, midsummer and late summer-fall. A few species emerged during winter or throughout most of the year. The species composition of the community showed affinity with Oregon, the Great Basin, and the Rocky Mountains but very little similarity with the California Coast Range. Four of the most abundant species in the basin have very restricted distributions. Ecological separation of several groups of closely related species could be explained by major differences in larval habitats or by different emergence periods.

Trichoptera from the Sierra Nevada of Cali- The primary objectives of the present study fornia have been collected and described in were to determine the species composition, papers over the last several decades, but no emergence periods, and habitat preferences comprehensive studies of species abundance of the Trichoptera community of the Sagehen and distribution exist for the Sierra Nevada or Creek basin. As the study progressed, sec- for any part of it. The Sagehen Creek basin on ondary objectives emerged, such as a com- the east side of the Sierra Nevada has been the parison of collecting methods, development site of a University of California field station of emergence collection techniques for re- and of numerous aquatic biology studies since mote areas, and examination of some taxo- 1951. Trichoptera from this watershed have nomic problems. been critical to several svstematic (i.e., Den- ning Wiggins 1970, 1973, 1977, Parker and Study Area Wiggins 1985) and behavioral studies (Erman

1981, 1984, 1986, 1987). Other aquatic inver- Sagehen Creek basin is on the east side of tebrate work in the Sagehen Creek basin is the northern Sierra Nevada, Nevada and extensive. A comprehensive list of the stone- Sierra counties, California, and on the west- flies was published by Sheldon and Jewett ern edge of the Great Basin in the Lahontan

(1967) and updated by R. Baumann, W. Shep- drainage. It includes 2,700 hectares from its ard, B. Stark, and S. Szczytko for the first headwaters (elevation 2,256 m) to its end in North American Plecoptera Conference in Stampede Reservoir (elevation 1,804 m).

1985 (unpublished). Plecoptera material from Mean annual precipitation is 93 cm, most of the Sagehen Creek basin has contributed to which falls as snow. Mean annual temperature many systematic (Jewett 1966, Surdick 1981, at the field station measuring station (1,943 m) Szczytko and Stewart 1979, 1984) and ecologi- is 4.9 C, and temperatures below freezing can cal studies (Sheldon 1969, 1972, 1980). occur in any mouth. Within the Sagehen Creek

The Turbellaria have been studied (Kenk basin is a wide diversity of aquatic habitats. 1970, 1972) as has the amphipod genus Stygo- In addition to the second-order, spring-fed bromas (Holsinger 1974); and the Cricotopus- Sagehen Creek, there are many permanent, Nostoc relationship was first described in constant-temperature springs (3.5-9 C) and Sagehen Creek (Brock 1960). The aquatic in- spring streams of various sizes and physical- vertebrate community and secondary produc- chemical conditions, several minerotrophic tion in peatlands have been examined (Erman peatlands (fens) in different stages of evolution and Erman 1975). (Erman 1976, Bartolome et al. in press), a

Department of Forestry and Resouree Management. Universitj ol < ialifornia, Berkeley. California 91720

186 April 1989 Erman: California Trichoptera 187

small cirque pond at the head of the basin that solved have been or will be sent to the Royal dries completely about two out of three years, Ontario Museum, Toronto, Canada. and temporary streams that exist each year for varying time periods. Results and Discussion Species Habitats and Distribution Methods A total of 77 species representing 14 families and 41 genera were collected in the

Adults were collected with emergence basin (Table 1). was the most traps, black lights, hand nets, and fish traps. abundantly represented family with 32 spe- Emergence traps were of two types: one, a cies, followed by with 13 spe- 1 X 1-m base pyramidal trap with collecting cies. Larval habitats were determined for bottle (80% alcohol) that was emptied at one- most species and are given in Table 1. Habi- or two-week intervals; the other, a smaller tats could not be determined for 12 species folding trap that could be easily moved to caught only by black light and for whom larvae more remote sites. During the six years of the were unknown. study over 800 one- or two-week emergence Twenty-six species occurred in one small trap samples were collected in 22 locations in water body or a combination of spring the basin, including spring sources, spring sources, spring seeps, spring streams, tempo- streams (first-order streams), an intermittent rary ponds, and intermittent streams but not stream, and Sagehen Creek. Fish traps, used in Sagehen Creek. At least one-third of the in other studies, were closed screen boxes, species in the basin, then, were restricted to open only on the ends. They extended above smaller aquatic habitats. Conversely, at least the water and were emptied from the hinged 27 other species, approximately another one- top (see Erman and Leidy 1975). They were third, were confined to Sagehen Creek and/or useful for collecting both larval and adult cad- near the mouths of the two largest spring disflies. Black lights were used in the vicinity streams only. Twelve species were found in of the U.C. field station. One black light was both general habitat groups, though they may run intermittently in the same location (about have shown a preference for one or the other. 30 m from the nearest water) at the field sta- Fifteen species from six families showed a tion from as early as April to as late as October variety of adaptations for life in variable habi- from 1980 to spring 1987. This light was not a tats at the land-water interface and are dis- trap; specimens were sampled at approxi- cussed in more detail elsewhere (Erman 1981, mately half-hour to one-hour intervals on 111 1986, 1987). Larvae of Clostoeca disjuncta nights. Other portable black lights were used were never found in permanent water. Adults in remote areas. Hand-netting and sweep- of designatus and Wormaldia netting of vegetation were done at many pachita emerged from an intermittent stream aquatic sites in the basin. just days before it dried completely. Lim- Larvae were collected with kick screens, nephilus peltus left permanent spring streams Surber-type samplers, and scoops. They were and pupated in damp moss; in laboratory rear- also collected by hand-picking. The objective ing studies it could not emerge if left in per- of the combination of sampling techniques manent water. A few species (i.e., Goeracea was to collect, if possible, all Trichoptera spe- oregona, Allomyia cidoipes, Lepidostoma er- cies in the basin and to determine the larval manae) were limited to one or a few springs.

habitats of all species collected. To that end, The distribution of the species in springs and many larvae were also reared. Quantitative their requirements will be discussed further sampling was not an objective; however, for in a future paper, the result of a separate study comparative purposes, the emergence traps on Sierra Nevada springs. provided data on relative abundance of spe- Species Abundance cies. A reference collection of most species will Relative abundance of species given in be placed in the California Academy of Sci- Table 1 is based roughly (a) on number of ences, Golden Gate Park, San Francisco. adults caught during the study, (b) on distri- Some of the more rare species and those for bution in numbers of habitats based on emer- which taxonomic questions have yet to be re- gence trapping, and (c) on an assessment of 188 Cheat Basin Naturalist Vol. 49, No. 2 larval abundances. The ratings shown are, of emergence, and attraction to light, affect trap- necessity, somewhat subjective. The 10 spe- ping success and could be considered on a cies designated as abundant were either species-by-species basis. The objective of this caught in very high numbers at black lights study, however, was not to study reasons for and were present as larvae in large numbers in trapping success but to collect all species pos- Sagehen Creek (7 species), or were caught in sible, and no attempt was made to quantify high numbers in emergence traps and in trapping effort for each technique. many locations (3 species). In this latter Fish traps were considered emergence group, one species, bethula, inhab- Desmona traps for the foregoing comparison. They ac- its newly flooded areas and fluctuates greatly counted for collection of only four species that in numbers from year to year (Erman 1981). were not otherwise caught in emergence Thus, it may not always be abundant. Both it traps. They may catch , however, and Limnephilus peltus, discussed above, during up- or downstream flight rather than range in the Sierra have an apparently limited during emergence. Nevada but occur, at least periodically, in great abundance in the Sagehen Creek basin Adult Flight Periods and (see also Denning 1965). Community Emergence Patterns Thirty-six species are listed here as com- mon, and 31 are considered rare. In the rare Emergence periods are shown for each spe- group are 2 species that occur in rather high cies in Figure 1. Species are listed in order of numbers but were found in only one habitat or emergence (and, within that category, alpha- a few small ones within the basin. These are betically by family and species) beginning Goeracea oregona (in one spring only) and with January. The year was divided into 52 Parthina linea (a few spring areas). Parthina numbered weeks by eliminating 29 February linea exhibits an elaborate, nearly flightless and 31 December, as in the Rothamsted In- mating behavior on riparian vegetation (Er- sect Survey (Crichton 1971). To save space in man 1984), which may partly explain its lim- this paper (Figs. 1, 2), I have begun with week ited distribution. Females may be flightless. 14 (2-8 April) and ended with week 44 (29 Similar behavior has been observed in the October-4 November). Traps were run, how- limnephilid Psychoronia costalis (Scott Herr- ever, at some sites year-round, and the follow- mann, personal communication), also con- ing four species were collected in late au- fined to spring seeps and small streams. tumn, winter, and early spring: Psychoglypha klamathi, P. mazamae, P. ormiae, and Wor- Trapping Success and Differences maldia occidea. The four Sagehen Psycho- Fifty-seven species were caught in emer- glypha species (including P. bella) do not gence traps, and 52 were caught at black lights emerge in the summer period. Wormaldia

(Table 1). Thirty-three species were trapped occidea, on the other hand, was collected at by both methods. Thus, 24 species caught in constant-temperature springs during every emergence traps were never caught at black month of the year and apparently has a non- lights, and 19 species caught by black lights seasonal life cycle. Rhyacophila oreta began were not caught in emergence traps. One spe- emerging in early February and was collected cies, gilvipes, was caught only in every week through September. It had a by hand-netting and by larval rearing. These similar long, nonseasonal emergence period results give a trapping similarity of 43% (using in the Salmon River basin of (Smith Jaccard's Index, Pielou 1984) between the 1968). Four other species (Desmona bethula, methods and indicate the value of using differ- Rhyacophila ardala, R. vaccua, and R. ver- ent trapping methods to determine total spe- nda ) were occasionally trapped in November cies in a given geographical area. and December. Most species not caught at black lights were There are two major peaks in adult emer- from the springs and spring streams (Table 1) gence in the Sagehen Creek basin Trichoptera and/or emerged during a season other than community. One is the midsummer peak in summer (Fig. 1). Many factors, such as loca- late June and July. The other is the late tion of trap in relation to microhabitat of summer-fall peak that begins as the tempera- a species, diel flight activity, time of year of ture cools in August. At least 20 species in the April 1989 Erman: California Trichoptera 189

i 1 1 1 1 r Rhyacophila orela Hydroptila rono

Apatania sorex Cryptochia excella Psychoglypha ormiae * 1 1 1 1 »- Lepidostoma rayneri

Psychoglypha klamalhi Lepidostoma roafi Polycentropus Psychoglypha mazamae _J halidus

Wormaldia occidea < 1 1 1 1 1- Amiocentrus aspilus Psychoglypha bella Hydropsyche occidentalis almota Anagapetus n. sp. Lepidostoma ermanae Allomyia cidoipes Hydatophylax hesperus Yphria calilornica unicolor Rhyacophila brunnea Pedomoecus sierra

Limnephilus peltus Rhyacophila rotunda Rhyacophila valuma Micrasema bactro ) 1 1 1 1 I Heleroplectron calilormcum Agapetus taho Glossosoma alascense Limnephilus externus Goeracea oregona Limnephilus frijole grandis Wormaldia gabriella Parapsyche elsis Rhyacophila tucula Clostoeca disjuncta Rhyacophila verrula Ecclisomyia maculosa

Lenarchus nllus I 1 1 1 1 t Neophylax occidentis americanus Oligophlebodes sierra Arctopsyche calilornica Dolophilodes aequalis Desmona bethula Wormaldia pachita Dicosmoecus pallicornis Rhyacophila grandis Homophylax rentzi Rhyacophila nevadensis Homophylax sp.

-i 1 1 1 1 1 1- Hydropsyche amblis I I Ochrotrichia trapoiza Lepidosloma cascadense Rhyacophila ardala Lepidostoma unicolor

Hesperophylax designatus -t 1 1- Limnephilus mornsoni Neophylax rickeri Rhyacophila vaccua Neophylax splendens Rhyacophila vao ghseola Hydroptila n. sp.

-I 1 1 1 1 1 Allocosmoecus partitus Hydropsyche oslah H 1 1- Lepidosloma verodum Chyranda centralis Rhyacophila vuzana Cryptochia calilca/shasta (- Parthina linea I I Rhyacophila harmstoni Amphicosmoecus canax

15 20 25 30 35 40 15 20 25 30 35 40 Week Week

Fig. 1. Adult emergence periods ofTrichoptera from the Sagehen Creek basin by week, 2 April—4 November.

basin are in this latter category. Adult activ- cies. There is, however, an obvious associa- ity patterns for rarely caught species with no tion between the overall emergence pat- larval information must be considered incon- tern and these three physical factors. The clusive. The combined species emergence solid bars represent all species in the basin; only. (Fig. 2) for 1979-1986 shows a pronounced the open bars, Sagehen Creek species increase in emergence in week 24 (11-17 It should be remembered that the water temperature curve in Figure 2 does not ap- June). This pattern is plotted against air streams, temperature, water temperature in Sagehen ply to species living in temporary springs, etc. Creek (means from 1979 to 1986), and solar radiation (taken as the mean monthly radia- Geographical Comparisons tion published for a similar latitude—Reno, Nevada). No correlation is obvious to explain Presence or absence of Sagehen Creek the sudden rise in numbers of emerging spe- basin species in other geographic areas is 190 Great Basin Naturalist Vol. 49, No. 2

Table 1. Species composition, habitat, relative abundance, and capture method for Trichoptera in the Sagehen Creek basin, California. [SC - Sagehen ('reek. S spring origins, SS = spring streams, LSS = largest spring streams only (near mouth), TS = temporary streams, TP temporary ponds.]

Relative Capture method Reared or Species Habitat abundance Emergence trap Black light associated larvae

Brachycentridae aspilus (Ross) 1938 ? Brachycentrus americanus (Banks) 1899 Micrasema bactro Ross 1938 Calamoceratidae Heteroplectron californicum McLachlan 1871 Agapetus taho Ross 1947 Anagapetus sp. Glossosoma alascense Banks 1900 Goeridae Goeracea oregona Denning 1968 Arctopsyche californica Ling 1938 Arctopsyche grandis (Banks) 1900 Hydropsyche amblis Ross 1938 Hydropsyche occidentalis Banks 1900 Hydropsyche oslari Banks 1905 Parapsyche altnota Ross 1938 Para psyche elsis Milne 1936 Hydroptila rono Ross 1941 Hydroptila sp. Ochrotrichia trapoiza Ross 1947 Lepidostomatidae Lepidostoma cascadense (Milne) 1936 Lepidostoma rayneri Ross 1941 Lepidostoma roafi (Milne) 1936 Lepidostoma unicolor (Banks) 1911 Lepidostoma verodum Ross 1948 Lepidostoma ermanae Weaver 1988 Limnephilidae Allocosmoecus partitus Banks 1943 Allomyia cidoipes (Schmid) 1968 Amphicosmoecus canax (Ross) 1947 Apatania sorex Ross 1941 Chyranda centralis (Banks) 1900 Clostoeca disjuncta (Banks) 1914 Cryptochia califca/shasta Denning 1968,1975 Cryptochia excella Denning 1964 Desmona bethula Denning 1954 Dicosmoecus atripes (Hagen) 1875 Dicosmoecus gilvipes (Hagen) 1875 April 1989 Erman: California Trichoptera 191

Table 1 continued.

Relativi Capture method Reared or Species Habitat abundance Emergence trap Black light associated larvae

Dicosmoecus pallicornis Banks 1943 ? Ecclisomyia maculosa Banks 1920 Hesperophylax designatus Banks 1943 Homophylax rentzi Denning 1964 Homophylax sp. Hydatophylax hesperus (Banks) 1914 Lenarchus rillus (Milne) 1935 Limnephilus externus Hagen 1861 Limnephilus frijolc Ross 1944 Limnephilus morrisoni Banks 1920 Limnephilus pelt us Denning 1962 Neophylax oceidentis Banks 1924 Neophylax rickeri Milne 1935 Neophylax splendens Denning 1948 Oligophlebodes sierra Ross 1944 Banks 1897 Pedomoecus sierra Ross 1947 Psychoglypha bella (Banks) 1903 Psychoglypha klamathi Denning 1970 Psychoglypha mazamae Denning 1970 Psychoglypha ormiae (Ross) 1938 Odontoceridae Parthina linea Denning 1954 Dolophilodes aequalis (Banks) 1924 Wormaldia gabriella (Banks) 1930 Wormaldia occidea (Ross) 1938 Wormaldia pachita Denning 1956 Phryganeidae Yphria californica (Banks) 1970 Polycentropus halidus Milne 1936 Rhyaeophilidae Rhyacophila ardala Denning 1965 Rhyacophila brunnea Banks 1911 Rhyacophila grandis Banks 1911 Rhyacophila harmstoni Ross 1944 Rhyacophila nevadensis Banks 1924 Rhyacophila oreta Ross 1941 Rhyacophila rotunda Banks 1924 Rhyacophila tucula Ross 1950 Rhyacophila vaccua Milne 1936 Rhyacophila valuma Milne 1936 Rhyacophila vao Milne 1936 Rhyacophila verrula Milne 1936 Rhyacophila vuzana Milne 1936 Gumaga griseola (McLachlan) 1871 192 Great Basin Naturalist Vol. 49, No. 2

40 I All Species 16 _/N 35 Q Sagehen Ck. 14 •£"

30 12 !_ \o CO -- Light f? 25 \ 10 | if _i CO V\ ft 20 > 8 o • —• 0)

1 5 6 3 I TO z c5 N«A Q. : 10 ^° ,•—• Water vXx 4 E H

5 2

M i |B3 |MJ |»J |BJ |BJ |»j |M |M |M LM BJ |BJ |BJ ,! |M |, P-P^iJJLjjjJJLi p | | | k 15 20 25 30 35 40 (Week)

I I I Apr I May I Jun I Jul I Aug Sep Oct

Fig. 2. Weekly emergence of caddisflies from all habitats in the Sagehen Creek basin and from Sagehen Creek only. Air and water temperatures (Sagehen Creek) are weekly means; light is monthly mean solar radiation taken from the nearest recorded site of similar latitude (Reno, Nevada).

Fifty-nine Sagehen Creek basin species are prove otherwise. Except for those three, all also in Oregon; 36 of these in the western Sagehen Creek basin species found as far east Cascades of Oregon. Thirty-eight species are as Utah or Colorado are also found in Oregon. in common with Utah, 33 with Colorado, 18 Hesperophylax magnus was collected from with southeast Alaska, 15 with the Yukon, and Sagehen Creek on 8 October 1966 (Parker and only 8 with the Pacific Coast Range in Califor- Wiggins 1985), but it was not collected during in nia. This latter result may at first seem sur- the present study. Its earlier presence prising, except that the California Coast Sagehen Creek was the only California site extension Range has a Mediterranean climate (McEl- reported and was the westernmost ravy and Resh 1987) very different from the of its distribution. short growing season and long, cold winters of Taxonomic Considerations the eastern Sierra Nevada. It is apparent that Some of the Sagehen Creek basin species certain species are ubiquitous in western show slight variations from described forms; a North America (i.e., Hydropsyche oslari, few are apparently undescribed species for Polycentropus halidus, Rhyacophila vao), which descriptions are planned. and others are probably confined to a small Anagapetus sp. is an undescribed species area in the Sierra Nevada. Fifteen Sagehen similar to A. chandleri and A. aisha, but it Creek basin species are found on none of the differs from both in a consistent way. above regional lists. Of these locally dis- Hydropsyche amblis is the form designated species, it is interesting to note that 4 tributed H. ahella by Denning (1952). I am considering the 10 abundant species in are among most it H. amblis here, based on information (Pat the Sagehen Creek basin (Anagapetus n. sp., Schefter, personal communication) that the Desmona bethula, Limnephilus peltus, and species varies widely. My designation is ten- Rhyacophila ardala). tative until further work is done. Three species (Ochrotrichia trapoiza, Rliy- Lepidostoma crmanae, recently described acophila harmstoni, and R. rotunda) have a by Weaver (1988), is similar to L. hoodi and wide distribution to the east (Utah and/or Col- L. spicatum. orado) but not northward. Of course, more Clostoeca disjuncta differs somewhat in extensive collections in eastern Oregon may both larva and adult from those described. April 1989 Erman: California Trichoptera 193

The larva has a sclerite on the lateral hump of Wormaldia, occur in completely different abdominal segment I, unlike that described aquatic habitats. Wormaldia pachita was

by Wiggins (1977), and makes a case of sedge found in one temporary stream where it pieces. The adult has a prominently marked emerged just prior to the complete drying of wing. The species shows wide variation over the stream. Wormaldia occidea, on the other its range (Glenn Wiggins, personal communi- hand, occurs in constant-temperature springs cation). and emerges year-round. And W. gabriella The few specimens of Cryptochia califcal lives in Sagehen Creek and large spring shasta from the Sagehen Creek basin have streams only. characteristics of both species. They were all Other related species were found in the caught in one spring stream over a span of a same habitat but were ecologically separated few weeks. Further work is underway to col- by the timing of their life cycles. Agapetus lect more specimens by emergence trapping taho, Anagapetus sp., and Glossosoma alas- to help resolve the taxonomic questions. Both cense, with similar larvae, all occur in Sage- of these species were originally named from hen Creek but are separated by emergence single males (Denning 1968, 1975). periods. Anagapetus sp. emerges primarily in I collected Ecclisomyia larvae, but no early summer, G. alascense in midsummer, adults, in very cold springs, 3-4 C, at the head and A. taho in late summer and fall. In addi- of the basin (elevation 2,408 m). These may be tion, A. sp. is adapted to large and small a different species from the Ecclisomyia mac- spring streams as well as Sagehen Creek; ulosa collected from the larger, warmer G. alascense is restricted to Sagehen Creek streams at a lower elevation (1,943 m). (Table 1). Other species of these three genera Neophylax rickeri and Neophylax splen- often occur in the same streams (Anderson dens warrant further taxonomic and ecological and Wold 1972). work in other sites where they occur together It is interesting that so many Dicosmoeci- and in greater abundance than in the Sagehen nae are found in the Sagehen Creek basin, Creek basin (only 11 males and 6 females total including the three possible species of Dicos- were caught during this study). The males moccus and the closely related Allocosmoe- were difficult to separate, and separating the cus. None of these four species was caught in females was little more than guesswork. The emergence traps, but larval rearings showed two species have the same emergence period that, as predicted by Wiggins and Bichardson in the Sagehen Creek basin. Anderson and (1982), D. gilvipes and D. atripes tended not Wold (1972) reported a similar finding in Ore- to be at the same site. Dicosmoecus gilvipes gon where the two species appeared in the larvae live farther downstream in more open, same emergence traps during the same inter- warm areas of Sagehen Creek, and adults val in October. Neophylax splendens may be a emerge later than D. atripes (Fig. 1). Larvae synonym of N. rickeri. of A. partitas were infrequently found in the A similar situation is true for Rhyacophila same section of Sagehen Creek as D. atripes, brunnea and R. vao. These species are al- but the preferred larval habitats of A. partitas ready known to vary widely (Smith and Man- and D. pallicornis are unknown in the Sage- ual 1984). Three distinct forms were found hen Creek basin. during this study, two variants of R. brunnea The four species ofPsychoglypha have simi- and one of R. vao. A few specimens seemed lar emergence periods, but P. bella is re- intermediate between the two species. Both stricted to Sagehen Creek and P. mazamae species were collected in the same emergence (larvae and adults) is found in warmer down- traps at the same time and in the same sweep stream sections of small spring streams than nets. I have separated the males but think are the adults of P. ormiae and P. klamathi. I they very likely may be one highly variable have not successfully reared or associated species. with adults the larvae of these latter two and do not know if the larvae can be distinguished Ecological Separation of from each other. Closely Belated Species Some The habitat differences of Chyranda cen- Some closely related species in the Sage- tralis and Clostoeca disjuncta are worth not- hen Creek basin, like the three species of ing because of their similar larvae and cases. 194 Great Basin Naturalist Vol. 49, No. 2

Table 2. Presence ofTrichoptera species of the Sagehen Creek basin in other western North American areas.

Western Coast 2 Sagehen Creek basin Cascades Southeast Range' 3 4 5 species Oregon (Oregon) Utah Colorado Alaska Yukon'' (Californi

Brachycentridae April 1989 Erman: California Trichoptera 195

Table 2 continued.

Western Coast Sagehen Creek Basin Cascades" Southeast Range' 3 4 1 6 species Oregon (Oregon) Utah Colorado Alaska Yukon (California)

Humophijlax rentzi Homophylax sp. Hydatophylax hesperus X Lenarchus rillus Limnephilus externus Lim neph ilus frijole Limnephilus morrisoni Limnephilus peltus Neophylax occidentis Neophylax rickeri Neophylax splendens Oligophlebodes sierra Onocosmoecus unicolor Pedomoecus sierra Psychoglypha hella Psychoglypha klamathi Psychoglypha mazamae Psychoglypha ormiae Odontoceridae Parthina linea Philopotamidae Dolophilodes aequalis Wormaldia gabriella Wormaldia occidea Wormaldia pachita Phryganeidae Yphria calij"arnica Polycentropodidae Polycentropus halidus Rhyacophilidae Rhyacophila ardala Rhyacophila brunnea Rhyacophila grandis Rhyacophila harmstoni Rh yacoph ila nevadensis Rhyacophila oreta Rhyacophila rotunda Rhyacophila tucula Rhyacophila vaccua Rhyacophila valuma Rhyacophila vao Rhyacophila verrula Rhyacophila vuzana Sericostomatidae Gumaga griseola

Total species in common 196 Great Basin Naturalist Vol. 49, No. 2

Key to larvae of Nearetic species groups of Literature Cited Rhyacophila, available from author). Rlu/a- Anderson. N H 1976. The distribution and biology of cophila ardala and R. vaccua are the only the Oregon Trichoptera. Oregon State Univ. other pair of closely related Rhyacophila. Agric. Expt. Sta. Tech. Bull. 134. 152 pp. in Smith's key They keyed to the same place Anderson, N H , G M Cooper, and D G Denning. of Experi- but could be separated by head length vs. 1982. Invertebrates the H. J. Andrews western Cascades, II. An head width and by correlation with adults mental Forest, Oregon. annotated checklist of caddisflies (Trichoptera). from emergence traps. It was then apparent Oregon Agric. Expt. Sta. Tech. Publ. 5772. that their habitats are different. Rhycophila trap Anderson, N H., and J. L. Wold 1972. Emergence ardala occurred in cold spring sources, and collections of Trichoptera from an Oregon stream. R. vaccua was primarily in Sagehen Creek Canadian Entomol. 104: 189-201.

Bartolome. , D C. Erman, and C F. Schwarz. and near mouths of spring streams close to J W 1988. Stability and change in minerotrophic peat- Sagehen Creek. lands. Sierra Nevada of California and Nevada. Pacific Southwest Forest and Range Expt. Sta., Conclusions Forest Service, USDA, Berkeley, California. (In press.) BAUMANN, R \V and D Unzicker 1981. Preliminary Though the eastern Sierra Nevada is dry J checklist of Utah caddisflies (Trichoptera). Ency- and has a brief growing season, a wide diver- clia 58: 25-29. sity of small aquatic habitats, both permanent Brock. E W 1960. Mutualism between the midge Crico- and intermittent, seems to account for a large topus and the alga Nostoc. Ecology 41: 474-483. and diverse Trichoptera community with spe- Crichton. M. I. 1971. A study of caddisflies (Trichoptera) of the family Limnephilidae, based on the Roth- cies adapted to nearly every aquatic possibil- amsted Survey, 1964-68. J. Zool., London ity. detailed studies in other Sierra Similar 163: 533-563. Nevada basins on both east and west sides and Denning, DG 1952. Descriptions of several new species in eastern Oregon and Nevada are needed to of caddis flies. Canadian Entomol. 84: 17-22. increase our understanding of California Tri- 1965. New Rhyacophilids and Limnephilids (Tri- choptera: Rhyacophilidae and Limnephilidae). choptera communities. It is unfortunate that, Canadian Entomol. 97: 690-700. although Trichoptera systematists have col- 1968. New and interesting North American Tri- lected in California for a long time, so little choptera. Pan-Pac. Entomol. 44: 17-26. detailed information about commu- 1970. The genus Psychoglypha (Trichoptera: Lim- Entomol. 102: 15-30. nities is known for this state. With ever- nephilidae). Canadian 1975. New species of Trichoptera from western increasing demands being made on California North America. Pan-Pac. Entomol. 51:318-26. waters, and recently on the smaller streams Erman, D C. 1976. Peat depth of Sierra Nevada fens, and and springs of the Sierra Nevada, a great profile changes from 1958 to 1972 in Mason Fen. wealth of evolutionary, ecological, and bio- Great Basin Nat. 36: 101-107. geographical information concerning Tri- Erman, D C, and N. A. Erman. 1975. Macroinverte- brate composition and production in some Sierra choptera may be lost before it is ever docu- Nevada minerotrophic peatlands. Ecology 56: mented. 591-603.

Erman. D C , and G. R. Leidy. 1975. Downstream move- Acknowledgments ment of rainbow trout fry in a tributary of Sagehen Creek, under permanent and intermittent flow. Trans. Amer. Fish. Soc. 104: 467-473. I thank the late Don Denning, Glenn Erman, N A. 1981. Terrestrial feeding migration and life Wiggins, Pat Schefter, Stamford Smith, and history of the stream-dwelling caddisfly, Desmona John Weaver for their help with verification of bethula (Trichoptera: Limnephilidae). Canadian 59: 1658-1665. species and resolution of taxonomic ques- J. Zool. 1984. The mating behavior of Parthina linea (Tri- tions. I am grateful to the following people for choptera: Odontoceridae), a caddisfly of springs emptying traps at different times during the and seeps. Pages 131-136 in J. C. Morse, ed., study: Chris Kellner, Vernon Hawthorne, Proc. 4th Int. Symp. Trichoptera. Dr. W. Junk Mike Yoder-Williams, Joe Thornton, Wayne Pub., The Hague. Spencer, and Lynn Decker. And my thanks to 1986. Movements of self-marked caddisfly larvae, Chyranda centralis (Trichoptera: Limnephilidae), Don Erman for editorial and field assistance. in a Sierran spring stream, California, U.S.A. Partial provided by the Univer- funding was Freshwater Biol. 16: 455-464. sity of California Water Resources Center, 1987. Caddisfly adaptations to the variable habi- Project UCAL-WRC-W-645. tats at the land-water interface. Pages 275-279 April 1989 Erman: California Trichoptera 197

in M. Bournaud and H. Tachet, eds., Proc. 5th 1972. Comparative ecology of Arcynopteryx and Int. Symp. Trichoptera. Dr. W. Junk Pub., The Diura (Plecoptera) in a California stream. Arch. Hague. Hydrobiol. 69: 521-546.

Herrmann, S. J , D E Ruiter, and J. D. Unzicker. 1986. 1980. Coexistence of perlid stoneflies (Ple- Distribution and records of Colorado Trichoptera. coptera): predictions from multivariate morpho- Southwestern Nat. 31: 421-457. metries. Hydrobiologia 71: 99-105.

Holsincer. J. R 1974. Systematics of the subterranean Sheldon, A L . and S G Jewett, Jr 1967. Stonefly amphipod genus Stygobromus (Gammaridae), emergence in a Sierra Nevada stream. Pan-Pac.

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