Micaomsmaunon or - 1, ‘ -. PYCNOPSYCHE‘LEPtDMflAGEN) , .. AND .PY-CNOPSYCHE comma (WALKER); _ 7' , e:_:2_:}1.:-_:1;;i (TREHOPTERA; LIMNEPHlLIDAE) _- g _ g, IN VERMILLI‘ON CREEK. MICHIGAN; """" ........ ,,,,,,, "(basis for the Degree 01M. 8. - MICHIGAN STATE UNIVERSHY ROGER WILUAM MUSTAUSH 1973 ‘ ABSTRACT MICRODISTRIBUTION OF PYCNOPSYCHE LEPIDA (HAGEN) AND PYCNOPSYCHE GUTTIFER (WALKER), (TRICHOPTERA: LIMNEPHILIDAE) IN VERMILLION CREEK, MICHIGAN By Roger William Mustalish The microdistribution of two species of Pycnopsyche larvae (Trichoptera: Limnephilidae), was investigated inathat portion of Vermillion Creek which flowed through the Orange sector of the Rose Lake Wildlife Aesearch Area, Calhoun Co., Michigan. The study consisted of: (a) an eight-week monitoring program, during which pH, turbidity, total carbon, dissolved oxygen, tem- perature, depth and current at three sites were closely observed; (b) a general survey of the stream to establish the degree of dispersal of the two species; (c) a quantitative stratified random sampling of the three sites for the larvae. Head capsule widths at the eyes and dry weights were determined for all organisms collected. Graphic analyses of the monitoring data revealed no major influence by any parameters on the microdistribution of the species, with the possible exception of dissolved oxygen.’ Roger William Mustalish Similarly, no conditions were found during the general survey that would indicate that conditions existed along Vermillion Creek to prohibit the existence of either species. Both were con- sistantly observed throughout the survey. Quantitative frequency distribution information, coupled with chi-square analysis, revealed no intraspecific influences on individual species microdistribution in a given habitat, while the application of a coefficient of numerical separation demonstrated the significance of substrate type in species segregation. The general conclusion of the study was that the micro- distributional pattern of E, guttifer and E, lgpigg, in Vermillion Creek, was a consequence of: (a) a temporal lag in their respec- tive life cycles; (b) the instinctual need for optimal pupation sites; (c) the active migration by E, lgpjg2_as a response to this innate stimulus. MICRODISTRIBUTION OF PYCNOPSYCHE LEPIDA (HAGEN) AND PYCNOPSYCHE GUTTIFER (WALKER), (TRICHOPTERA: LIMNEPHILIDAE) IN VERMILLION CREEK, MICHIGAN By Roger William Mustalish A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Zoology 1973 ACKNOWLEDGMENTS I wish to thank the following for their contributions: Dr. T. w. Porter for his guidance and advice throughout my graduate studies at Michigan State University. Dr. Clarence C. McKnabb for his advice and aid in pro- curing laboratory space and equipment. Dr. Kenneth w. Cummins for his critical appraisal of the biological aspect of the study and his advice of sampling procedures. Dr. Charles S. Thornton and the Zoology Department for financial assistance. Dr. Niles R. Kevern and the Fisheries and Wildlife Department for the use of facilities at the Limnological Research Laboratory. ii TABLE OF CONTENTS Page LIST OF TABLES ..................... . . . v LIST OF FIGURES . . . . . ..... . ..... . ...... vi INTRODUCTION . . . . ........... . ..... . . . . l STUDY AREA . . . . . . . . . . . ..... . ..... . . . . 4 METHODS . ..... . . . . . . . . . . . . . . . . . . . . . ll Physical and Chemical Monitoring .......... . . . ll Dissolved Oxygen . . ..... . ..... . . . . . . . ll Turbidity ......... . ..... . ..... . . . ll Total carbon . . . . . . ........... . . . . . l2 pH ......... . ................. l3 Depth .......................... l3 Current .................. . ...... l3 Temperature . . . . . ........ . . . . ..... . l4 General Stream Survey ..... . . . . ..... . . . . . l4 Quantitative Site Sampling ..... . ..... . . . . . l4 RESULTS ........................... l6 Physical and Chemical Monitoring ............ . l6 General Stream Survey .............. . . . . . l6 Quantitative Site Sampling ........... . . . . . 2l DISCUSSION .......................... 24 Physical and Chemical Monitoring ............. 24 Depth ..... . . . . ..... . . . . . . . ..... 25 Current . . . . ..... . ...... . . . . . . . . . 27 pH ....... . ................. . . 28 Turbidity .............. . ........ . 29 Total Carbon . ........... . ..... . . . . 3l Temperature . . ..... . ........... . . . . 34 Dissolved Oxygen . . . . ........ . . . . . . . . 34 Other factors . . . . . . . . . . . . ..... . . . . . 38 Quantitative Site Sampling . . . . . . . . . . . . . . . . 40 Frequency Distribution of Head Capsule Widths . . . . . . 4O Frequency Distribution of Dry Weights . . ..... . . . 4O Coefficient of Numerical Separation . . . . ....... 47 CONCLUSION ............. . ............. 5l SUMMARY ..... . . . . . . . . . . .‘ ....... . . . . 53 BIBLIOGRAPHY .................. . ...... 54 Literature Cited ............... . ..... 54 General Reference . . . . ........... . ..... 57 APPENDICES ............. . ............ 59 Appendix A . . . . ..... . ..... . . . . . . . . . 59 Appendix B .................. . ..... 67 iv LIST OF TABLES Table Page l. Weekly averages of stream monitoring . . . . . . . . . . . l7 2. Weather data ..... . . . . ..... . . . . . . . . . 20 3. Summary of biological collections . . . . . . . . . . . . 23 4. Intrasite variation in 0.0. (ppm) . . . . . . . . . . . . 39 5. Coefficients of numerical separation . . . . . . . . . . . 49 6. Results of all determinations during monitoring . . . . . 59 7. Results of biological collections . . . . . . . . . . . . 67 LIST OF FIGURES Figure Page l. Rose Lake wildlife area . . . . ..... . . . . . . . . 5 2. Site l ................. . . ..... 7 3. Site 2 ...................... . 8 4. Site 3 .................. . . 9 5. Stream profiles . . . . . . ........ . . . . . . . lO 6. Depth; current . . . . . . . . . . ..... . . . . . . . 25 7. Turbidity (ppm) . . . .................. 30 8. Total carbon (ppm) . . . . ..... . . . . . . . . . . . 32 9. D.O.; temp. . . . . . . ........... . . . . . . 35 10A. Frequency distribution of head capsule widths-— E, guttifer ...................... 4l 108. Frequency distribution of head capsule widths-- E, lgpjga_ ....................... 42 llA. Frequency distribution of dry weights--E, guttifer . . . . 43 llB. Frequency distribution of dry weights--E, lepida . . . . . 45 vi INTRODUCTION Species of the genus Pycnopsyche belong to the trichopteran family Limnephilidae and are found throughout northeastern North America. They are holometabolous organisms, the larvae of which are aquatic in lotic environments. .Despite their large size (up to 25 mm. long), distinctive cases and relative frequency, few investigators have chosen them as subjects for detailed ecological studies. : Early works were taxonomic in nature and characterized by misleading descriptions and general disorganization (Ross, l944; Lloyd, l921; Betten, l934). Betten (1950) reorganized the genus by uniting a number of species which had previously been placed in other genera. The revised genus now consisted of fourteen species, but larval determination was still difficult due, in large part, to the lack of distinctive morphological characteristics. Reliable identification of seven terminal instars of Eycnopsyche became possible, however, following the study by Flint (l960). Although many of these early works contained notes on the biology of the genus, no detailed investigations concerning the ecology of the larvae were conducted until Cummins (l964), Feldmeth (1970) and Mackay (unpublished Ph.D. thesis). It is quite common to find two or three Pycnopsyche species in the same habitat, and in the case of E, lepida and E, guttifer, this is nearly always the rule (Flint, l960; Cummins personal communication). Upon hatching in early September, the larvae of E, lgpjga_ begin to construct a case composed of vegetable matter and spend their early instars feeding on allochthonous organic material within the marginal, slow-water areas of the stream. As terminal instar approaches, the larvae leave the margins of the stream for the swifter flowing central portions characterized by a substrate of gravel. Their food now becomes mostly algae. They remain active on the substrate surface until February when they cease feeding, burrow into the substrate and enter diapause. Diapause lasts until late summer after which they come to the surface of the gravel, pupate and finally emerge in late July, August and early September. It is during this time of emergence that they are most susceptible to predation, especially by fish (Cummins, l964; Muttkowshi and Smith, l929; Needham, l927; Betten 1934; Mackay, 1969). The life cycle of E, guttifer is somewhat simpler than that of E, 122193, The new 3, guttifer population becomes estab- lished in early December. They, too, remain in the marginal areas of the stream and feed on allochthonous organic material, but their cases are composed of equal amounts of sticks and leaf material. As they mature, their cases become exclusively stick material, though some mineral material may be used at times. They do not, however, venture into the central portions of the stream at terminal instar. Instead they remain in the slower areas of the stream and in late spring attach themselves to sticks, cobbles and submerged branches of shore plants. Feeding ceases and they remain in diapause until late September and October when pupation and emergence occur (Cummins, l964; Mackay, l969; Betten, l934; Muttkowski and Smith, l929). One reason they are able to coexist in the same general habitat is because they are separated temporarily due to their staggered life cycles, but all of the factors which influence the microdistribution of these species in a given system have only begun to be understood. The purpose of the present study was to monitor the chemical and physical parameters of a nearby stream, and follow- ing a quantitative sampling for E, igpiga_and E, guttifer larvae, attempt to illustrate some of the factors which exert influence on the microdistribution of these insects. STUDY AREA Vermillion Creek is a warm, slow flowing stream located twelve miles northeast of Lansing, Michigan, and eventually flows into the Looking Glass River. The water table in the area is quite high and the major source of the stream's flow, at times emptying into it as open springs. In periods of dry weather, 80-90% of the flow is due to this groundwater seepage, according to John Byerlay of the Water and Environment Section of the United States Geologi- cal Survey Division (personal communication). Additional flow is in the form of terrestial runoff and effluent from Rose Lake, septic tank overflow, small intermittent streams and a 60#7O acre man-made flooded area. A three and one-half mile section of the stream, which flows through the Orange sector of the Rose Lake wildlife Area, was studied (see Figure 1). Three monitoring sites were chosen along the length of the stream. The first was at the junction of Woodbury Road. The stream originally flowed several meters to the north, but highway construction near the stream resulted in rerouting to its present position. Site 2 was located at Peacock Road. A concrete and steel-pipe walkover bridge is located here and the stream bed is characterized by boulders and cobbles, apparently as a result of 3.x 3.1 . ...: «A. {ll 9“. . o . ....mnWJ 9.5.1.... .4... -. mace “keen: . .. II o wH 1 mad. IUm A :35 22:53:. II ..I. ...m 92:... .3..— -Iall o!< octton E r 3113 0.11! 25: :35 ill; 32.5 552...: _ .4 . a . ... . J . . .. .. .80: coon—Sm 20: I 8:25; 8—0.. 25: b M n . . . . < 3“! "an...“ a O . ”<4. ... . .. S W .3 . q .... u _ a 3 .... a. .... . $0 IS- 6 4. . 3%»?qu .. ..p .. a. 3, .. v... ... .. . 2.5.3.4 flaw“. t «UN... .4 l o. . . oaoz oaaumlu ~... \ . ..‘b'l—JWKI lllllhh< :4; at ‘Q ...o o-~‘ ‘. no. . Cir.” .......1..on......4....... , a a... a . QIJLWJOWO'i "Oxlmflfdvlfld‘xr. Don‘t-(,0)-L.. 3d. . . .o s no . .f . . .ru..u4:.w n. a. , 3..., a 036:5.(4 u_z ...!» the bridge construction (no other section of the stream has such substrate characteristics, sand and silt being the standard). Overflow from Rose Lake enters one-quarter mile upstream from this site. The last site was at the junction of Bath Road. This section is subjected to effluent from the flooded area southeast of the site. Figures 2, 3 and 4 represent the three sites, respectively, and illustrate the substrate characteristics as well as the surrounding area. Cummins (1964) has suggested the use of the "phi-scale" as a classification for substrate sizes. Phi is the negative log to the base two of the particle size diameter in millimeters. Using such a scale, the boulders of Site 2 are -8 phi, the cobbles, -6, -7 and the pebbles, -5, -4. The gravel at all of the sites ranged from -3 to —1, while the sand was very coarse to medium and, therefore, 0 to 2. Finally, the silt ranged from 5 to 8, Using a method outlined by Leopold, §t_gl, (1964), a line was secured across each site and depth measurements taken every six inches. The result was a profile of the stream bed at each site. These are reproduced in Figure 5. gravel use vomoaxm x X X Hzmzamm \\\ ll \‘\ I” \\\\\ I’ll, \\\|\ D .l e V a N r g A S 1/ IIIIIIIIIIIIIIIIIIIIIIII \\ HzmzHomm \A D X X X X X X X X X x O X X X X X Figure 2. tree Site 1. lowland brush X ’ suaplnog , I / \ ," “~‘-___—-_______—‘,”’ [BAPJQ "’-.__.-—c——'-—---- -----—--~-~-fl-" A (I) Q, 3 a: ' ...- A S. U) .Q LIJ 0) v l—' .D D .Q 0 D. _.I I W a) D I— .Q .9 O O U V CO ----§----- ~-- -“‘ ‘s. ‘\\ ‘ “~ ”- “~‘\ '2 \ \ LLI z \ H ---- D ----—------- -% “-—~- 3; I" squeld paBJaqus Cf» X X x x \ X X x O x X X X X X X X X 0 = tree lowland brush X Site 2. Figure 3. X x x X X x X x x x X X X X . x x X x X x X x . . X X X X \\ . X x x x . X X \\ . SE: Swfizmzm x . Emzamm " x x x \ x X \ . x x .\\ ‘I’lu' I x x \\_\\\\ -l" \‘I I I III“ """"""\‘ I, I""“ “\ ” |“““ ”" --'|-“‘|““ D 9 AU 1 .. e N e V V ., .d 3...! r A r g g S ‘\"'|ll \\\\ Iii-Ill, \\\ ”"I-"||‘l|l|"‘-' \ ..\\\ Emzsmm \ ‘I‘ \ IIII'|-II|"l-'"I.IIIII|I'\ \IIel-I'" \\s\x \\ I, \\ \ I \\ x \ ‘ x I \ x -DDDII“WA x (y AIINA‘ x x l ‘IIII x x x s x x X x ..I||\\..\11 X X X X 0 X X X X X X x x X x x x x x x x x x X x x x x X x X x x X x X x X tree lowland brush 0 X Site 3. Figure 4. IO .mmFPmoLa Emmgpm .m we:m_m m wuwm N muwm I ll‘! F mpwm xeam “seem xeem pee. METHODS Physical and Chemical Monitoring For an eight-week period (April 15-June 10, 1973) seven ffi physical and chemical parameters were monitored at each of the three sites. Dissolved Oxygen Dissolved oxygen was determined using the azide modifi- cation of the standard Winkler test. The modification was employed as a safeguard against possible interference due to high degrees of organic nutrients in the stream water. To account for diurnal fluctuations in the dissolved oxygen levels, determinations were made at sunrise, when the levels would tend to be at their lowest, and in the late afternoon, when the levels would tend to be at their highest. Determinations were made three times per week. Turbidity One hundred mls. of water were collected at each site three times per week and returned to the laboratory for turbidity determinations. Each sample was shaken and 50 mls. drawn off for analysis in a Model 900-3 Klett-Summerson colorimeter equipped with a number 42 blue filter. Another 50 mls. were filtered first 11 12 and then placed in the colorimeter. The difference between the two readings was then plotted against a standard turbidity curve and the value in parts per million (ppm) obtained. Total Carbon One hundred mls. of water were collected at each site three times per week and analyzed for total carbon by the phenol- sulfuric acid method (Dubois, et_al,, 1956). The method was originally designed to be performed on purified samples obtained through partition chromatography using a phenol-water solvent, but can be modified (Mustalish unpublished) to yield reliable estimates of the total carbohydrate content of crude water samples. The modification is based on the results of studies by Larson (personal communication) of the Stroud Water Research Laboratory in Avondale, Pennsylvania. He found that approximately 85% of the total carbon determined in water samples in the White and Red Clay Creek systems can be attributed to carbohydrates. There- fore, by multiplying the results of the phenol-sulfuric acid method by a correction factor, a fairly reliable estimate of total carbon can be obtained. The test itself was quite simple. To 10 mls. of sample were added 5 mls of 5% phenol and 25 mls. concentrated sulfuric acid. The mixture was allowed to stand for ten minutes while a deep yellow color developed. It was then placed in a 23°C. water bath for ten to twenty minutes, after which it was placed in the same colorimetric apparatus as for turbidity. The values were l3 plotted against a standard glucose curve and the results corrected to yield total carbon in parts per million (ppm). L” This was determined at each site through the use of a series of p-Hydrion papers and compared to a standard color chart (correct to :_.025). Three determinations were made per week. Depth Depth markers were placed in the center of the stream at each site and readings were taken three times per week. Current Surface velocity was determined each week through the use of a simple float timed over a predetermined distance. Cummins (1964) seriously questioned the value of such determinations as they reveal little of the conditions at the stream bed. This draw- back was realized from the outset, but equipment designed for stream-bed velocity determinations was unobtainable. Therefore, it was hoped that although no significant conclusions could be drawn from the velocity determinations, at least the nature of the surface current at the three sites throughout the monitoring period could be established. 14 Temperature A Taylor minimum-maximum thermometer was placed at Site 2 with readings being taken each week. Spot checks at the other two sites throughout the monitoring period agreed well with the tempera- ture ranges being found at Site 2. General Stream Survey To determine the extent of dispersion of the two species throughout the entire length of the stream, hand net samples were taken at intervals of twenty meters, from Woodbury Road to Bath Road. No attempt was made to quantitatively sample the entire stream. Only presence or absence of each species was recorded. Quantitative Site Sampling, Larvae of both species were collected at each site using a stratified random sampling scheme. The stratification was on the basis of substrate type as studies by Cummins (1962, 1964), Mackay and Kalff (1969), Mackay (unpublished Ph.D. thesis) and others have demonstrated the significance of substrate types to insect distribution. The sampling was done in June; both species were in diapause and therefore, either burrowed into or attached to substrate. Within each substrate type, ten 1 square foot (.092 m2) samples were randomly taken using a coarse mesh hand net (Mecan, 1949-1950). The procedure was as follows. For each sample a one square foot frame was placed on the substrate and all exposed 15 larvae were collected. A stiff box-shaped wire mesh net, mounted on a wooden rod, was then pUshed several inches into the stream bed and a one square foot area of substrate removed. This techni- que enabled burrowed larvae to be collected. When sampling submerged branches of shore plants, the branches were lifted out of the water, the frame was placed over a section and the larvae attached to the branches within the frame were removed. As all the sites were not the same, the number of sub- strate divisions and therefore the total number of samples differed from site to site. At Site 1, twenty samples were taken (ten in sand, ten in sediment), at Site 2, forty (ten each in sand, sedi- ment, gravel and submerged plants) and at Site 3, thirty (ten each in sand, sediment and submerged plants). Head capsule widths at the eyes were determined for all larvae using an ocular micrometer and dry weights calculated after placing the organisms in an oven for twenty-four hours at 100°C. Apparently this sampling technique would best account for the environmental irregularities found at the sites and do so on a scale which Cummins (1962) has stressed must correspond to the size and habits of the particular animals being investigated. RESULTS Physical and Chemical Monitoring The average values per week of the physical and chemical .. parameters monitored at the three sites are summarized in Table 1. Appendix A provides a complete listing of determinations for each 2‘ sampling day throughout the monitoring period. For the sake of completeness, Table 2 lists the weather data for the period as compiled by the National Weather Service (precipitation data were available only to May 30). General Stream Survey Vermillion Creek from Woodbury Road to Bath Road was remarkably homogeneous in nature. With the exception of Site 2, the stream bed was predominantly sand, though interspersed with gravel in the central regions and silt along the margins. No true riffle areas were found. There were very few straight sections as meandering was the rule. The stream's drainage area was generally lowland woods and brush though marshy areas were sometimes encountered. Woody plants grew quite close to the bank and at times, bushes were growing within the stream. Similarly, fallen trees across the channel and woody debris were quite com- mon. The two characteristics that varied the most were stream 16 17 Table 1. Weekly averages of stream monitoring. Site 1 Site 2 Site 3 Week l 0.0. (ppm) A.M./P.M. 8.2/12.l .3/ll.7 7.5/10.9 Turbidity (ppm) 9.512 17.118 19.672 Total carbon (ppm) 7.073 8.719 7.398 PH 6.0 6.0 6.0 Depth (m) .940 .836 1.176 Current (m/sec) .423 .726 .346 Temp. range (°C.) 8.0-18.3 .0-18.3 8.0-18.3 Week 2 0.0. (ppm) A.M./P.M. 9.2/1l.7 .2/ll.2 8.l/ll.3 Turbidity (ppm) 18.327 11.654 13.659 Total carbon (ppm) 12.090 15.778 12.319 PH 6.0 6.0 6.0 Depth (m) .978 .911 1.027 Current (m/sec) .215 .610 .435 Temp. range (°C.) 6.7-16.7 .7-16.7 6.7-16.7 Week 3 D.0. (ppm) A.M./P.M. 8.6/11.7 .2/10.9 8.5/10.3 Turbidity (ppm) 18.052 10.309 24.065 Total carbon (ppm) 13.514 12.384 11.653 pH 6.0 6.0 6.0 Depth .932 .817 1.319 Current (m/sec) .254 .663 .314 Temp. range (°C.) 5.6-17.8 .6-17.8 5.6-17.8 18 Table 1. (Continued) Site 1 Site 2 Site 3 Week 4 0.0. (ppm) A.M./P.M. 8,0/11.2 7.6/11.2 7.9/10.6 Turbidity (ppm) 21.676 11.819 22.967 Total carbon (ppm) 16.957 15.473 15.903 pH 6.0 6.0 6.0 Depth (ppm) .826 .661 1.029 Current (m/sec) .242 .508 .331 Temp. range (°C.) 9.4-18.3 9.4-18.3 9.4-18.3 Week 5 0.0. (ppm) A.M./P.M. 8.3/10.2 8.4/10.3 8.4/9.6 Turbidity (ppm) 38.920 34.170 51.907 Total carbon (ppm) 15.899 14.982 15.304 pH 6.0 6.0 6.0 Depth (m) .665 .491 .902 Current (m/sec) .148 .435 .339 Temp. range (°C.) 6.7-16.7 6.7-16.7 6.7-16.7 Week 6 D.O. (ppm) 7.0/8.4 6.7/8.0 7.0/7.9 Turbidity (ppm) 46.745 56.575 58.414 Total carbon (ppm) 16.189 15.821 15.982 pH 6.0 6.0 6.0 Depth (m) .762 .614 .965 Current (m/sec) .381 1.220 .423 Temp. range (°C.) 11.7-18.9 11.7-18.9 11.7-18.9 Table 1. (Continued) 19 Site 1 Site 2 Site 3 Week 7 D.O. (ppm) A.M./P.M. 7.0/8.2 6.4/8.2 6.5/7.9 Turbidity (ppm) 13.412 18.024 31.533 Total carbon (ppm) 15.416 15.660 21.753 pH 6.0 6.0 6.0 Depth (m) 1.050 1.046 1.333 Current (m/sec) .288 .677 .359 Temp. range (°C.) 10.0-21.1 l0.0-21.1 l0.0-21.l Week 8 D.0. (ppm) A.M./P.M. 5.4/8.1 5.1/8.3 5.4/7.7 Turbidity (ppm) 40.649 14.784 9.238 Total carbon (ppm) 13.328 11.461 10.565 pH 6.0 6.0 6.0 Depth (m) .859 .760 1.058 Current (m/sec) .321 .762 .508 Temp. Range (°C.) 16.1-23.3 16.1-23.3 16.1-23.3 width and water depth. Width ranged from a little over a meter to nearly ten meters, while depth ranged from less than one-half meter to over one and one-half meters. Throughout the stream it was always possible to collect E, lepida and P, guttifer. would not seem to support their existence. At no time were conditions found that 20 Table 2. Weather data. Week nggggaggre Precip$§ation l l.67-26.67 2,39 2 -2.78-19.44 .20 3 .56-22.22 1.55 4 1.67-23.33 1.93 5 -3.89-21.11 .08 6 5.00-22.78 4,14 7 4.44-26.67 1.91* 8 16.67-27.78 .. *Exclusive of June data. **Data not available. In the course of the survey, a number of E, guttifer larvae were collected that appeared to be parasitized, though no attempt was made to identify the agent. Muttkowski and Smith (1929) reported 60% parasitism of caddis flies in a stream in Yellowstone National Park, and Stewart, et a1. (1970) found fungi, algae and protozoans on six emerging species in three different trichopteran families. On several other occasions, young, predaceous Phasgano- phora (Plecoptera: Perlidae) larvae were seen to emerge from E, guttifer cases. 21 Caddis larvae are recognized by many (Needham, 1927; Muttkowski and Smith, 1929; Lloyd, 1921; Betten, 1934; Ross, 1944) as important as a source of food for fish. This relationship was certainly established at Vermillion Creek for numerous fish were observed. Quantitative Site Sampling The results of the quantitative sampling are summarized in Table 3. Appendix B provides a complete listing of the results per sample per substrate type per site. 22 Table 3. Summary of biological collections. W No Ave. head Ave. dry No. larvae .capsule weight empty w1dths (mm) (mg) cases Site 1 Sediment Eycnopsyche lepida 2 1.730 41.45 3 E, guttifer 7 1.803 39.51 10 Sand 3, lepida 8 1.736 36.51 7 E, guttifer 11 1.847 41.56 Site 2 Sediment E, lepida 3 1.667 35.17 7 E, guttifer 17 1.774 36.18 14 Sand 3, lepida 2 1.785 42.50 0 E, guttifer 2 1.900 29.25 1 Gravel E, lepida 31 1.784 41.55 E, guttifer 12 1.759 37.37 Submerged plants 2, lepida 0 - - l E, guttifer 43 1.819 31.52 8 Site 3 Sediment P. lepida 1 1.610 36.00 E, guttifer 2 1.810 39.80 10 23 Table 3. (Continued) No Ave. head Ave. dry No. larvae capsule weight empty widths (mm) (mg) cases Sand E, lepida 3 1.647 31.73 24 E, guttifer 10 1.842 42.78 31 Submerged plants E, lepida 0 - - 0 P. guttifer 24 1.771 37.03 1 DISCUSSION Physical and Chemical Monitoring Depth The upper portion of Figure 6 illustrates the weekly fluctuations in depth at the three sites throughout the monitor- ing period, Site 2 was consistently the most shallow, followed by Site 1 and then Site 3. In the case of sites 1 and 3 the stream channel was relatively deep with a distinct drop from each bank to the bed, while the contrary was true at Site 2. In comparing the curves, the responses of sites 1 and 2 are seen to be quite similar, while Site 3 exhibited wide deviations, especially during Week 3. The reason for this is the flooded area to the southeast which empties into the stream at Site 3. The area was created by damming the natural drainage pathways in that portion of the Orange Area. A channel was then dug leading from the northern portion of the flooded zone west to the stream. The 3.48 cm. of rain which fell from Week 3 to Week 4 was more than enough to cause an overflow of the flooded area thereby delivering a disproportionate amount of runoff to Site 3. The amount of rain was apparently not enough to cause a substan- tial overflow of Rose Lake, for Site 2, which receives that over— flow, did not exhibit a response similar to Site 3. However, such 24 Depth (m) Current (m/sec) 25 Figure 6. Depth; current. Week number 1.3;- 1.1.. .9' .7- '5 ’ —Site 1 '- d-Site 2 .3- ---Sne3 .1 -— .. A —Site1 1 - ,/ \ ---51te 2 . ---Site 3 .9 r- .7 - L. .5 -' .3 P .1 - 1 1 1 1 1 1 1 1 2 3 4 5 6 8 26 a response is observed during Week 7 as all sites recorded a sharp increase in depth. During this period, more than 6.05 cm. of pre- cipitation fell on the area. The flooded area, Rose Lake and various intermittent streams all delivered additional flow to Vermillion Creek. The question arises as to the effect of these wide fluctuations on bottom organisms. Moffett (1936), Mottley, gt_gl, (1939), Jones (1951) and Hynes (1970) offered evidence that periods of high water greatly reduce the benthic populations of streams. To be sure, the extent of the effect varies from one stream to another and often within a stream itself due to local conditions. In Vermillion Creek, active P, guttifer larvae could always be seen moving about submerged tree trunks and in the marginal areas during weeks 1 and 2. Following the rise in water level at Site 3, the number of active larvae on the trees and in the margins was noticeably reduced. Many migrated onto the banks with the rising water, often to be stranded as the water level dropped (Hamilton, 1940). Others apparently were swept away by the rising water. Similar depreciations in Pycnopsyche popula- tions were not observed at the other sites. By Week 7, and the second great increase in depth (this time at all sites), 3, guttifer larvae had entered diapause and were firmly attached to a substrate, enabling them to remain during the flooding. At all sites, 3, lgpjda_was in diapause the entire time and unaffected by the variations in water levels. 27 Current As the lower portion of Figure 6 illustrates, the surface velocity exhibited little variation with the exception of the extreme value for Week 6 at Site 2. The concrete bridge there acted as a dam, especially in times of high water. The two large steel drainage pipes beneath the bridge did allow the stream to flow through, but when levels were high, the velocity of the water that did pass through under these Circumstances was greatly increased. Flow velocity is, technically, zero at the stream bed, usually increasing with distance above the bed. Many factors govern the velocity, among which are energy gradient, roughness and depth. At a depth 0.6 of the distance from the surface to the bed, local velocity measurements will yield the mean velocity of the vertical column as a whole owing to the logarithmic change of velocity with distance from the bed (Leopold, gt_gl,, 1964). One consequence of water flow in a channel is the carry- ing of dissolved and suspended material downstream. The effect on water flow of the shape and size (i.e., roughness) of the bed material, the velocity at which individual particles will settle out (settling velocity) and the velocity at which particles are lifted from the bed (threshold velocity), along with channel shape and depth determine what the stream load and substrate will be (Cummins, 1962; Leopold, et a1., 1964). 28 In addition to determining, in part, the substrate characteristics of a stream, water flow has been shown to act directly (Scott, 1958; Hynes, 1970) and indirectly (Gaufin, 1959) in influencing the nature of the benthic fauna. Cummins (1962), however, has shown that an environment with grain sizes approaching .18 mm. and a velocity of 2 cm/sec. will remain relatively stable with very little transportation of sediment. One final aspect of water current is that of driftihg, the significance of which was described by Elliott (1968) and Anderson (1967). Early in the monitoring period P, guttifer larvae were seen to be drifting, though no attempt at numerical estimations was made. With the onset of diapause, drifting in E, guttifer ceased. P_H_ Throughout the eight-week period, the pH at all sites averaged 6.0 with very little variation. Similarly, no diurnal fluctuation was noted. As the stream receives effluent from Rose Lake, the flooded area and marshes, one would expect the pH to be lowered. Slack and Feltz (1968) have demonstrated that pH decreases as organic leaf material is broken down in streams, and Hynes (1960) reported a decrease in pH of waters that flowed through peaty areas. In addition, C02 from the respiration of stream organisms would contribute to the reduction of pH. 29 Apparently a buffering system was operating at Vermillion Creek, maintaining the pH level at 6.0. This value is only slightly lower than those reported by Jones (1949), Macan (1949-1950) and Mackay and Kalff (1969). From a survey of 100 stations on 25 rivers in the United States and Canada over a ten-year period, Roback (1965) reported the pH range of Eycnopsyche to be 6.0-9.0 with the most frequent occurrence of the genus at a value of 8.0. Even though the value of 6.0 for Vermillion Creek is on the lower end of this scale, the presence of stable Eycnopsyche populations indicated a tolerance for that level, and implied that pH is not a major influence on their microdistribution. Turbidity Figure 7 depicts the trends in turbidity for the three sites during the monitoring period. All three positively responded to the high water and increased velocity-~as per Figure 6--during the sixth week. These trends illustrate very well the significance of water flow on the suspended load of a system as discussed earlier. The response for Site 1 can be attributed to terrestial runoff and the scouring effect on the stream bed by the water flow. The same holds for sites 2 and 3 but with the respective additions of overflow from Rose Lake and the flooded area. Increased turbidity can affect the amount of oxygen being produced in a system by reducing the degree of light 30 ‘ 6O - /\ ——Site 1 / \ . l/ .A\‘ —-—S1te 2 /// I 1“ - ---Site 3 . \ V 50 " I, -/ \‘1‘ 7 1 1., I, ,l ‘1‘ 1 [/7 I 11 A 1 E40- ’ \ 1 I, 7/ / 1 \ 5? X I 1 1 7 1 13 ,7 . 1 . a 0 , ./ 1 \ |_3 1- ’ o I, l 1 1 / l 1 \ // l \ 1 I O 20.. ,’ ' 1 \ x / l 1 \ \\ I ' \ \\ \ /I \ \ ./ I T\ \7 0 \ \x / \ \.\' '/\’ \ 10" “"” \ 1 1 1 1 1 1 1 1 1 2 3 4 5 6 7 Week number Figure 7. Turbidity (ppm). 31 penetration, and thereby inhibiting algal photosynthesis (Hynes, 1970). The levels of turbidity and dissolved oxygen found in Vermillion Creek did not seem to indicate that this phenomenon was occurring. Mackay and Kalff (1969), in Canada, found maximum tur- bidity levels following spring thaw and subsequent snow melt, while Slack and Feltz (1967), in Virginia, reported maximum levels in the fall due to increased leaf fall. At best, then, one can state that turbidity levels are a result of a complex of local conditions and are not subject to gross generalizations. Gaufin (1959) found turbidity to be insignificant as a factor influencing the distribution of the bottom fauna of the Provo River in Utah. The same seems to apply to Vermillion Creek. Total Carbon The trends in total water carbon based on the data are presented in Figure 8. Though all three are similar in shape, without further data, their interpretation is difficult. Slack and Feltz (1968) reported that the solute load of a stream can be altered by an increase in leaf fall, but for the time period under investigation here, this is all but negligible. Mustalish (unpublished, 1972) observed in Red Clay Creek, Pennsyl— vania, that total water carbon was intricately related to colloi- dal particles in the stream bed. Through positive adsorption, colloidal particles remove organic material from solution, 32 21 ,1 / 1 / , 1 I I / l / / I / I 19 ’/ ' r l l‘\ // 1 / \\ / ' I \ . l 17 ,’ 1 - 1 l I I 15 ' l E | A Q. 3 13 1 : s 1' ‘E /l/ 'l (U U ‘1 '3 11 K g I 1~ I 9 . 7 Site 1 \ Site 2 N‘ 5 Site 3 “‘ 3 l l 2 3 4 5 6 7 8 Week number Figure 8. Total carbon (ppm). 33 continuing until all adsorption sites are filled. It was found, however, that adsorption of organic carbon was greatly reduced in sandy soils owing to the lack of colloidal particles in such substrates. As the stream bed of Vermillion Creek was nearly exclusively of sand, the trends revealed in Figure 8 cannot be interpreted in terms of substrate adsorption of organic carbon. Conversely, Hynes (1970) states that the amount of organic carbon increases in a stream that drains marshy areas. The overflow from Rose Lake and the flooded zone, and the exis¥ tence of marshes in the drainage area suggest that these are the factors to be considered when attempting to explain the trends. For sites 2 and 3, peaks are recorded during Week 2 and Week 6, the two time periods when, due to much rainfall, overflow from the various external sources was indicated. This is particularly evident at Site 3 during Week 6, and is further substantiated by the results of the trubidity investigation. It would be expected that a rise in turbidity (reflecting an increase in suspended organic and inorganic material) would be positively correlated with the amout of carbon in solution. Again, local conditions must effect the results of such interactions and it is best to be conservative with any generalizations. In Roback's (1965) survey, Pycnopsyche were collected within a 8.0.0. range of 0.5-10.0 ppm, with most found between 1.0-5.0 ppm. Total carbon, in Vermillion Creek, does not seem to be a significant factor in influencing the species' distribution. 1 34 Temperature The lower portion of Figure 9 represents the range of water temperature during the monitoring period. The trend is quite simple, reflecting a general increase in minimum and maxi- mum as the days became progressively warmer with the approach of summer. ".‘I The role of temperature in an aquatic environment is quite varied. It directly influenCes the rate of an organism's 1 development (Hynes, 1970; Flannagan and Lawler, 1972; Gower, 1967; Ulfstrand, 1968), inversely alters the oxygen carrying capacity of water (Hynes, 1970; Hutchinson, 1957) and when variations are wide, decreases overall production in bottom fauna (Gaufin, 1959). In Vermillion Creek, however, it did not appear to influence distributional activity of either species. Pycnopsyche were collected by Roback (1965) in tempera- ture ranges of 12-28°C. with 28°C. exhibiting the highest frequency. Vermillion Creek's lower value of 5.6°C. extends this temperature range and agrees well with Scott's (1970) findings in Africa that many caddis fly species are capable of wide tolerances. Dissolved Oxygen The diurnal dissolved oxygen curves for the three sites over the test period are presented in the upper portion of Figure 9. A general downward trend at all sites was noticed. This, in large measure, was due to the gradual increase in water temperature 35 12- 11» 101- D.O. (ppm) ————-Site 1 . “‘"Site 2 -_-- Site 3 Max. 20- v—\/ . Min. 1 1 1 L 1 4 5 6 7 8 Week Number ml" 1 1 2 Figure 9. 0.0.; temperature. 36 resulting in a subsequent increase in the rate of oxygen consump- tion by the bottom fauna and a decrease in the oxygen carrying capacity of the stream water. In sluggish streams like Vermillion Creek, this situation is compounded by the absence of turbulence. Oxygen is not readily absorbed from or given up to the air. Increasing the organic load only serves to continue the reduction of available oxygen. Two factors balance these effects, though. The first is the fact that the rate of oxygen uptake by the water increases proportionately as the saturation percentage decreases, and the second is the pro- duction of pure oxygen through photosynthesis by green plants in the water. In fact, in times of bright sunshine, sluggish waters can become supersaturated (Hynes, 1960). Since oxygen is a necessity for respiration in most aquatic organisms, and as it often becomes a limiting factor, it has drawn the attention of many investigators (Morgan and O'Neil, 1931; Norris, §t_al,, l964; Wigglesworth, 1966). Knight (unpub- lished Ph.D. thesis) and Gaufin and Gaufin (1961) have conducted extensive studies on the oxygen requirements of stoneflies. A distinct behavioral pattern characterized by body undulations was observed when oxygen levels were reduced under laboratory condi- tions. Similar results using other organisms were reported by Waterman (1960), Philipson (1954), Fox and Sidney (1953) and Mustalish (unpublished, 1973). The series of experiments by Mustalish were conducted on P, guttifer larvae collected at Site 2 of Vermillion Creek. Based on the data, it was concluded that 37 the larger members of the species could tolerate a lower dissolved oxygen level before beginning undulatory movements than could the smaller individuals. Similarly, once these movements began, larger organisms sustained them to lower dissolved oxygen levels before complete metabolic breakdown. Sporadic undulatory movements are common in caddis larvae (Philipson, 1954) but in this series of experiments, 7.63 ppm appeared to be the pivotal value, under which movements became continuous and increased in rate as dissolved oxygen was reduced until a peak was reached. A rapid decline in rate then occurred, followed by death of the organism. The results of this study raise an interesting point, for toward the end of the monitoring period, the morning dissolved oxygen values were well below 7.0 ppm implying that the organisms during the evening hours might be under oxygen stress. It is perhaps a coincidence, though advantageous nonetheless, that the life cycle of E, guttifer is such that it enters diapause at just about the time dissolved oxygen values begin reaching these low levels. P, lgpjgg, on the other hand, had been in diapause since February and with emergence in late summer would also miss these low oxygen periods. This conclusion, though, is made with reser- vation for caution must be exercised when making ecological state- ments based on laboratory experiments. Gaufin (1959), working with turbulent waters, found dis- solved oxygen to be unimportant in explaining differences in 38 bottom fauna. This would be expected as turbulent waters are nearly always at the saturation point (Hynes, 1960). The range of dissolved oxygen values found by Roback (1965) for Pycnopsyche was 7.0-15.0 ppm with 9.0—10.0 ppm exhibit- ing the highest frequency of organisms. In Vermillion Creek, therefore, oxygen levels were low toward the end of the period a by Roback's standards, and they can be expected to continue their decline somewhat more as water temperature and biological influ- 1 ences continue to rise. To explore the possibility of local variations in oxygen levels at each site, a series of determinations were made over each substrate division. The results are listed in Table 4, and indicate that only minor variations were observed. Distributional influence was not apparent during the eight week period, but the downward trend in oxygen levels cer- tainly suggests a potentially perturbational situation, especially in light of past evidence. Final resolution, however, must await additional investigation. Other Factors A number of parameters often considered important facets of aquatic systems were not monitored in this study, among which were 002, M.0.-alkalinity and total hardness. Evidence by Mackay and Kalff (1969) and Roback (1965) suggested that Pycnopsyche were quite tolerant with respect to these parameters, and as it was 39 Table 4. Intrasite variation in C.0. (ppm). A.M. P.M. Site 1 Sediment 5.72 7.16 Sand 5.51 7.00 Site 2 Sediment 5.99 7.69 Sand 6.15 7.84 Gravel 6.20 8.00 Submerged plants 6.10 7.74 Site 3 Sediment 6.20 6.92 Sand 6.31 7.31 Submerged plants 6.47 7.42 judged unlikely that any extremes would be encountered at Vermillion Creek, the decision was made not to monitor them. For the sake of completeness, however, the ranges for the above as reported by Roback (1965) are listed below. The value in parenthesis was that with the highest Pycnopsyche frequency: 002: 5-50 ppm (10 ppm) M.O.-a1ka1inity: 10-550 ppm (100 ppm) Total hardness: 5-1000 ppm (150 ppm). 40 Quantitative_Site Sampling Frequency Distribution of Head Capsule Widths To determine the degree of intraspecific interaction in habitat distribution, the head capsule width data were subjected to a quantitative frequency distribution analysis. If the result- ing distributions could be considered asymetrical, it might indi- cate that there was selection for Or against particular associa- tions by individuals with members of their own species. The data were treated on three levels-~the stream as a whole, the individual sites and the particular substrate types-- the results of which are presented in Figure 10A for P, guttifer and Figure 108 for E, lgpjgg, With a null hypothesis that there was no intraspecific influences on individual habitat distribution, a chi-square test was performed on the frequencies. On all levels, and for both species, there was no significant evidence, with 95% confidence, to cause a rejection of the null hypothesis. FreguengygDistribution of Dry Weights The frequency analysis and chi-square evaluations con- ducted on the head capsule measurements were performed on the dry weight data as well for both species. Figure 11A and 118 present the histograms for E, guttifer and E, lepida respectively. Once .Lmewppam ”m11mgpnwz mfizmamo 6mm; mo cowpsawepm_v xucmzcmgm .mgm 1ooF ucmm -oo_ pcmewvmm ..oop m.P o.P m.p m._ m._ P r m. .., .. ..lfl FL 1 om m._ - m._ .m._ [ I l L L__ uotaelndod [9101 go auao Jad LLLI cm l CD 1.0 l J I I I m 66.6 cop N eeam - co, 2 a... 1 co. Eeeeem Loo_ 42 .enwmmF um-1;pnw3 m_:mamo vmm; mo :oepznwepmwu xucmzcmgm m at... AEEV :puwz mpzmamo cam: .mo_ mgsmwm m.F m._ m.— m.~ m.— m.~ m._ o.F m.p . _ — - .4 . n 1 q . , n q q A _ . . q . 1cm - om . om 1 . Fl 1 Pm>mem .Hoc_ ucmm H 00? ucmewumm 1 oop m.F m._ m.F m._ o.— m.~ . - m.F m._ m._ . - , d n d a 4 _IL - a q - - q . IL I 8 - om I 1 8 ..0m {cop uoIleLndod [9101 go quao Jad 43 “ -. 4 r‘ .mewppsm am--mp;mwm3 zgc mo cowuznvgpmwu xucmzcmgm .<__ mgsmwm Amev mpcmwmz zgo 8. 8 S + om , -om . S W —J n u - h a q . [TL - L x 4 L 0m 1 0m 1 J l I d 9 ... l J I 3 V J ..UU m 3.; . 2: N 3.; . 2:1 . O .... om om S 8 , - .8 E m 44 . ‘ - - , m . . J m... _ _ - d - L m J I m. l P. n u“ I om L om w 4 - - OS -2: _ mpwm Emmgpm 44 - , om om o— L mucmpa vmmLmEnzm . , pm - om - - op uld ‘ — — — [‘4 4 vcmm L om oop om cop AmEV pgmwmz xgo Aum=prcouv .<__ mLsmwu om - - ,om w c_ Fm>mgm . om - . - om . o_ L pcmsvvmm 1 om cop om ooF uogqelndod [9101 ;o nuao Jed 45 .muwmm— um--ma;m:m3 zgv we covpznwgpmwu :ocmzcmgm .m:: mgzmwm Amsv mu;m_m2 ago om om 2 09 cm 2 4. a u q q a . a; a 4 - ‘ - a . L ‘ tom .8 L ‘ I. d ‘ a I In . m m 3.3. . 2: N 8.; .52 w m. om om E + - om , - 0m. 0.: 1 _||_ _ fl 4 - - - ‘ . q . a 0 I 17 1 m. l ‘ . ... d 1 J m L m. - 8 .. om m. 0 l l u L l : 3.; H 2: =52: . 2: 46 .8 Amev uzmwmz Ago om om op ucmm FLHILII om oop :m>mgm Aumzc:p=ouv .m__ mgamwm om oo~ om om or q fi, —_q- pcmswvmm l uogqelndod [2101 ;o nuaa Jed 0 L0 cop 47 again, chi-square indicated there was no statistically significant support for rejection of the null hypothesis. Coefficient of Numerical Separation For two closely related species to coexist in the same environment, it is generally accepted (Ulfstrand, 1968; Grant and Mackay, l969) that they must develop different exploitation pat- terns. The virtual exclusion of E, lgpjga_larvae from submerged plants; yet their relative abundance in graveled areas, in Ver- million Creek, suggested a need for statistical evaluation of interspecific habitat preferences. Egglishaw (1964), Nielson (1942), Décamps (1967), Ulfstrand (1968) and Mackay (l969) each found some aspect of food preference a major microdistributional influence, but with E, lgpiga_and E, guttifer spending a large part of their larval existences in nonfeeding diapause, it seems unlikely that food requirements influence their distributional activities to any great degree. Cummins (l964) used the case-building requirements of the two (mineral particles for E, lgpigg, stick material for E, guttifer) to conclude, in part, that the nature of the sub- strate was limiting their microdistribution. To investigate this possibility, the sampling data were evaluated using the coefficient of numerical separation, as modi- fied by Grant and Mackay (l968), as the test statistic. The coefficient, GS, is given by the formula: GS = loo-iooznmx), 48 where y is the number of the rarer species, x is the number of the more common species and n is the number of subdivisions in each collecting category. A coefficient value of 100 would mean total segregation, while a value of 0 would indicate that each species was represented in equal numbers. The data were subjected to this analysis on three levels-- the stream as a whole, on the basis of sites and on the basis of substrate types. The results are summarized in Table 5. The 60.94 value for the stream was, according to guide- lines established by Grant and Mackay (1969), evidence that large- scale spatial separation existed between the species. The degree of separation, however, was not statistically significant. Examining the values for medium-scale spatial separation, the same conclusion can be drawn with the exception of Site 3 which exhibited a high level of separation. Partitioning the data into substrate categories revealed total segregation at the submerged plants subdivision, a conclusion easily verified by the collection data. Sediment offered the next degree of separation, though the value was not exceedingly strong. The 61.29 entry for gravel was positive but weak, evidence of segregation, while the 43.48 tabulation for sand was low enough to imply distinct species interaction. In evaluating the averages for the three levels, substrate type emerged weakly as the most influential distributional factor, followed by the site and stream levels respectively. Segregation between E, lepida and E, guttifer did not become statistically 49 Table 5. Coefficients of numerical separation. G s Stream 60.94 Site 1 44.44 Site 2 51.35 Site 3 88.89 Average , 61.56 Sediment 76.92 Sand 43.48 Gravel 61.29 Submerged plants 100.00 Average 70.42 apparent until quantitative data were analyzed on a microhabitat level, in this case substrate type. A question now arises as to the possible stimulus for such segregating activity in Vermillion Creek. An answer is sug- gested by the life cycles of the insects, and in particular, the active migration of terminal instar E, lgpiga_larvae. The stimulus apparently manifests itself as an instinctual need to obtain an optimal site for pupation. With E, lgpiga_migrating out of the marginal areas, these slower water habitats are made available for exploitation by the newly hatching E, guttifer population. The evolutionary advantages given to a species that migrates under these circumstances is obvious, for through migration it removes 50 itself from the possibility of direct competition with closely related species while at the same time allowing maximum energy to pass through a given habitat. Viewed in these terms, the microdistributional pattern of E, lgpjga_and E, guttifer observed in Vermillion Creek, was a consequence of : (a) a temporal lag in their respective life cycles; (b) the instinctual need for optimal pupation sites; (c) the active migration by E, lgpjga_as a response to this innate stimulus. CONCLUSION Percival and Whitehead (1929), Anderson and Hold (1972), Flannagan and Lawler (1972), Hynes (1970), Mackay and Kalff (1969) and Cummins (1964) established the significance of substrate in the microdistribution of aquatic insect; oxygen studies have been conducted by Philipson (1954), Gaufin and Gaufin (1961), Fox and Sidney (1953) and Knight (unpublished Ph.D. thesis); the importance of temperature has been demonstrated by Ulfstrand (1968) and Scott (l970); the influences of water flow has been outlined by Feldmeth (1970) and Philipson (1954); and the role of alkalinity indicated by Armitage (1958). To be sure, these factors and others all enter into the dynamics of an aquatic ecosystem. Many such investi- gations, however, have concentrated on a particular parameter to the exclusion of others. In a natural system nothing can be excluded. The attempt was made in this study to include as many of the significant factors as was feasible, monitor interactions in the system and draw conclusions as to their possible influence on the microdistribution of E, lgpig3_and E, guttifer. This was generally accomplished. Although such factors as pH, temperature, turbidity, total carbon and dissolved oxygen may often be limiting factors in other systems, they were not demonstrated to be signifi- cant in Vermillion Creek during the study. The one possible excep- tion was dissolved oxygen, for while the levels monitored indicated 51 -n I"? 52 no detrimental effect to the organisms, the downward trend, in light of experimental evidence, offered a possibility of the occurrence of oxygen stress to the species being studied. What did emerge was an awareness that substrate characteristics, and indirectly, stream flow and the local geomorphological conditions in association with instinctual requirements within the organisms were a significant influence on the microdistribution of E: lgpjga_ and E, guttifer. Cummins (1964) collected 2, lgpjga_in coarse gravel and pebbles, while photographic analyses of the study area determined the importance of nonmineral substrates to E, guttifer. The strati- fied random sampling, the histogramic analyses of head capsule widths and dry weights and the determination of coefficients of separation in the Vermillion Creek study have substantiated his findings and provided additional information as to the microdis- tributional characteristics of these two aquatic insects. By the very nature of ecological studies the results of this investigation should be viewed with caution, for as Muttkowski and Smith (1929) have expressed, local conditions beget local results. It was a starting point, however, and deemed valuable despite its limited scope. SUMMARY With the possible exception of dissolved oxygen, none of the physical and chemical parameters monitored at the three sites during the eight-week period appeared to influence the micro- distribution of E, lgpjga_and E, guttifer in Vermillion Creek. Throughout the stream, it was always possible to collect E, lgpjg3_and E, guttifer. At no time were conditions found that would not seem to support their existence. Chi-square analyses of head capsule widths and dry weights of all organisms collected indicated no intraspecific influences of the microdistribution of individual organisms. Segregation between E, lgpigg_and E, guttifer, based on sub- strate type, was statistically demonstrated using a coefficient of numerical separation. The microdistributional pattern of E: lgpjgg_and E, guttifer, was a consequence of: (a) a temporal lag in their respective life cycles; (b) the instinctual need for optimal pupation sites; (c) the active migration by E, lgpigg_as a response to this innate stimulus. 53 BIBLIOGRAPHY ‘l .11 BIBLIOGRAPHY Literature Cited Anderson, N. 1967. Biology and downstream drift in some Oregon Trichoptera. Egg, E33, 99: 507-521. Anderson, N., and J. Wold. 1972. Emergence trap collections of Trichoptera from an Oregon stream. 104(2): 189-201. Armitage, K. B. 1958. Ecology of riffle insects of Firehole River, Wyoming. Ecology. 39: 571-580. Betten, C. 1934. The caddis flies or Trichoptera of New York State. W: 1, State figs, Bull. 292: 1-576. Betten, C. 1950. The genus P cno s che (Trichoptera). Ann. ent. Soc. Amer. 43: 508—522. Cummins, K. 1962. An evaluation of some techniques for the collection and analysis of benthic samples with special emphasis on lotic waters. Amer. Midl. Nat. 67: 477- 504. 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The life cycle and ecology of Pycnopsyche gentilis (McLachlan), E, luculenta (Betten), and E, scabripennis (Rambur), (Trichoptera: Limnephilidae) in West Creek, Mont St. Hilaire, Quebec. Unpublished Ph.D. Thesis, McGill University. Mackay, R., and J. Kalff. 1969. Seasonal variation in standing drop and species diversity in a small Quebec stream. Ecology. 50: 101-109. Macan, T. 1949-1950. Survey of a Moorland fish pond. J, Anim. Ecol. 19: 160-186. Moffett, J. 1936. A quantitative study of the bottom fauna in some Utah streams variously affected by erosion. Bull. Univ. Utah. biol. S25, 26: 224-225. Mottley, C. et a1. 1939. The determination of the food grade of streams. Trans. Amer. Fish. Soc. 68: 336-343. Mustalish, R. Unpublished 1971. A survey of the major techniques for determining the amount of dissolved organics in water. Univ. of Penn. Mustalish, R. Unpublished 1972. An analysis of the variations in organic carbon in Red Clay Creek. Univ. of Penn. Mustalish, R. Unpublished 1973. Oxygen tolerance studies of the limnephilid caddisfly genus Pycnopsyche. Mich. St. Univ. Muttkowski, R., and G. Smith. 1929. The ecology of a trout stream in Yellowstone National Park. Roos. Wild. Ann. 2(2): 241-263. Needham, J., and R. Christenson. 1927. Economic insects of some streams of northern Utah. Utah Agric. Exp. Sta. Bull. 201: 3-36. Nielson, A. 1942. Uber die Entwicklung und Biologie der Trichop- teren. Arch. Hydrobiol. Suppl. 17: 255-631. 57 Norris, W., et a1. 1964. Oxygen consumption by caddisfly larvae. ng, J, ng, 16(1): 72-79. Philipson, G. 1954. The effect of water flow and oxygen concen- tration on six species of caddis fly (Trichoptera) larvae. Proc. Zool. Soc. Lond. 124: 547-564. Roback, S. 1965. Environmental requirements of Trichoptera. in Biological problems ig_water pollution. U.S. Dept. H.E.W.: “8'1260 Ross, H. 1944. The caddis flies, or Trichoptera, of Illinois. . Ill. Nat. Hist. Surv. Div., Bull. 23(1): 1-326. Scott, K. 1970. Some notes on the Trichoptera of standing waters in Africa, mainly south of the Zambezi. Hydrobiol. 35(2): 177-195. Slack, K., and H. Feltz. 1968. Tree leaf control on low water quality in a small Virginia stream. Envir. Sci, agg_ Tech. 2: 126-131. Stewart, et a1. 1970. Dispersal of algae, protozoans and fungi by aquatic Hemiptera, Trichoptera and other aquatic insects. Ann. ent. Soc. Amer. 63(1): 139-144. Ulfstrand, S. 1968. Life cycles of benthic insects in Lapland streams. Oikos. 19(2): 167-190. Ulfstrand, S. 1968. Benthic animal communities in Lapland streams. Oikos, suppl. 10. Waterman, T. 1960. Physiology of crustaceans. Acad. Press, New York. Wigglesworth, V. 1966. Insect physiology. Methuen, London. General References American Public Health Association. 1962. Standard methods for the examination of water and wastewater, 11th ed. American Public Health Association, American Water Works Association and Water Pollution Control Federation of Water and Wastewater, New York. Mendenhall, W. 1968. The design and analysis of experiments. Duxbury Press, Belmont, California. 58 Mendenhall, W. 1971. Introduction to probability and statistics. Duxbury Press, Belmont, California. Pennak, R. 1953. Fresh-water invertibrates of the United States, Ronald Press, New York. Sokal, R., and Rohlf, F. 1969. Biometry. W. H. Freeman and Co., San Francisco. United States Department of the Interior. 1970. Study and inter- pretation of the chemical characteristics of natural water. U.S. Government Printing Office, Washington. Usinger, R., ed. 1971. Aquatic insects of California. University of California Press, Berkeley. APPENDICES APPENDIX A APPENDIX A Table 6. Results of all determinations during monitoring. Week 1 Site 1 Site 2 Site 3 0.0. (ppm) A.M./P.M. 1 9.03/13.51 8.73/13.33 8.56/12.74 2 9.38/12.10 9.03/12.69 8.38/10.62 3 6.17/10.62 5.43/8.97 5.49/9.38 Turbidity (ppm) 1 7.014 7.014 8.909 2 8.661 21.017 17.063 3 12.862 23.324 33.044 Total carbon (ppm) 1 6.350 8.437 7.017 2 5.541 6.118 5.260 3 9.329 11.603 9.917 pH 6.0 6.0 6.0 Depth (m) 1 .953 .876 1.168 2 .953 .800 1.193 3 .914 .832 1.168 Current (m/sec) 1 .423 .726 .346 2 .423 .726 .346 3 .423 .726 .346 Temp. range (°C.) 8.0-18.3 8.0-18.3 8.0-18.3 59 60 Table 6. (Continued) W Week 2 Site 1 Site 2 Site 3 0.0. (ppm) A.M./P.M. 7.49/11.92 7.26/12.21 6.67/11.80 9.29/11.45 8.61/10.86 8.91/10.97 8.67/11.80 8.73/10.62 8.79/11.27 Turbidity (ppm) 1 23.818 9.979 13.027 2 15.334 11.133 16.652 3 15.828 13.851 11.297 Total carbon (ppm) 12.218 12.763 11.290 13.525 21.249 13.321 11.290 12.345 13.321 pH 6.0 6.0 6.0 Depth (m) 1 1.156 1.105 1.400 .965 .967 1.245 .813 .660 1.003 Current (m/sec) .215 .610 .435 .215 .610 .435 .215 .610 .435 Temp. range (°C.) 6.7-16.7 6.7-16.7 6.7-16.7 61 Table 6. (Continued) r 1 Week 3 Site 1 Stie 2 Site 3 0.0. (ppm) A.M./P.M. 1 7.91/11.39 7.32/10.27 8.38/9.09 2 8.32/11.09 8.08/10.27 7.91/10.27 3 9.44/12.51 9.20/12.04 9.09/11.62 Turbidity (ppm) 13.686 9.238 44.493 18.381 6.437 10.227 22.088 15.251 17.475 Total carbon (ppm) 1 12.534 13.111 11.578 2 13.470 11.972 11.621 3 14.538 12.070 11.761 pH 6.0 6.0 6.0 Depth (m) .940 .864 1.556 .991 .851 1.283 .864 .737 1.118 Current (m/sec) .254 .663 .314 .254 .663 .314 3 .254 .663 .314 Temp. range (°C.) .6-17.8 5.6-17.8 5.6-17.8 Table 6. (Continued) 62 Week 4 Site 1 Site 2 Site 3 0.0. (ppm) A.M./P.M. 1 7.43/11.45 6.37/11.33 7.49/10.15 2 6.96/11.33 6.49/10.97 6.49/9.97 3 9.68/10.74 10.09/11.33 9.62/11.62 Turbidity (ppm) 1 32.467 9.073 17.805 11.956 9.403 26.371 20.605 16.981 24.724 Total carbon (ppm) 1 23.392 17.384 17.334 13.504 14.847 16.224 13.975 14.151 14.151 pH 6.0 6.0 6.0 Depth (m) .864 .711 1.092 .825 .668 1.028 .787 .610 .965 Current (m/sec) 1 .242 .508 .331 .242 .508 .331 .242 .508 .331 Temp. range (°C.) 9.4-18.3 9.4-18.3 9.4-18.3 F 63 Table 6. (Continued) Week 5 Site 1 Site 2 Site 3 0.0. (ppm) A.M./P.M. 1 8.16/9.91 8.00/10.07 8.16/9.49 2 8.69/10.81 8.90/10.34 8.90/10.02 3 7.95/9.75 8.27/10.60 8.06/9.33 Turbidity (ppm) 52.319 38.563 41.857 21.759 23.077 50.259 42.681 40.869 63.604 Total carbon (ppm) 14.516 15.591 16.133 15.872 15.381 15.205 17.279 14.397 14.151 pH 6.0 6.0 6.0 Depth (m) .711 .533 .851 .673 .495 .838 .610 .445 1.016 Current (m/sec) .148 .435 .339 .148 .435 .339 .148 .435 .339 Temp. range (°C.) 6.7-16.7 6.7-16.7 6.7-16.7 Table 6. (Continued) 64 Week 6 Site 1 Site 2 Site 3 0.0. (ppm) A.M./P.M. 1 7.21/9.01 7.31/8.59 7.37/8.27 2 7.53/8.48 7.00/8.22 7.42/8.37 3 6.15/7.79 5.72/7.26 6.30/7.05 Turbidity (ppm) 1 35.103 35.103 29.666 2 60.144 82.714 85.350 44.988 51.907 60.227 Total carbon (ppm) 1 13.132 14.607 14.115 14.326 16.069 16.505 21.109 16.787 17.327 pH 6.0 6.0 6.0 Depth (m) 1 .711 .521 .991 2 .711 .533 .864 3 .864 .787 1.041 Current (m/sec) 1 .381 1.220 .423 2 .381 1.220 .423 3 .381 1.220 .423 Temp. range (°C.) 11.7-18.9 11.7-18.9 11.7-18.9 65 Table 6. (Continued) W Week 7 Site 1 Site 2 Site 3 0.0. (ppm) A.M./P.M. 1 6.78-8.27 6.25/8.43 6.47/8.11 2 7.42/8.48 7.00/8.27 7.10/8.06 3 6.68/7.95 6.04/7.79 6.04/7.53 Turbidity (ppm) 1 14.016 14.510 66.569 14.510 22.747 13.521 11.709 16.816 14.510 Total carbon (ppm) 17.455 20.434 20.722 19.105 16.646 19.528 9.688 9.899 25.009 pH 6.0 6.0 6.0 Depth (m) 1 1.118 .991 1.422 2 1.105 .991 1.397 .927 1.156 1.181 Current (m/sec) 1 .288 .677 .359 .288 .677 .359 .288 .677 .359 Temp. range (°C.) 10.0-21.1 l0.0-21.l 10.0-21.l 66 Table 6. (Continued) W Week 8 Site 1 Site 2 Site 3 0.0. (ppm) A.M./P.M. 1 5.83/8.16 5.35/7.84 5.41/7.31 2 5.41/7.90 4.98/8.37 5.62/8.11 3 4.82/8.16 5.04/8.64 5.19/7.53 Turbidity (ppm) 33.785 11.050 14.345 34.114 11.050 4.955 3 54.049 22.253 8.414 Total carbon (ppm) 9.828 15.254 10.075 20.080 9.020 12.141 10.075 10.110 9.478 pH 6.0 6.0 6.0 Depth (m) .927 .794 1.143 .864 .750 1.067 .787 .737 .965 Current (m/sec) .321 .762 .508 .321 .762 .508 .321 .762 .508 Temp. range (°C.) 16.1-23.3 16.1-23.3 16.1-23.3 APPENDIX B APPENDIX B Table 7. Results of biological collections. —: _:‘ Head Dry capsule weight width (mm) (mg) Case only None Site 1 Sediment Sample 1 Pycnopsyche lepida E, guttifer Sample 2 E, lepida E, guttifer Sample 3 E, lepida E, guttifer Sample 4 E, lepida E, guttifer 1.78 37.5 Sample 5 p, lepida 1.73 37.9 " 1.73 45.0 E, guttifer Sample 6 E, lepida E, guttifer 1.84 31.0 Sample 7 E, lepida E, guttifer 1.78 33.0 67 68 Table 7. (Continued) Head Dry capsule weight width (mm) (mg) E, guttifer 1.67 47.6 " 1.96 54.0 " * Sample 8 E, lepida * E, guttifer 1.75 38.0 " 1.84 35.5 Sample 9 E, lepida . * E, guttifer * Sample 10 E, lepida * E, guttifer * n * Sggg_ Sample 1 E, lepida 1.84 35.1 P, guttifer 1.79 42.9 Sample 2 E, lepida * P, guttifer 1.84 40.0 Sample 3 E, lepida 1.68 42. " 1.68 33. " 1.84 40. " 1.96 65. E, guttifer * II * 010C301 II * 69 Table 7. (Continued) m: cagggIe weIENt Case None width (mm) (mg) only Sample 4 E, lepida * E, guttifer 1.96 47.0 n * Sample 5 E, lepida ' * E, guttifer * Sample 6 E, lepida 1.61 29.0 n * E, guttifer 1.84 48.2 " 1.84 52.0 " 1.84 50.0 " 1.73 32.5 Sample 7 E, lepida * u * E, guttifer * Sample 8 ' E, lepida 1.67 25.0 " 1.61 22.0 H 'k E, guttifer 1.90 37.5 " 1.90 34.5 ” 1.84 33.5 " 1.84 39.0 Sample 9 E, lepida * E, guttifer * 70 Table 7. (Continued) _ 4 Head Dry capsule weight width (mm) (mg) Sample 10 3, 1e ida E, uttifer n Site 2 Sediment Sample 1 E, le ida E, uttifer f .50 8.1 1 Sample 2 2, 1e ida E, guttifer 1.84 34.0 Sample 3 P. 1e ida P. uttifer 1.79 42.9 1° 11 Sample 4 P. 1e ida E, guttifer Sample 5 E, 1e ida E, guttifer 1.84 49.4 " 1.84 49.4 ” 1.79 38.5 " 1.78 33.5 1° I“ 71 Table 7. (Continued) M caggfiIe weIQNt Case None width (mm) (mg) only Sample 6 E, lepida 1.73 48.0 " 1.61 24.5 E, guttifer 1.73 27.5 " 1.73 25.8 " 1.98 22.5 " 1.73 36.0 " 1.86 42.3 " 1.63 32.0 ” 1.73 55.5 " 1.61 24.5 ll * Sample 7 E, lepida * ll * II * u * ll * E, guttifer * Sample 8 E, lepida * E, guttifer * II * Sample 9 E, lepida * E, guttifer 1.87 49.1 ll * ll * Table 7. (Continued) Head capsule width (mm) (mg) 72 Dry weight Case only None Sample 10 E: lepida 1.66 E, guttifer 1.90 Sand Sample 1 E, lepida 1.79 E, guttifer Sample 2 E, lepida E, guttifer Sample 3 E, lepida E, guttifer Sample 4 E, lepida E, guttifer Sample 5 E, lepida E, guttifer Sample 6 E, lepida E, guttifer Sample 7 E, lepida E, guttifer Sample 8 E, lepida E, guttifer Sample 9 E, lepida 1.78 E, guttifer 1.96 33.0 44.5 38.0 47.0 23.4 Table 7. (Continued) 73 Head capsule width (mm) Dry weight (mg) Case only None Sample 10 E, lepida E, guttifer Gravel Sample 1 E, lepida E, guttifer Sample 2 E, lepida E, guttifer Sample 3 E, lepida E, guttifer Sample 4 E, lepida E, guttifer Sample 5 g, lepida E, guttifer 1.84 1.84 1.73 1.84 1.78 1.68 1.61 1.96 1.78 1.75 35.0 45.4 35.5 38. 47. 26. 43. 44. 49. 01000010 40.3 Table 7. (Continued) 74 caUESIe weggfit Case None width (mm) (mg) °"‘Y Sample 6 E, lepida 1.82 53.1 " 1.84 46.2 " 1.84 42.0 ” 1.84 42.4 ” 1.84 44.9 " 1.93 55.2 11 * E, guttifer 1.86 46.0 " 1.61 27.5 " 1.84 50.2 Sample 7 E, lepida 1.70 28.5 " 1.61 27.0 E, guttifer 1.84 55.0 " 1.61 32.5 " 1.78 37.0 " 1.84 29.0 Sample 8 E, lepida 1.70 32.0 ” 1.84 48.0 " 1.79 48.0 " 1.86 47.5 ll * E, guttifer 1.84 30.5 " 1.73 26.0 Table 7. (Continued) 75 caESSIe weEgfit Case None width (mm) (mg) only Sample 9 E, lepida 1.61 37.4 " 1.96 56.3 ” 1.93 61.3 " 1.70 29.0 " 1.93 46.5 " 1.63 25.0 " 1.75 31.5 " 1.78 38.5 ” 1.84 49.0 E, guttifer 1.73 40.5 " 1.68 34.0 Sample 10 E, lepida 1.73 33.5 " 1.61 36.5 11 * E, guttifer * Submerged plants Sample 1 E, lepida * E, guttifer 1.75 32.5 " 1.96 28.6 " 1.96 36.5 " 1.78 18.0 11 * 11 * 11 '1: Sample 2 E, lepida * E, guttifer 1.75 20.0 ” 1.61 14.0 76 Table 7. (Continued) caEESIe weggfit Case None width (mm) (mg) °"‘Y Sample 3 E, lepida * E, guttifer * Sample 4 E, lepida * E, guttifer 1.63 22.0 " 1.84 19.5 ” 1.98 33.0 " 1.63 18.0 " 2.07 29.5 ” 1.85 39.5 11 * ll * Sample 5 E, lepida * E, guttifer 1.84 32.0 " 1.79 37.0 " 1.61 35.5 " 1.98 33.5 " 1.84 35.5 " 1.84 32.5 " 1.84 32.5 " 1.84 46.0 " 1.96 27.0 H * Sample 6 E, lepida * E, guttifer 1.91 25.0 ” 1.73 29.0 Table 7. (Continued) 77 caEESIe weggfit Case None width (mm) (mg) only Sample 7 E, lepida * E, guttifer 1.73 45.5 ” 1.86 32.5 Sample 8 E, lepida * E, guttifer 1.84 37.5 " 1.84 35.0 " 1.87 28.0 ” 1.96 35.0 ” 1.84 39.0 ” 1.84 52.5 " * 11 * Sample 9 E, lepida * E, guttifer 1.61 33.5 " 1.84 40.6 ” 1.84 21.0 " 1.73 40.0 " 1.77 28.5 " 1.84 31.0 Sample 10 E, lepida * E, guttifer 1.78 26.0 " 1.63 26.0 " 1.84 28.0 “ 1.84 25.5 ” 1.90 48.0 " 1.84 23.5 78 Table 7. (Continued) __- -_ Head Dry capsule weight width (mm) (mg) Case only None Site 3 Sediment Sample 1 E, lepida * E, guttifer * Sample 2 E, lepida * E, guttifer * Sample 3 P, lepida * E, guttifer 1.84 45.0 Sample 4 E, lepida * E, guttifer * Sample 5 E, lepida * E, guttifer * ll * Sample 6 P. lepida 1.61 36.0 E, guttifer * Sample 7 P. 1e ida * E, guttifer 1.78 34.6 H * 1° 79 Table 7. (Continued) caSSSIe weISNt Case None width (mm) (mg) °”‘y Sample 8 E, lepida * E, guttifer * Sample 9 E, lepida * E, guttifer * ll * Sample 10 E, lepida * E, guttifer * Sand Sample 1 E, lepida * E, guttifer * ll * n * Sample 2 E, lepida * n * II * 1| * E, guttifer * Sample 3 E, lepida * II * ll * 11 'k P. guttifer * u * 80 Table 7. (Continued) Head Dry capsule weight 52?; None width (mm) (mg) E, guttifer * u * 11 'k n * ll * Sample 4 P. lepida * " * " * n * E, guttifer 1.96 51.1 “ 1.82 34.5 11 *- n * " * Sample 5 E, lepida 1.61 36.2 " 1.72 33.5 " * E, guttifer 2.01 50.1 " 1.75 38.5 " 1.84 46.6 Sample 6 E, lepida * E, guttifer 1.61 31.5 " * n 'k 81 Table 7. (Continued) Head Dry capsule weight 3:16 None width (mm) (mg) y Sample 7 E, lepida * u * 1| * 11 * II * ll * n 'k E, guttifer * II * n * ll * Sample 8 E, lepida 1.61 25.5 II it E, guttifer 1.84 48.0 n * 11 * Sample 9 E, lepida * u * E, guttifer * n * 11 * II * II * 1| * 82 Table 7. (Continued) Head Dry capsule weight width (mm) (mg) Case only None Sample 10 E, lepida * E, guttifer 1.84 33.5 " 1.96 43.0 " 1.79 51.0 Submerged plants Sample 1 E, lepida * E, guttifer 1.84 38. " 1.66 37. ” 1.84 29. " 1.96 30. " 1.78 48. Sample 2 E, lepida * E; guttifer 1.81 45.5 " 1.61 30.0 H * h-DOOO Sample 3 E, lepida * E, guttifer * Sample 4 E, lepida * E, guttifer * Sample 5 E, lepida * E, guttifer * 83 Table 7. (Continued) caSEEIe weIQNt Case None width (mm) (mg) only Sample 6 E, lepida * E, guttifer 1.79 33.5 Sample 7 E, lepida * E, guttifer 1.55 14.0 " 1.84 44.2 ” 1.96 44.1 " 1.84 53.5 ” 1.61 33.8 " 1.79 35.4 Sample 8 E, lepida * E, guttifer * Sample 9 E, lepida * E, guttifer 1.84 52.0 " 1.91 34.0 " 1.96 37.5 ” 1.86 45.1 ” 1.38 17.0 Sample 10 E, lepida * E, guttifer 1.50 24.6 " 1.73 50.0 " 1.75 43.6 " 1.87 37.0 " 1.84 31.0 ‘..12- ‘OF— 1.4--..0 x: _ MICHIGAN STATE UNIVERSITY LIBRARIES '1! II III 3 1293 | 3111111111 43 0402