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1988 Invertebrates Associated with the Pincushion ( glaucum) and Underlying Soil Timothy R. Leffler Eastern Illinois University This research is a product of the graduate program in Zoology at Eastern Illinois University. Find out more about the program.

Recommended Citation Leffler, Timothy R., "Invertebrates Associated with the Pincushion Moss (Leucobryum glaucum) and Underlying Soil" (1988). Masters Theses. 2578. https://thekeep.eiu.edu/theses/2578

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Date Author

m Invertebrates Assoc iated with the Pincushion

Moss (Leucobryum glaucum) and Underlying Soil

(TITLE)

BY

Timothy Leffler R.

THESIS

SUBMITIED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF

Masters of Zoology

IN THE GRADUATE SCHOOL, EASTERN ILLINOIS UNIVERSITY CHARLESTON, ILLINOIS

1988 YEAR

I HEREBY RECOMMEND THIS THESIS BE ACCEPTED AS FULFILLING THIS PART OF THE GRADUATE DEGREE CITED ABOVE

l!�l'fg'ATE ABSTRACT

A faunal study of soil invertebrates collected from pincushion moss (Leucobryum glaucum) and underlying soil

in a mixed deciduous forest of Charleston, Illinois, was

performed because it was discovered that moss habitats

have been largely overlooked by investigators.

The results from this study suggest that moss habitats

provide a buffer area from the external physical environment,

provide protection from predators, a place for oviposition

and may be used as a food source. Twenty-seven orders

of invertebrates were collected (66,023 individuals).

Acarina (57.9%) and Collembola (37.9%) made up the majority

of the invertebrate population. Three abiotic factors

(temperature, moisture content and pH ) were considered

in relation to their effect on the fauna. Collembola

populations fluctuated more than the Acarina population

possibly indicating that they were more sensitive to

environmental fluctuations than were the Acarina. This

sensitivity may be attributed to the behavioral and

anatomical diff erences. It is evident from the literature

and this research that there is a need for more work on

moss/invertebrate associations. INTRODUCTION

This paper is the result of a one year study of the invertebrates associated with the pincushion moss,

Leucobryum glaucum, and the underlying soil. Both abiotic and biotic factors which affect soil invertebrates are considered.

Gerson (1972) reported that Ledermuellaria frigida

(Acarina: Stigmaeidae) could not survive on the leaves of g. glaucum. This was the only reference found mentioning

L. glaucum and soil invertebrates.

The association between and arthropods has been largely neglected by botanical as well as zoological ecologists (Gerson, 1969). Aoki (1967) stated that most soil zoologists put soil samplers on the surface of the forest floor or in soil at flat places, intentionally avoiding stones, rocks, tree stumps, rotten wood, patches of moss, etc. These avoided habitats contain some peculiar which are scarce or absent in other habitats.

Cloudsley-Thompson (1958) dismissed mosses as being a natural habitat as "it is clear that moss does not form a biotope with a natural microclimate." However, Richardson

(1981) states that the microclimate in moss cushions and carpets is much less extreme than that found immediately

1 2

above the moss. Norgaard (1951), investigating spider distributions in a Danish sphagnum bog, discovered that moss carpets modified temperature and humidity, thus affecting the spatial distribution of arthropods.

Bry-0phytes have the ability to ab sorb large amounts of moisture. Corbet Lan (1974) soaked Grimmia pulvinata & for thirty minutes in distilled water and discovered that the clumps of dry moss took up, on the average, 4.38 times their own weight of water. Water is held between the stems, between the leaves on each stem, and within the cells. Moisture content seems to be one of the most important abiotic factors governing the success or failure of many different types of soil invertebrates. Verhoef

Witteveen (1980) stated that, for the most part, Collembola & are very susceptible to dessication. Because many species inhabit moi st areas, neither control and limitation of diffusion of water through the cuticle, nor speciali zed adaptations to dessication are found. Testerink (1982) found that Tomocerus minor would die when it was separated from its food by a dry area. Bhattacharya Raychaudhuri & (1979) considered moisture content of the soil to be the most important abiotic factor responsible for microarthropod population fluctuations in a wasteland of Santiniketen,

India.

Soil invertebrates utili ze bryophytes for food. Byers 3

(1961) found that bryophytes harbor craneflies, including the genera Tipula, Liogama, Erioptera, Dolichope za, and

Orope za. Some species in these genera feed on the bryophytes with which they are associated. Gerson (1972) collected one hundred-sixty moss samples, representing fifty- five species of mosses; he found thirty-eight species that contained Ledermuellaria (Acarina:Prostigmata:

Stigmaeidae). Ledermuellaria frigida, k· rhodomela, k· clavata, and k· schusteri were found to feed on various mosses. Loria Herrnstadt (1980) found harvester & ants in Israel climbing the setae of Aloina aloides and

Bryum bicolor and cutting off their capsules. It was surmised that the capsules served as a primary food source, until phanerogam seeds became available. Woodring (1963) reared certain cryptostigmatid mites on a diet consisting entirely of mosses.

Gerson (1982) stated that invertebrates which feed

on bryophytes, usually oviposit there. Ledermuellaria

frigida usually laid eggs in the axils of the moss leaves

(Gerson, 1972). The cranefly Dolichope za deposits eggs

within certain mosses such as Sphagnum, Tetraphis, and

Orthotricum on which larvae will feed. Keeley (1913)

found mite eggs in the empty capsules of Orthotricum

pusillum.

Richards Davies (1977) stated that twenty-one out & 4

of twenty-nine insect orders are associated with bryophytes.

Butcher et. al. (1971) stated that biologists in the United

States, with the exception of taxonomists, have concentrated their research on soil arthropods which directly affect yield and quality of crop . Noneconomic forms are given little attention even though they are important contributors to fundamental humification processes. Edwards et. al. (1970) stated that "Soil animals contribute to the breakdown of organic matter by disintegrating and animal tissues, making them easier to invade by microorganisms, selectively decomposing and chemically changing parts of organic residues, transforming plant residues into humic substances, increasing the surface area available for bacterial and fungal action, forming complex aggregates of organic matter with the mineral part of the soil and mixing the organic matter thoroughly into the upper layers of soil. " 5

MATERIALS AND METHODS

The study was conducted at Lakeview Park which is located in Coles County three kilometers southeast of

Charleston, Illinois. Lakeview Park is hilly terrain covered by a mixed deciduous forest. The dominant trees are Quercus alba. The understory consists mainly of Marus rubra, Acer saccharum, Fraxinus americana, Ostrya virginiana, and Amelanchier arborea. The soil type within the study area is a strawn soil type which is light colored, well drained and silty (Hallbick, 1968).

A total of one hundred-eighty samples of Leucobryum

glaucum were taken between December 4, 1982, and November

30, 1983. Five 7.5cm x 7.5cm x 2.5cm samples of glaucum 1· with soil substrate were taken each month from north-, south-

and west-facing slopes. Two or three samples were taken

during the first half of the month with the other two

or three taken during the second half of the month.

Moss samples were chosen from pure stands of L. glaucum

measuring not less than 12cm x 12cm. All samples were

taken between 10:00 a.m. and 2:00 p.m. and placed in a

Glad Ziplock bag (Union Carbide Corporation), labeled

and then returned to the lab. The samples were weighed

on an oHaus triple beam balance and placed in a Tullgren 6

apparatus to extract invertebrates from each sample.

Glass jars containing 70% isopropyl alcohol were placed under each funnel so that the invertebrates would fall into the alcohol after being driven down by heat created by a fourty watt light bulb above the funnel. Samples were left in the Tullgren apparatus for at least seventy­ two hours. Following extraction, the invertebrates were stored in 70% isopropyl alcohol, and labeled for later identification. Potassium hydroxide was used to clear some of the invertebrates for proper identification.

Air, moss, and soil temperatures were taken at the collection site with a battery operated Markson IV thermometer. Moisture content was measured in grams by subtracting the dry weight of the sample from the wet weight.

Hydrogen ion concentration of the moss samples was investigated at the end of the study. Three gram portions of moss were taken from each sample and placed in 250ml

beakers which were washed thoroughly with distilled water.

Samples were saturated with thirty-five milliliters of

distilled water and allowed to stand for approximately

one hour. The pH of the sample was then measured with

an Orion Research !analyzer/Model 407A.

Statistical analysis was performed using the Pearson 7

Product moment correlation. Correlation analyses between pH and number of individuals; temperature and number of individuals; and percent moisture content and number of individuals were compared.

Temperature and precipitation readings were recorded throughout the entire study at Dr. Dalias Price's weather station, located approximately 2.0 miles from the study site (Figure 1). Three temperature readings were also taken at the study site (for each sampling date); one inch above the moss, within the moss, and at the moss/soil interface. 8

RESULTS

Sixty-six thousand twenty-three invertebrates were collected from one hundred eighty moss samples between

December 4, 1982, and November 30, 1983 (Figure 2) ; sixty moss samples were taken from each of the north-, south­ and west-facing slopes. (Table 7). Twenty-seven orders of invertebrates were collected (Table 1). Twenty-nine thousand one hundred thirty-seven invertebrates (44.1% of the total population) were taken from the south slope,

20, 714 invertebrates (31.4%) were taken from the west slope, and 16, 170 invertebrates (24.5%), were taken from

the north slope. Mites accounted for 57.9% of the total

invertebrate population and Collembola for 37.9% (Figure

3) .

Of the mite population, 29.5% was associated with

the north slope, 32.7% with the south slope, and 37.8%

with the west slope. Oppioidea and Oribatuloidea made

up 43.0% and 41.0% respectively, of the total mite population

(Figure 4) .

Of the Collembola population 15.8% wa s associated

with the north slope, 63.2% with the south slope, and 21.0%

with the west slope. The Collembola population was

dominated by Hypogastruridae and Isotomidae which constituted

61.0% and 22.0% respectively, of the total Collembola 9

population (Figures 5, 6, and 7).

The class Insecta accounted for 27,059 individuals.

Ninety-three percent (25, 194) were of the subclass

Apterygota, and 7.0% (1, 865) were of the subclass Pterygota.

The ma jority of the apterygotes were Collembola.

Seventy-nine percent (4 orders; 1, 467 individuals) of the pterygote insects were endopterygotes and 21.0%

(6 orders; 398 individuals) were exopterygotes.

Three hundred thirty-six immature insects were collected; 62.0% (207) were of the Order Diptera. The family Chironomidae accounted for 33.0% (68 individuals) of the Diptera larvae. One hundred and eighteen immature insects were associated with the north slope, one hundred sixteen with the west slope, and one hundred two with the south slope. The greatest number of immature insects was found in December and November; the fewest number found from June to September (Figure 8).

The greatest amount of precipitation occurred during

March and October; the smallest amount occurred during

July and August. The highest average temperatures occurred during July and August; the lowest occurred during January and February.

During the winter months, air temperature averaged

2.39°C cooler than the temperature found within the moss and 3.49° C cooler than the temperature found within 10

the moss/so il interface. Dur ing the spring months air temperature was on the average 2.42°C cooler than that found within the moss and 2.95°C cooler than that found with in the moss/soil interface. During the summer months the air temperature was on the average 1.07°C warmer than that found within the moss and 1. 88°C warmer than that found with in the moss/so il interface. Dur ing the fall months the air temperature was on the average 1.26°C warmer than that found within the moss and 1.49°C warmer than that found with in the moss/so il interface.

Average percent mo isture content of the moss samples was highest in the winter and spr ing seasons. A general decrease followed from June to September, followed by an increase in October to November (Figure 9). The lowest percent mo isture content reading for a single sample was recorded from a west-fac ing slope on August 30, 1983, (7.8%), wh ile the highest reading for a single sample was recorded from a south-fac ing slope on July 31, 1983, (67.9%) follow ing a ra in storm.

Hydrogen ion concentrat ion was recorded for north-, south- and west-fac ing slopes. The highest pH recorded was 5.2, on north- and west-facing slopes during February,

1983. The lowest pH recorded was 3.6 on a west-fac ing slope dur ing August.

Correlat ion analyses between pH and number of ind iv iduals 11

yielded two significant correlations, involving Collembola from west-facing slopes for the entire year and also for winter-spring seasons only (Table 3). Analyses comparing temperature and the number of individuals yielded two sig­ nificant correlations. The first involved the total number of Collembola found associated with the north slope during the winter-spring seasons. The second significant correl­ ation involved the total number of Isotomidae during the winter-spring seasons (Table 4). Analyses between percent moisture content and number of individuals yielded nine significant correlations (Table 5). These included total number of immature insects for the entire year; inverte­ brates associated with west-facing slopes during winter­ spring seasons and invertebrates from all slopes during winter-spring seasons. Also significant were Collembola associated with south-facing slopes during spring-summer seasons; with south-facing slopes during spring-summer­ fall seasons; with west-facing slopes during spring­ summer-fall seasons; with west-facing slopes during the entire year and total Collembola during the entire year.

Finally, Hypogastruridae associated with west-facing slopes during the entire year was also significant. There were no significant correlations involving Acarina and any of the above abiotic factors. 12

DISCUSSION

The greatest number invertebrates occurred in February,

1983, (Figure 2) . Fifteen thousand seven hundred ninety­ three individuals were counted. Collembola accounted for 83% (13, 094 individuals) of all invertebrates found during this month. Two Collembola aggregations were found on February 26, 1983; a total of 11, 135 hypogastrurids were collected from the south slope and 1, 003 hypogastrurids were taken from the west slope. One other aggregation was observed which occurred on October 13, 1983, and included

1, 649 individuals of Oppioidea from a west-facing slope.

A variety of studies pertaining to aggregations of invertebrates, especially Collembola and mites, have given reasons for this phenomenon. Usher (1975) concluded that the aggregation tendencies noted in the literature can be attributed to "water, temperature, time of day, microclimate, season, food source, microflora, vegetation, clustering of eggs." One possible function of aggregations

may be to help aid in reproduction of the species. Verhoeff et al. (1977) stated that as mating in Collembola is indirect, aggregation of both sexes may enhance the efficiency of reproduction. To find places conditioned

by Collembola (aggregation sites) a pheromone, called

an "arrestant", is released (Dethier et al., 1960) . This 13

causes an orthokinetic behavioral reaction to the pheromone sources (Verhoef f et al. , 1977).

The average air temp erature from February 14, 1983, to February 28, 1983, was 6. 05°C. This is in the range

(6° C to 12°C) stated by Christiansen (1964) as required for normal full develop ment. Snider (1972) has found considerable evidence which suggests that temp erature has an ef f ect on Folsomia candida in relation to sp eed of growth, maturation, and egg production.

Two signif icant correlations were observed when analyzing temp erature and number of organisms (Table 4).

Total Collembola on north-f acing slop es during winter and spring and Isotomidae on north-, south- and west­

facing slop es during the same period showed significant

correlations. This indicates that these two group s from

their resp ective slop es and seasons increased in number

as the temp erature increased. All other combinations

(varying slop es and seasons) showed no signif icant results.

It has been shown that, fluctuations in invertebrate

pop ulations are due to the ef f ects of physical and chemical

environmental factors. Levings and Windsor (1984) found

a positive correlation between litter moisture content

and ants, beetles, collembolans, homop terans, isop ods,

and millip edes. Durant and Fox (1966) found that arthrop od

pop ulation densities showed no signif icant correlation 14

with soil temp eratures. However, pop ulation densities were inf luenced by soil moisture. Joose (1981) stated that "In a habitat where moisture is heterogeneously distributed and wet and dry seasons alternate and wh ere

Orchesella cinta and Tomocerus minor co-exist, Orchesella dominates, " due to various cap abilities (i. e. physiological resistance to desiccation, flex ible rep roduction, etc. ).

Plowman (1979) found that the distributions of Collembola and mites were inf luenced by the percentage of water and the dry weight of litter in two Australian subtropical

forests. Mukharji and Singh (1970) at Varanasi, India,

found that the microarth rop od pop ulation increased when

both the moisture content and temp erature were high.

Bhattacharya and Raychaudhuri (1979) stated that, the

various factors studied (rainf all of the previous month,

relative humidity, air temp erature at time of samp ling,

soil temp erature at 5cm dep th, moisture content of soil,

organic carbon, conductivity and pH, moisture content

and the temp erature of the soil, rainf all of the previous

month, mean monthly relative humidity and air temp erature)

at the time of samp ling showed a signif icant positive

correlation with the total microarthrop od pop ulation.

The average moisture content for the moss/soil samp les

during February was app roximately 49.0%, which was the

highest reading for any of the months during this study. 15

This is the same month that the two aggregations of

Collembola were discovered. Christiansen (1964) said that the cause of these outbreaks is still uncertain, but it seems clear that they are associated with persistent, unusually damp conditions. This is an imp ortant conclusion, due to the fact that moisture allows organisms to commute from one area to another more freely, whereas, hot and dry conditions restrict the movements of soil inhabiting organisms. Thus, persistent damp conditions would allow the Collembola to utilize areas with food and also utilize other areas which may be sheltered from harsh environmental conditions and/or predators. A total of six ty-six predators

(chilop ods, spiders, beetles, ants, and mites) were collected during February, 1983. This number rep resents only 3. 0% of the total number of predators collected throughout the study. Thus, it appears that during February predation would be at a minimum.

The number of Collembola found in December (477 individuals), January (408 individuals), and Sep tember

(483 individuals) was low compared to other months.

The average temperature for the two winter months was

3. 3°C for December and 1. 2°C for January. These low average temperatures may have had some ef fect on the collembolan population. Christiansen (1964) stated that an effect of low temp eratures start to bring the animals into a state 16

of inactivity somewhere between 4°C and -4° C. To remain active through the winter months it may be necessary for

Collembola to migrate vertically. Hale (1967) stated that in most cases vertical migration occurs during periods of adverse climatic conditions in the up per layers of the soil. Thus, if vertical migration did occur, many of the Collembola may have been below the dep th (2. 5cm) that the samp le was taken.

A decreased collembolan population also occurred in

September. There was an observable decrease in percent moisture content of the samples from June to September.

This would result in a decrease of relative humidity.

Christiansen (1964) stated that Collembola usually require

a relative humidity of over 89%. Thus, to avoid dessication

it may have been necessary for certain groups of Collembola

to migrate vertically into more humid areas. As a result

of this vertical migration these Collembola would be absent

from the samples taken. Mukhurji and Singh (1970) observed

that high temp erature and low moisture decreases the

population of soil arthropods during the months of May

and June in Varanasi, India, due to vertical migration

below 22. 86cm of soil. Of all the abiotic factors tested,

percent moisture content when compared to Collembola showed

the greatest number (six ) of positive correlations (Table 5) .

That is, as the amount of moisture increased so did the 17

collembolan population. Many investigators have interpreted moisture relative humidity as being one of the primary ( ) factors af f ecting soil invertebrate populations. Davies

(1928) has shown that distribution of Collembola is a direct result of humidity and thus Collembola may be used to indicate the amount of soil moisture. Hale (1967) states that Collembola are strongly inf luenced by physical factors which bring about changes in the water content of the substrate and McE. Kevan (1962) stated that. moisture content of the soil is very important to the soil fauna. Darlong and Alf red (1982) , investigating dif f erences in arthropod pop ulation structure in soils of forest and Jhum sites of northeast India found that the mean total population of soil arthropods in all sites was correlated with the moisture content of the soil. Bhattacharya and Raychaudhuri

(1979) stated that moisture content of the soil was considered to be the most important single factor responsible for the population fluctuation of the microarthropods inhabiting soil. Other investigators believe that as moisture content remains at higher levels along with increasing temperatures the soil invertebrate pop ulation will also increase. Usher (1971) said that the two factors of temp erature and precip itation are the most likely to af f ect the populations of soil arthropods.

Predation may also have played a role in the depressed 18

pop ulation level of Collembola. Niijima (1971) observed

a fairly good correlation between the seasonal changes

in numbers of Collembola, and the predators extracted with

them; whether this correlation was directly related to

the prey/p redator interaction or to other conditions wasn't

obvious. The Collembola pop ulation decreased from May

through Sep tember. During this time a total of 1, 815 predators were collected. This number rep resents 59. 7%

of the total number of predators collected during the

year long study. The total number of predators collected

during September, 1983, was 16. 2%. The exp ected decrease

in the predator pop ulation did not occur until October.

The decrease may have been due to a combination of the

decrease in the collembolan pop ulation and the increasingly

cooler temp eratures. Table 6 lists predators and numbers

found per month.

The Collembola pop ulation seemed little af f ected

by pH. Signif icant correlations were found associated

with the west-f acing slop e. However, there were no

signif icant dif f erences in pH readings between slop es

from month to month. Booth and Usher (1984) found

that samp le dry weight, percentage water content and

calcium correlated with the number of arthrop ods in samp les;

whereas, nitrogen, phosp horus and pH were least imp ortant 19

to arthropod communities in a maritime antarctic moss- turf habitat. Christiansen (1964) stated that pH conditi ons have little eff ect upon many of the soil fauna. Bhattacharya and Raychaudhuri (1979) found that pH of the soil showed signi ficant negative correlation with Cryptostigmata,

Collembola and all other invertebrates.

An interesti ng observation was made concerning the collembolan families Sminthuri dae and Neelidae. The

Sminthuridae were totally absent from samp les taken from

June through September. Neeli dae were absent from January through Apri l. Thus it app ears that the Sminthuridae were present during periods of cooler temperatures and higher moisture content while Neelidae are present during warmer temperatures and drier periods. Wallace (1967) reported that, although numerical variation of Sminthurus viridi s in West Australia was aff ected by weather, soil type, botanical composition of the pasture, predation by Bdellodes lapidaria, and scavenging on dead bodies by newly hatched nymphs, pop ulation collapse may actually occur due to accumulation of uric aci d. The absence of the Sminthuridae from the samples coincided with an increase in temperature and decrease in moisture. The greatest number of predatory mites (Eupodidae, Bdellidae, Cunaxidae, and Rhagi diidae) was also found duri ng this time (1, 047 out of 1, 848). The effects of increased temperature, 20

decreased moi sture, uri c aci d accumlati on, and predati on may have caused th e sminth urids to find oth er habitats wh ich would be ph ysi ologically less stressf ul. Th e Neeli dae are th e smallest of th e Collembola collected during th is study. Th eir absence from samples from January th rough

April may be due to th e fact th at th ey may have been below th e 2. 5cm collecti ng dep th . Neelidae wh ich are found deep in the litter layer may subsist on decomp osed leaf material. Anderson and Healey (1972) stated th at Orchesella flavescens (Entomobryidae) seemed to feed fairly deep in th e li tter layer and to include in its diet large quantiti es of parti ally humif ied leaf material. Smi nth uridae and Neelidae may be in di rect competition with one anoth er.

Both were found in samples during May. In June, th e

Sminthuri dae were absent while Neelidae were sti ll pres ent.

Both Sminth uridae and Neelidae were found in samples in

October through Nov ember. This may be due di rectly to leaf fall wh ich would produce more favorable habitats in wh ich to feed and reproduce. Th us, th ere would be less competiti on between th ese two families.

The greatest number of mi tes were found during October and Nov ember. The fewest were found during March and

April. Mite populations increased by more th an a factor of two from September to October (3, 170 indiv iduals to

7, 889 indiv iduals). Th e increase may be due to leaf fall 21

which may lead to increased microf lora activity. In fact,

Luxton (1981) say s that some phthiracarids (e. g. Phthiracarus anonymous) have organized their lif e cycles where the peak period of prelarval dep osition coincides with the peak period of litter inp ut onto which the prelarva are laid. Loots and Ry ke (1967) believed that abundance of

Cryp tostigmata is strongly correlated with rich organic matter. Butcher et al. (1971) stated that the abundance of oribatids may be related to organic matter as a source of food. Gist and Crossley (1975) gave a possible exp lanation for increases in microarthrop od populations as being due to increased microf lora activity commensurate with leaf fall. The increase in microf lora may lead to an increase in the fungivorous Collembola and Cryp tostigmata.

The average temperature during October, 1983, was 13. 9° C.

The total amount of precip itation for that month was

6. 39 inches. These two abiotic factors along with low pH readings from the moss samp les may have sp eeded up

the breakdown of leaf fall allowing for greater microf lora activ ity, thus resulting in an increase in the mite pop ulation.

The ef f ect of predation on mites during this study

is uncertain. Cloudsley -Thomp son (1958) rep orts that

cryp tostigmatid mites are not pref erred by predators such

as spiders, due to their strongly sclerotized exoskeleton 22

or distastefulness. Madge (1965) found that the numbers of predatory and cryptostigmatid mites showed the same trends in fluctuations and that the decrease of cryptostigmatid mites in spring and early summer was not caused by predation. Madge (1965) also found a marked increase in numbers of cryptostigmatid mites during July and August, possibly ow ing to high faunal and bacterial activity in decaying leaves and hence greater availability of food during that period. The population of Collembola also increased during the fall months but not to the extent that the mite population increased. Niijima (1971) stated that it is generally accepted that predation is one of the main factors which regulates populations of small animals. This may suggest that the Collembola (also fungivores) were preyed upon more heavily by predators

than were the mites. There may be several reasons for

this. Collembola lack the hard ex oskeleton found on the

cryptostigmatids thus it would take less energy for predators

to feed on Collembola than on mites. Also, Collembola

such as Hypogastruridae are just as large or larger than

the cryptostigmatid mites collected in this study and

thus could provide the same or greater amount of nutrient

material for predators. McBrayer et al. (1977) found

mites to be less mobile than the entomobryid Collembola.

However, when compared to other collembolan families, 23

the entomobryids are least restricted to any environment due to their mobility. The Hypogastruridae, for example, being "plump" and having a rather shortened furcula, could be assumed to be less mobile and just as easy to prey upon as cryptostigmatid mites, making them more suitable to predators.

Abiotic factors seemed to have less effect on the mite population as no positive correlations were found when comparing temperature (Table 4) and moisture (Table

5) to number of mites.

There was only one positive correlation found when comparing pH to the number of mites on a south-facing slope during the spring and summer months. These observations, however, do not conclusively demonstrate that mites are not affected by these abiotic factors.

Other invertebrate groups collected during this study are briefly discussed below:

Protura:

A total of sixty-five Protura were collected.

The greatest number of proturans taken from any one sample was seventeen on October 31, 1983, from a south-facing slope. The order Protura is considered to be a primitive one and are usually collected from moist soil, moss, leaf mold, under bark or rotting wood. 24

Diplura

One hundred thirty-five diplurans were collected.

The majority of diplurans collected belonged to the family

Japygidae (97) . The greatest number of dip lurans were collected from May to September. Diplurans are usually found under bark, stones, logs, in soil, and leaf litter .

Thysanura

Only one immature thysanuran was taken during the samp ling period on November 30, 1983, from a west-facing slope. Most species are found in leaf litter, under bark and stones, in debris and some may be found in buildings.

Orthopt era

Three Orthoptera were collected. One each from the families Gryllidae, Gryllacrididae, and Blatellidae.

Isopt era

Three worker termites were collected during August from north-facing slopes. They belonged to the family

Rhinotermidae. They are subterranean termites which nest in the ground and burrow into wood, usually fallen trees and stumps.

Psocoptera

Twelve Psocidae were collected. Nine were taken from July to September. Levings and Windsor (1985) found that the Psocoptera and Thysanoptera increased during the dry season and the size of the Psocoptera pop ulation 25

is related to the length of the dry season.

Thysanoptera

A total of two hundred twenty-three Thysanoptera were collected. Two hundred six were taken from a single sample on July 14, 1983, from a west-f acing slope. All

Thysanoptera collected belonged to the family

Phloeothripidae. This family may feed on plants, fungal spores or be predaceous. Durant and Fox (1966) found a large number of thrips in a hardwood forest in the South

Carolina Piedmont Region. Grimm (1958) stated that

Thysanoptera migrate into the litter in response to low humidity. The findings in this study seem to agree with this observation.

Hemiptera

Twenty Hemiptera belonging to four families (Coreidae,

Dipsocoridae, Tingidae, and Aradidae) were collected.

The Coreidae (leaf footed bugs) feed on plants or are predaceous. The Dipsocoridae (jumping ground bugs) are found in moist places usually under leaves. The Tingidae

(lace bugs) feed on foliage of trees and shrubs. The

Aradidae (f lat bugs) are usually found under decaying bark or woody fungi.

Homoptera

One hundred thirty-seven Homoptera were collected with one hundred twenty-nine belonging to the family 26

Coccidae. Other families include Aphididae, Cicadellidae, and Membracidae.

Coleoptera

Two hundred fifty Coleoptera were collected. The families with the greatest number of individuals found include the Elateridae (52), Staphylinidae (65) , and

Psclaphidae (31). All the Elateridae (click beetles) collected were larvae. The larvae (wire worms) feed on plant or animal materials and are found in rotten wood

or soil. The majority of the staphylinids collected were larvae. Most adults and larval staphylinids are predaceous

on other insects but some feed on decaying organic matter

and a few are parasitic. All pselaphids collected were

adults. The short winged mold beetles are said to feed

on fungi.

Lepidoptera

Twenty-four larval and pupal Lepidoptera were

collected. Eight belonged to the family Tortricidae.

Tortricids are common moths, many of which are called leaf

rollers.

Diptera

A total of two hundred sixty Diptera were collected.

The families with the greatest number of individuals

collected included: Cecidomyiidae (46) , Chironomidae (85), and Dolichopodidae (29). All but one cecidomyiid (gall 27

gnats) were in the larval form. The larvae may be parasitic, predaceous or phytophagous (gall producers). All the chironomids (midges) collected were larvae which are typically associated with very moist habitats. This is probably why only eight chironomid larvae were collected from June through September. Larval dolichopodids are found associated with wet soil, under bark, in decaying vegetation, and in water. Adults, of which two were collected, are predaceous on smaller insects. Forty-one fly larvae remain unidentified.

Hymenoptera

Nine hundred thirty-three Hymenoptera were collected of which five hundred sixty were Formicidae. Ants vary

in feeding habits with some being carnivorous and others herbivorous or scavengers. Other Hymenoptera collected

included a variety of tiny wasps. Twenty-six individuals

belonging to the family Encyrtidae were collected on

September 12, 1983, from a south-facing slope. This was

the only time that encyrtids were collected. Most encyrtid

larvae are parasites of scale insects and white flies.

Twelve Eurytomidae were collected. Eurytomid larvae feed

on plants or are parasitic on other insects; a few are

parasites when young and plant feeders when older. Nine

Scelionidae were collected; these wasps are parasitoids

of insect or spider eggs. 28

Araneae

One hundred five sp iders were collected. Linyp hiidae

(10), Hahniidae (32), and Thomisidae (35) were the most commonly collected families. The Linyp hiidae, (line weaving sp iders) construct a snare and take up a position underneath the snare to wait for prey. Hahniidae are small sp iders which build delicate webs usually near water or in moss.

Thomisidae (crab sp iders) are wanderers and secure their prey by stealth. Many individuals were observed with egg sacs. It would seem reasonable for Thomisidae to use the moss as a safe retreat to protect their young from predators. Young crab spiders af ter hatching may find it easier to find prey and, still, remain in a protected environment.

Oligochaeta

A total of one hundred seventy-eight oligochaetes were collected. All belonged to the genus Lumbricus.

They were young individuals which were present during the wetter, warmer months (March through June).

No individuals were collected from July through

Sep tember probably due to higher temp eratures and the

lack of moisture.

Nematoda

Twenty-two nematodes were collected. Ten Nematoda were collected in April. Nematodes are common inhabitants 29

of soil and litter but are generally too small to be collected with the techniques used in this study.

Chilopoda

A total of one hundred nineteen chilop ods were collected of which fifty-five were Brachygeophilus (Geop hilidae) and sixty-four Cryptops (Cryp top idae). Eighty-four centip edes were collected from March through July.

Centip edes are predaceous and feed on insects, sp iders, and other small invertebrates.

Pauropoda

One hundred ninety-two paurop ods were collected, of which, one hundred ninety-one belonged to the family

Polyp aurop odidae. The other individual collected belonged to the genus Eurypauropus (Euryp aurop odidae) . Fifty-one individuals were collected during Sep tember . Paurop ods occur under stones, in leaf litter and in similar microhabitats.

Diplopoda

Twelve dip lop ods were collected and all belonged to the genus Abacion (Lysiop etalidae). Most millip edes are scavengers and feed on decaying plant material.

Isopoda

Eight isop ods were collected. All were identified as Trachelipus rathkei (Porcellionidae). Isop ods are found in moist environments and are scavengers. 30

Chelonethida

Twenty-f ive pseudoscorpions were collected. They were absent from the samples from December through

April, and agai n in June. These predators are found under

bark, under stones, in moss, and in similar situations.

Phalangida

Two phalangids were collected; one in March and the

other May. Harvestmen feed on plant juices and dead in or live insects.

Symphyla

Eighty-one symphylans were found in samples from

March through November. All belonged to the family

Geophilelli dae. Symphylans occur in humus, soil, under

st ones, decaying wood, and other damp situations.

Gastropoda

Two Palli fera (Philomycidae)1 two Retinella (Zoniti dae) ,

and three undetermined slugs were collected. 31

CONCLUSIONS

Pr evious author s have stated that moss habitats do not contain the qualities necessar y for a fruitf ul existence.

Also, many have said that moss habitats ar e limited in their variety of or ganisms. My results suggest that moss habitats may ser ve as one of the mor e impor tant micr ohabitats for soil and non-soil or ganisms. For many soil or ganisms, the br yophytes play a ver y important role in their lif e cycles. The br yop hytes pr ov ide a habitat which buffers them from the extr emes of a mor e exp osed envir onment.

Dur ing dr y periods, some br yophytes ser ve as an "oasis"; containing mor e moistur e than the ar eas sur r ounding them.

The moss envir onment also supp lies protection from pr edator s and a site for aggr egation of the sp ecies for repr oduction.

It prov ides a place to lay eggs, a supply of food and war mer temper atur es dur ing the colder months. The var iety of inver tebr ates found with the moss/soil inter face was gr eater than that noted in other studies.

The intent of this inv estigation was to sur vey the fauna and to consider the ef f ects of abiotic/biotic factor s on the inver tebr ate population thr oughout the year . The findings from this study indicate that Collembola ar e mor e sensi tive to the changes in the envi r onment than ar e the Acar ina. This is probably due to behavior al and 32

anatomical differences. The Collembola, lacking a highly sclerotized exoskeleton, are more susceptible to env ironmental factors (relative humidity, temperature, etc. ) than are mites. This factor may also make Collembola more suitable than the acarines as prey. Such abiotic and biotic factors may cause Collembola populations to fluctuate more than those of Acarina. The Acarina may be more indep endent of these abiotic and biotic factors and theref ore exhibit more stable pop ulations.

Moss/soil invertebrate relationships seem to be in need of further study. Only a few pap ers pertaining to this area of investigation have been found and there are numerous avenues of research. For example, 41 fly larva collected during this study could not be identified, indicating a need to correlate immature stages with adults.

Many invertebrate life cycles are not known or are incomp letely known. agree with other invistigators I who believe there is a need for more eff icient methods of extraction; for considering predator-p rey relationship s and for studying horizontal and vertical migrations, aggregation, habitat, partitioning, etc. 33

ACKNOWLE DGEMENTS

There are a number of individuals who helped me throughout this study and to whom am very gratef ul. I would like to acknowledge Dr. Kipp Kruse for his advice I in biostatistics and the use of a computer. Dr. William

Weiler is thanked for the use of the water quality lab and his input pertaining to pH testing. Dr. John Ebinger identif ied the flora. Dr. Garland Riegel identif ied various

Hymenoptera. Dr. George Godf rey identif ied lepidopteran larvae. I would also like to thank my committee members for all of their help; Dr. Michael Goodrich for the identif ication of various Coleoptera and collembolans;

Dr. Bill Ridgeway for his guidance and editorial assistance;

Dr. Steven Whitley for identif ication of the Oligochaetes; and Dr. Charles Arzeni for giving me the idea for this paper and getting me started. Speci al thanks are due to Dr. Richard Funk, who identif ied a variety of invertebrate groups and who provided guidance throughout this study. 34

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Average Temperature and Precipitation

Recorded Throughout the Study T T 30 35 LEGEND

-��!4'<:. I H Temperature

25 + I � � I 30 Precipitation

- 0(_) I I � � � + 25 - --...... 20 L E -� ...... u :::J • 0L 20 c: .... -+-'0 CL 15 -+-'0 � E ..... ·- C.:> I-- 0... ia. 1 5 u I I 0L a_ 10 I � I I I ·���� � � m + 1 0

5 5

rrl' 0 I 'd4I" 0 Nov Jan Mar May Jul Sep Nov Dec Feb Apr Jun Aug Oct Month FIGURE 2

Compar is on of Total Indi viduals to

Total Indiv idual s Without Hypogastr ur idae 16000 LEGEND

14400 • Total Individuals

12800 • W / out Hypogastruridae

112 00

. Cf) 0 :=i 9600 -0 N .> -0 c: 8000 .... - �Ill<

0 � . 6400

4800

3200

1600

0 I ""1"" •••"1"'••••·�� I Dec Fe b Apr Jun Aug Oct Jan M ar M ay Jul Sep �--J ov Month FIGURE 3

Compar ison of Total Collembola Populat ion to Total Mite Populat ion on a Monthly Ba si s Dec Feb Apr Jun Aug Oct

Jan Mar May Jul Sep Nov MONTH

_. FIGURE 4

Comparison of Oppioidea to Oribatuloidea

on a Monthly Basis # of Individuals . N N lN VJ � __.) __.) f Ul 0 01 0 U 0 Ul 0 0 0 0 0 0 0 0 - 0 0 0 0 0 0 0 0 0 0

0 (1) (_ 0 0 :::J 'l (1)

� ()

0 l )> ""O .., �� QO z'< (_ � c I ::J (_ c )> c ({) (/) (I) u 0 0 ..-+ z 0 <

r r1l C) I f1l 0 0 z ..., -0 0 CJ -0-· 0 ,-+ 0 c Q. (1) 0 0 Q. (l) 0

FIGURE 4 FIGURE 5

Compar ison of Isotomidae to Hypogastruridae

on a Monthly Basis 12000 LEGEND

110 00 • lsoto midae

10000 Hypogastruridae

9000

8000 . Cf) 0 :=:> 7000 "U 11'1 "> -0 -c 6000 ...... 4-- . ca.� 0 5000 �

4000

3000

2000

1000

0 Dec Fe b Apr Jun Aug Oct Jan Ma r May Jul Sep Nov MON TH FIG URE 6

Compari son of Smi nthuridae to

on a Month ly Basis Neelidae 120 LEGEN D

1 1 Smint huridae 0 •

100 • Neelidae

90

80. (/) 0 ::J 70 "' -0 "> � "'O � c 60 Ai. ....__

0 50 ===**=::

40

30

20

1 0

0 Apr Dec Feb Jun Aug Oct Jan Mar May Jul Sep Nov MONTH FIG URE 7

Compar ison of Entomobr yidae to

Onychiuridae on a Monthly Basi s 500 LEGEND

450 • Entomobryidae

Onychuridae 400

350

(./)

0:::J 300

-0 ,... :> -0 c 250 .... - �r-. 0 � 200

1 50

1 00

50

0 I K

Jan Mar May Jul Sep Nov MONTH FI GURE 8

Total Number of Immature Insects

Collected Per Month # of Immature Insects

N 0J Ul ()) 0 0 0 0 0 0

0 fll 0

(_ )> z 'l fll OJ � )> :::lJ )> -0 �� 0 )> -f z�cz % (_ c r }> c () (/) fll -0 0 0 � z 0 <

FIGURE 8 FIGURE 9

Comparison of Percent Moistu re

Content of L. Glaucum on North-,

South- and West-Facing Slopes % Moisture content

N u .p.. (J1 0) 0 0 0 0 0 0 0

0 ro 0 (_ 0 :J ,., ro CY s: 0 -, )> u -, o�s: � :J (_ rt c :::y :J (_ c

)> c tO fJ) ro u 0 () r+ z 0 <

r f11 C) �II f11 � ()) z z (1) 0 0 0 (/) c -, r+ r+ r+ :r :::y

FIGURE 9 Table 1. Orders of inver'teb,r a"tes associa'te.d w1"th Leucobryum glaLt cum.

Order of Individuals Order o f Individuals * *

Pro"tura 65 Aranea 105 Thysanura Chelonee "thida 2 5 1 ... Oip lura 135 Pha l angida ..:. Col lembo la 2499 1 Acarina 38 197 Or"thop"tera Scolopendromorpna 3 63 Isop"tera Geophilomorpha 3 55 Pscop'tera 12 Chordeumida 12 Thysanop"tera 223 He'terogna"tha 192 Hemi p"tera 20 Cephalos"tigma"ta 81 Homop"tera 137 I sopoda 8 Col eop"tera 250 S"ty l omma'tophora 6 Lepidop"tera 24 Oxyuro1dea 22 Oip"tera 260 Opi s'thopora 178 Hymenop"tera 933 Tab-l e 2 Sea na l average 'tempe ra'tur•s w/ 1n - in'tesorface S u 't and We( ocs"t ) faforcin gair sl, opes . ITIOll ana mos s/soi l on Nor'th , o h

A1 r Sou'th Wes't Average Nor"th

II.Iin"ter 2.04 2.:6 1•16 .82 l S r i ng 11.55 11.62 10.70 11.29 p Summe r 25 .94 26 .04 24 .99 25 .66 Fal l 16.24 16.34 16.24 16.27

II.I/in Mo ss

i n e r 4.08 4.72 3.82 4 2 1 ll.l "t , Spr inQ 13.48 14.30 1 3.71 13.34 Summe r 24 .34 25 .15 24 29 24 .59 . Fa ll 14.86 4. 5 15.32 1 8 15.01

Mos s/Soi l in'ter'face

W1n'ter 5.24 5,41 5.29 5,31 Spr ing 14.43 14.06 13.64 14.24 Summe r 23 . 79 24 . 23 .44 23 .78 1: Fal 1 4 . 14.95 1 4.78 1 52 14.86 Table 3. Pearson Produc"t o n"t Cor rela"tion Anal s is be"tween and M me y pH organisms . - * of

Orga nism Slope Season r F'

Sp Su N s w 1.\1 F

Co l lembo la x x x x x - 0. 1231 x x x 0.0258 .9.7644106 x x x x - 0. 1071 . 23 x x x x x 0 2260 .479903 x x x 0 ..1035 x x x x 0.0667 .8870572 x x x x x 0.6549 . 01 71 x x x 0.5927 .2. 3768 * x x x x 0.4724 .2088 x x x o.8656 .0227 x x x 0. 1522 905 * x x x -0.0573 .9. 7 208 x x x x x x 0. 1849 x x x x x x x 0.1 198 ,5.3553595 Acarina x x x x x 0.0626 . 509 x x x -0. 4270 .48 294 x x x x 0.2781 .4842 x x x x x -0. 1075 .746 1 x x x -0. 8833 0160 x x x x -0.0857 . 332 * x x x x x 0. 1713 .68037 x x x 0.2668 .6359 x x x x 0.2267 .2860 x x x 0.5389 .2967 . x x x -0.4119 . 4482 x x x -0. 3820 485 x x x x x x 0.2169 . 8938 x x x x x x -0. 2144 .5. 5937

r = Pearson o n Cor rela"t1on F' roduc"t M me 't Coeffec1en"t = Two-sided S1gn1 f1c an"t Leve l p x = Ino1ca"tes pr on slope :: esence ano season( si I o1 a e s * n c "t S1gn1f1cance 4. Pearson Product' Moment'· Cor relat"ion Ana lysis be't'ween Tempera ure Tabla 't' (01 and of organisms . I

Organism Slope Season r p

Sp Su N s w w F

Inver'tebra'tes x x x x x x x -0.2302 .4818 Acar ina x x x x x x x . 79 15 o.oa19 Col embola x x x x x x x -0.3190 .3215

x x x 0 . 9 1 1 6 .0078 x x x -0.2977 .5949 * x x x 0.257 1 .6488 x x x x x .19 59 .7310 -o Acar 1na x x x 0.6435 .1857 x x x -0.6226 .2066 x x x 0.263 1 .640 7 Oppi odea x x x 0.6870 .1446 x x x -0. 7657 .0803 x x x 0.6134 .2160 Or ibu'tuloidea x x x 0.4207 .4372 x x x o.5146 .3244 x x x -0.0548 .9243 Iso'tomidae x x x x x 0.8854 .0153 * En't'omobry1dae x x x x x 0.3942 .4704 x x x x x o.7766 .0725 f'uYchiuridea x x x x x -0.2767 .6227 Sm1ejpogn't'ushurtruridae1dae x x x x x -o .1856 .7450

r = Pearson Product' Moment' Correla't'ion Coef fecien't' = Two-sided Significant' Leve l p x = Indica't'es presence on slop e and season = Indica't'es Signi ficance * Table Pearson Product Momen't- Correla't1on Analysis be'tween l Mois'ture s . Con'ten't and of organi sms . #

Organism Slope -Season r p

Sp Su N s Ill w F

Imma 'ture Insec'ts x x x x x 0. 98 .0093 x x 69 * Inver'tebra'tes x x x x -0.1392 .6743 x x x 0.2324 .5620 x x x x x 0.5232 .0815 x x x 0.2432 .5433 x x x x 0 1 462 .6588 x x x x 0.. 1 141 .7790 x x x 0.9516 .0014 x x 0.7803 .0700 * x x x x 0 3248 .5595 x x x x x o.. 5283 .0228 x x x x x 0.0653 .7074 * x x x x 0.1044 .6078 x x x x x x 0.3078 .1191 x Col lembo la x x x x x 0.5222 . 0 2 x x x x x x x 0.1113 .783752 x x x x 0.5561 x x x 0.5410 .1. 2942245 x x x 0.9 65 .0067 x x x x 0.89061 .0005 * x x x x x 0.5489 .0643 * x x x 0.6666 .1634 x x x x 0.6478 .0588 x x x x x 0.5812 .0463 * x x x x x x 0.6502 * .000 1 * En'tomoory1dae x x x x x -0.0497 .8814 x x x x 0.057 1 .8639 x x x x x 0.0781 x x x x x x x -0.0037 .8145 x x x x x 0.20 .9832 Onycbiuridae x x x x x O.Z3C?14l .5402 .4644 x x x x x 0.3878 . x x x x x x x 0 5 .12193 246 .2 6 1

r = Pearson P duc't o Cor rela't1on ro M men't Coeffecien't = Two-sided Sign1 f1c an't Leve l p x = Indica'tes o n s looe ano season< s> presence = Indica'tes Signi ficance * Tab le 5.

Orga nism l pe Seas n r S o o p

Sp N s "" "" Su F x x x x x 0.2088 .5250 Hypogastruridae x x x x x 0.5405 .0696 x x x x x 0.594 1 .0402 x x x x x x x 0.3074 .0680 * Iso'tomi dae x x x x x -0.0823 .8046 x x x x x 0.0103 .9752 x x x x x 0.344 1 .2818 x x x x x x x 0.0839 .6291 Acar1na x x x x x -0. 1225 .7120 x x x x x -o. 1894 .5652 x x x -0.4013 .4614 x x x x o.o74B .8543 x x x -0.2643 .6392 x x x x -0. 0734 .857 1 x x x -0. 2254 .6912 x x x x 0.0443 .9134 x x x x x x -0.0408 .8413 Opp101dea x x x x x 0. 1040 .7543 x x x x x -0.0343 .9181 x x x x x 0.0177 .9577 x x x x x x x -0.0086 .9605 Or iba'tuloidea x x x x x 0. 1279 .6996 x x x x x -0. 1355 .6824 x x x x x o.0484 .8844 x x x x x x x -0.0243 .8889

r = Pearson Proouc't Momen't Cor rela't1on Coeffecien't = Two-s1oed S1gnific an't Leve l p x = lndica'tes presence on slope ano season = lnd1ca'tes Signific ance * Table 6. Invertebrate predators a3sociated with LeueobrY\..Dll glaucl.Dll .

f'D.ITII TAXA 12-82 1-83 2-83 3-83 4-83 5-83 6-83 "7-83 8-83 9-83 10-83 11-83

Pselanhidae 6 5 6 1 1 3 5 2 1

Stanhvl inidae 5 2 11 10 7 4 6 4 1 4 6 5

Fonnicidae 3 3 28 2 8 86 79 137 55 141 25 13 8

Araneae 5 3 1 10 17 15 6 13 8 7 13 12

E1 mnrlidae 35 4 22 1 28 11 48 105 7 5 7

Bdellidae 8 16 4 3 9 29 22 238 144 171 146 144

Cun.axidae 2 4 8 3 1 28 10 20 10 18 4

Rhagidiidae 45 7 33 14 6 42 46 74 108 27 79 52

Ch ilonnchi 2 21 19 20 8 16 i 10 9 6

Chelonethida 1 5 7 7 3 2 Table Invertebrates associated with the glaucum and underlying soil. 7. L.

12-4-82 12-17-82 1 -4-83 1-17-83 2 -8- 8 3 2-26-83 3-6-83 3-27-8 3 DATE : SLOPE : w Apterygota N s !6M I I rl..Ll.JL.ffll..Ll.JLll..!!-l..Ll_w_ll-1Ll..Ll_w_ll-1Ll..Ll_W_ll-1Ll..Ll_w_ll-1Ll-Lll Protura = =1=1=11=1=1=11=1=1=11=1=1=11=1 =1=1r=1 =1_211=1=1= Thysanura =-2.= ·--· ---11--•--· ---11-- • --·---·- ·--· ---11--1--1---11--1--1 --- 11·-- ·--· ---11-- · -- · -- Oiplura - Carnpodeidae 2 s 2 ___ _ 1=1=11=1=1=11=1=1=1t::1=1 =11=1=1=11=1=1=11:_ 1=1=11_ 1 =1= Japygidae Collembola -- · ---1 1 -- • --·---11-- 1 --1---·- ·--· --- 11-- • --·---1 1 -- • -- · --- 11·-- ·--·---11-- • -- · -- -- 2 6 6 'lsotomidae 36 34 22 - - 3 11 3 136 4 21l3T· 3 11� 1 30 - Entomobryidae__ 26 56 1;4- - - 1 --zo -rs1 6-5 --1 36 -37 -n-21 -1 8 To-25 59 62 -12 6 -1 4 3 n- 11 9 1 -2 - - - - - lO - 1 --3 5 9 -- 3 Onychiurid� - 2 1 1- -6 12 -23 1 3 -21 - 4 -10 -45 -36 -26 28 -36 - 2 - - - 11 12 - 7 - 1 -ro 9 HypogastrMrida� 20 - 1 - 8 1 12 -50 --13 -22 -70 -33 -13 1003 -r2 329 -16 - 3 - -- -4- - - - - 9 - Sminthuridae 1 8- 1 --61--1 ------40 - - -43 42 14 -15 1 4 --· __ - 2 ______1 ______9 _ --_ _ Neelidae 1 l Ptl!rygota ------_- _ - . ------Exopterygota__ . Orthoptera Gryllacrictidae_ . Gryllidae

Blatellidae --- · lsoptera · --- 11--1--1 --- .. --·--·--- 1 1 --•--·---n-- •--·--- 11--•--·-- Rhinotermidae --·---11--•--·---11--•-- Psco tera ___ - =1=1= Psoc fdae ·=1=11=1=1-r11=1=1=11=1=1-r11=1=1=11=1=1 1 11=1=1=11 ·-- --·---11--•--·---u--·--· ---n-- • --·--- u-- · -- ·--- n-- •--·--- u --1--1--- 11-- •-- Thysanoptera__ Phlaeothripidae_ =1=1=11=1=1=11.J..1=1=11=1=1=11=1 1 1=11=1=1=11=1 =1=11=1=1= Hemiptera Aradidae Coreidae =1=11=1=1=11=1=1 111=1=1=11=1 = 1=11=1=1=11=1=1=11=1=1= __ Dipsocoridae --- 11-- •--• --- u-- 1 -- 1 --- Tingidae -- •--•---n-- •--•---n-- • -- • ---u -- 1 --1---11-- 1 --1--- 11--1--1 Acal;t2ta

Homoptera --- n • --- 11 1--•--- u -- • --•--- -- • --•--- 11-- •--• --- 11-- • -- • --- 11--1--1---11-- • --1 -- 1-- --

Aphididae 11 1 --- 11-- 1--• ---n --•--· --- Cicadellidae . -- · --- 11-- 1 --1---11-- •--·---11--1--1--- 11-- 1 -- 1 --- -- 1 -- Membracidae =1 Coccidae --- Cercopidae Endopterygota__ I II I I 11 I I II I l_J_ll I I II 1-'I�I I I I II I I Coleoptera Polyphaga -. I II I I II I I II I I II I I II I I II I I II I I II' Continued Table 7.

6-13- 83 4 - 12-83 4-30-83 5-8-83 5-31-83 7-14-83 7-31-83 DATE: 6-21-83 w N w ------I w --w - - -- SLOPE: w ------s s N s w N s w N s w N s N _L -1L _L N

------�pter ygota - rotura __ 3 1 2 2 5 ------hysanura - ,_l-- ,-1!. - l

------lu -- - - - ip ra -- 1 __ _ _ -- 1 3 2 3 2 l ampodeidae 2 1 0 __ 1 3 - - ---i -1 .J 3 2 1 s apygollemboidaela -1 3 3 --2 10 2 -.!l _l -1 _l -1. -1. 113 1 03 85 318 68 100 147 109 7 142 147 99 20 28 sotomidae - 1 23 1 02 141 71 1 56 243 5 1 3 56 37 62 3 7 39 1 37 ...11.30 10 ntomobryidae -1Sl 2 so 54 25 16 37 v 4 17 63 276 Ti17 18 29 nychiuridae _)].30 _];..2 15 58 '16ls 14 � 60 Ts 53 79 44 88 86 T6 -m 48 T3 2s 22 _D _s 173 � 71 114 14 _ H �pogas trur1daL_ 448 ls 9s 26 22 l 34 31 s !.!.Q3 _M s� 11� 5T8 194 -1 � � -1.2. __!! ___] rninthuridae - -- - 5 30 1 2 7 1 __ N eelidae _j_ Ii� - 1 31 1 14 l 2 5 1 l ,2- --1. - - -1. ------.-1 - p ------..- ry ota ------e - . - - - t g __ ------E ------Kopterygota ------

------0 - - -- - rthoptera ------

------G - - -- - ryllacrididae ------_ -

------G ------_j_- - -- ryllidae - 1 ------B ------latellidae ------

------so te a ------p r ------R :iinotermid·ae------_ - _ p ------scoptera ------l 1 . o c d -1. ae 1 s i ------· T ------hysanoptera ------3 p hlaeothripidae �

- - -- · ------H ------emiptera ------

radidae '

A -- · ------oreidae ------c - - -- - ipsocoridae_ 2 1 D ------i idae ------

T ------ng ------calypta ------

l -- A 2 -r ------omoptera ------

------ii ------1- - -- phididae ------A - icade llidae c -- - -r emb acidae H -- -r 6 occridae . c 1 7 l -r 15 . 2 � -· � -- - - -1. -- --1. ------ercopidae ------c - -- rdopterygota E. _ - e ------ol p a ------eo t - - c - -- r ------ol p aga ------p y h ------Table 7. Continued

DATE : 8-12-83 8-30-83 9-12-83 9-30 -83 10-16-83 10-31-83 11- 16-83 11-30-83 TOTALS s w N s w N s w N s w w s w N ------SLOPE: N w w -- - TAXA _§_ JL _§_ -1L ------.JL 2-

-- -- Apterygota ------Protura - - - - 4 2 _3 65

• ___! - _!_.-1 -- ....!2. -- _!_ -- Thysanura -- -1. -1. ------..l: ------Diplura ------__ __ ------1 - -- - - Campode idae_· - - - __ 1 - _ _l _l l l 1 2 37 Japygidae - 2 - 4 ------1 - - 98 _l _l Collembola -1 -1 __!_ - _1. -1 Isotomidae 128 32 1 19 6 41 66 2 5 51 104 108 3 40 9 3 75 4 83 96 7 - 74 52 Ento1110bryidae 7 ill. 18 --1.Jl26 �1 28 35 -1.U35 5 _12. 24 86 54 1---1 1 28 /;2 Si T4 29 29 Onychiuridae 28 .JL � 3 13 � -12. _!_ 4 21 �2100 39 3s 53 9 14 48 13 20 2 23 17 Hypoga strur 61 2 171 8 __..!!_8 69 10 3i2 33 39 5 1 ..1..!±. � 6 � 15 2 ts 12 32 7 33 53 42 __57 18 __ 15214 Sminthuridae� - _7 _!l - 11 5 4 1 _]] 21 -- � ...li _!.§ 1 l 4 ' J __ _!_ -1 -1. -1. l � _ Neelidae - _5 2 4 l - __!. 28 Pterygota __,!-- ,-1-- 1-1.�-- _l ------1-1 - -- - Exopterygota ------Orthoptera - ..,.--

__ ------GryllacrididatL,_ -- -- 1 - ! ------Gryllidae ------_l - - - -- ! ------1. -- Blatellidae ------1. - -- l ------. -- - Isoptera ------Rh inotermidae ------) Pscoptera -1. 2 ,__!- - - 1 1 ------1 I2 Psocidae ------Thysanoptera - -- - -1 ------Phlaeothripi�- - - Hemiptera _z. --1. -1. -2.. _2. _L -1.. � Aradidae ,__!.---1. -1.... 1 -- - -- Coreidae _ - - - -- 7 - 2_ Dipsocoridae __!. __...!. ------1.. --1.. - - - zi 1 Tingidae -- - -- Acalypta 4 ----a-

------Homoptera ------Aphididae ------l- - Cicadellidae ------_1_ Hembracidae ------Coccidae --r - - 4 _L_ - J._ � � � _ _ _a. __.&_ ...!L =_ .ll �==J_==___ _ -2.... =_ Cercopidae --1 ..ll _ --.- J_ _..a._ Endopterygota ------Coleoptera ------Polyphaga ------Tab l e Cont inued 7. 12-4-82 1 2-17-82 1 -4-83 1-17-83 2-8 -83 2-26 - 83 3-6 - 83 3 - 27- 8 3 DATE : l SLOPE : 11.JLI _w _ll.JLI ...LI_ w_ll.JLl...Ll _w _ll.JLI 2-l _w _ll.JLI j __ll I __ll l l __ ii ...L w .JL ...Lj w -1!.. -L w ll.JLI 2-l L TAXA · - Anth i c i dae -- -- - C nth i-d -· · -- ii --1- -- a ar ae ------r - - ,------� =1===== 1 = =1=T======------__ __ Ela ter ida ee --_ __ --______-- __- __ - __ - __ _ �__ _ � _z__ _l_ _l_ � -1.. ______l______!._ _ _1._ _1._ ___ ....L Endomychidae ______

Heteroceridae ------Histeridae ------= =- - = - - = = = = = _ ======Lamp;,1ridae __

= = _ a = ______L thridiidae ______l______Lycidae ______

______N i t idulidae ______...L ..2- ______z.______L ...l._ __ _ Ostomi dae ______1 -1. acrida.,_ ------1 - -- - - Pha ------_ -- -- ______--______-- ______-- Pse laphidae-.,.- __ - - - - - .Jt______4 ______Pt i lodac tylidae ______

______S carabae idae ______

______L ______S coly.tidae ______

______dm nid __ __ Sc y ae ae _l taphylin idae______S -1L -'-- -S- .Jt_ ..2... ______l______� ______g_ _L _L______Tenebri on idae_

Adephaga _L ______

______Car abidae ______

______L__ _L ______Lep i dopter a ______

______Coleophoridae _ I I _ ------111 ------______-- ______--_ __ -- _ ___ r a --______-- -- __ -- _-- -- _ _ _ ome t i d e ___ ------Ge ______--1..______Heliodin i dae______L . ______No u ae ______c t id _ . Her:nin i inae -- . Tort ric id a ======;--- . = = = =______l_ _ Diptera ______L _i . Nematocera = Sca tops idae ======. ======__ __ Bibionidae . = = _ Cec i do myiidae _ 1 _J_ .L -2... l _.2.__ ..J_ -2.. _J. ______= _ _ = = __ _J_ = = = __ __ _1______= _.JL _6_ ..L-1. __ M etop i l idae______yc h -l. ____l ______C ______hiromomidae T 2 ------5.._ -- _ _.L ______.!.. .L __ ..L ______..Ji. ______-- c od da ______-- -11....L. Psy h i e :1 Tab l e Continued 7. DATE : f 4-12 -83 4 -30- 83 5-8-83 5 -31 -83 6 - 1 3-83 6-21-83 7-14-83 7 - 3 1 - 8 3 sLOPE : 11..Lf-Ll.JLfl..Lf-Ll..!Lfl.JLlll.JLH..Ll-Ll.JLHJLril-Llf.JL(J....f _w _ft.JLf J....f_w_ff.JLrJ....l.JL TAXA ae 1 �icid Cantharida---e 11------111--- 0 1 = _.l.== = _L ==--1.. ======11

1 �i� _ �1:; o _ � _ --- lc� T � ------_L 11------_ ----__ __ _ ----- __ - - __ E later idaee______-1- ...i_ _L ______l______l_ _J.______J._ ...L Endomychidae __ J_

Heter oceridae ------H isteridae ------

' ______11--- Lampyridae __ __- --__ -- ______- --____ ------__ --__ __ -- -__ -__ -- __ __ -- --- _ __--_ -- -- _l______-· - __ La thr idiidae ll·--- ______Lycidae ______11 _ _

______Nitidulidae ______..L 11 _l_ -1.._ __ _ _ i ______Os tom dae __ Phalacridae -1.. _L _l_ _1_ 11 - -1 ------______Pse laph idae _ _-- --__ --_ __ -- ____- __ -______� - - -- 11--- __ � J_ ....!______!______J_ _Ji_ _L.L _ Pti lodactylidae______

J... ______S carab�e idae ______1 ll·--- _ _L .L ______S colyt idae ______11 _Ji______-r- ______1 ______Scydmaen idae _____ ------11·--- ______� ______S t aphylin idae 1 ______-- T 2 11 Tenebri n ae_ � J_ _J_ _!_ _l_ ....L +.- _L o id 1 � Adephaga ------= = ___ 111 ______a bi a ______:=-:-_ C ra d e = =__ . ---n---- Lepidoptera ______---ll·--- ______Co leo phoridae______111 _J._ , m r ----ll •-•---11 • • --- 11 • • --- 11 • • ---n • • ---1 1--1--1---11--1 --1---11--1--1---11;---- o d ------Ge et i ae -- He l iodin idae------1 11 Noctuidae ___11 __ ------Hermin i inae ------======---ll--- Tortricidae-- = _ = = = --ll---_ _J._ _ = = = ______= _ __= __ __ = _ __ .J._ D i ptera ______...z.. 11 -1 -1 _z_ _l_ ....1.... .J.______l..______l_ o __ Nemat cera _ _l_ --11--- ______Scatop idae______s 11 ______11 ___ B ibionidae ______l______Cec i domyi idae______L --·ll--- -1..______.-:- --1.. ______L Hycetophi l idae_ __ __ -·ll--- ______Chiromomidae ______J_ 11 _l__ _Jt______-1.. ___ --1.. _L ______l______Psychod i dae _ _-1.. ..L ..L ___ _ ,, Tabl e 7. Continued - - I 8�12-83 8 -30 -83 10 16-8 3 10-31-83 11-16-83 11-30 83 TOTALS DATE : 9-12-83 9-30-83 s w N s w N s w - - - SLOPE : - JL ...L JL ...L .JL JL ...L .JL JL ...L ....L. JL ...L .JL .lL ------TAXA 1 - - - - - icid 1e ------Airt'h ------5 Canthari :fae ...L - hrys me e ------C o L ida ...L nidae-- ..i. _l_ ...L -1. -1. ...L ....1. ------.....L -- --- l2 t.;urculio ..L _Jl --2. _L _l_ ..L 13 rid aee_ ------Elate -2._ ...L ..i. _L _i_ _J_ 52 _J_- _J_- - -- ..J..._ - - ..L. - - - - -1.- d - -- ch - - -- nd m l _ --3.. -...1.. --- _L E o y ae -- l ------l ------Heteroce ------_ - ridae ------...L_ -- � Histerid ae ------1 ------_1._ -- - -- ..L.- Lampyrid,, ae ------___ --- -- th i i ------La r d .dae 1 Lyc idae ------1 ------1 ------2 Nitiduli ae _ .,- -y- i� Ostomida - ...L.. _l._ lQ4 Pha lacri ..L - ...L - - - ae___,_ Pselaphi ae _ 1 -- _i_ -- 1 1 --- .....L ------..L 1 31 - - - - - ...!... - ...L - - _l_ -1.. Pti lodac ylidae_ - - - - ..L - - - -- � ------__ ------_L - - Scarabae dae 2 e ------J_ ...... L ------Scolytida ------l ------dae_ ------c n - -- -- S dmae ------y a S aphy l i idae_ ------_l_ -- ...L_ -- -- t ...L_ ..L ...L_ .L 6S - _1._ - _l._ -1.. - -L ....1. Tene ri idae - - - _ .....L- - b o -- -- � ------ph ga --- -- ....1. ------Ade a ------e ------Carabida ------...L_ 2 -1.. - -1.. - - -- - ..L ------Lepidoptera ------leopho ri e_ - - - - - Co da ------1 ------.. ------Geometri lae l He liod in __ ------1 ------idae 1 1 4 a - Noctuida ------� ------Herminilna e ------1 - 1 - - 2 :fae ------l' o rtrici 1 4 � � D ipter r r T _J_ 1 1 1 _.L 1 a T 2 .1_ ..L T - emato e ra - - ) - N c - -- -- · ------Sca ops dae ------t i ------) ------Bibionidae ------Cec idomy iid e_ T ------1 ------..L __L a ..J_ .L. 464 Hycetoph ilidae_ - - 1 4 4 10 idae ------1 -- -- _ 1 1 -- -- Chiromom _2 T 1 ..L ....L .L ....L as ------Psychodi dae -,- - 1 ------Table Continued 7. DATE : 12-4-82 1 2-17-82 1-4-83 1-17 -83 2-8-83 2 - 2 6-8 3 3-6-83 3-27-8 3 SLOPE : J..!!.. _§_ _w_ ..!!.. _§_ _w_ _§_ _w_ ..!!...... L _w_ ..!!.. _w_ II ..!!.. _§_ _L..!!...... L ..!Ll....Ll-Lll..!LI....L I .JL

------Sc iar dae ------� i - IL� ------1-1 - - Ti idae -- - --1... pu l ------1.. ------Brachycera ....1...- -- - -1... -...L Dolichopodidae_ =1=1 11=1 1 1 11 _ - - I I I 2 Emp ididae . � -1.. _j_ l

Rhag ionidae_ ------

S t,ratiomyidae ______L _____ - - Therevi ae ------d _ - - - - _ - 11-11 Drosoph ilidae - =1==-11-1 1 11-1 1- _ _ = ------Cyclorrhapha_ - - _ ------Huscidae - 1 Phoridae = - 1-1 l ll_J_ -- - - Hymenoptera ------1 - - _ 1 1 _ 1 1 1 - _

Symphyta ------Siricidae ------Ap�cr i ta - - - = =1 =1=11=1=1 l - - 11 ae - - - -- 1= Braconid -- - - =1 ·11=1=1=11=1 Ceraphronidae _ = Cynip i dae = -1-

------Diaphriidae ____ - -1.. - Eulophidae ------1 1 - -1...L =11 11 omidae _ 11 11 Eury 1 t 1 11= _ ======1=- - .J.. - -1. _£_ Mutillidae ------

· _ _ Platygasteridae _ --- _ __ _ _L ------Pteromalidae ------_ _ _J_ ------Scelionidae ------..L Encyrtidae -- - _ _ - -- - -

Eupelmidae ------_ _ - -

dae - Formici -- Fonnicinae = = = = _1._ -2... Myrmic i nae______....1... _ __ ------1

Araneae ------1=11=1=1 ·11 .. I 1-1.11 I I II I l_Lll I� 1-=t Lab i dognatha__ -

- -- -- Hypochi loidea � - - - -

__ Neoc ribe llatae - - --

Dictynidae ------

Lathys 1 11 1 4 ___ -L 1 1 1 -1-1 - Entelogynae 11 1=1=11 l - II l..LI II l=l=ll=l--1:1= = Table 7. Continued

DATE : 4 -12-83 4-30 -83 5 -8-83 5-31-83 6-1 3-83 6-21-8 3 7-14 -83 7-31- 8 3

SLOP : I __ E ______11.JLI ...LI W ll.JL I ...LI W ll.JLI ...Ll W ll.JLl .....L l W ll.JLLLl W ll.J:L12-l W ll.J:LI ..LI W ll.J:LI ..LI� � __ __ S c i r id e ______a a _L _l_ -1.. _L ..L ..L 1 TipBraculhyceidaea = =1 ======r = __: = =· �-= da ______Dolichopod i e_ __ _1_ --2. _L _L -- ...L - - .!!__--- _L ..L - - p d d e ------Em i i a --- - - ______ida ______Rhagion e--'-- ______� _ d ______a ______S t r t omy e ______a i --i-- i - - The vida ------re e ------� - -1 _ _ _ ida ______Dro�op i l e__ ___ h � l ______yc a ______C lorrh pha______Huscidae i _L �� = 1 ==__ = = == _ =�==== ______=====______== ___ _ == a __ ,__ Hymenop te r ______

___ ymphy a ______S t ______Si icidae r ------Apocrita ------11--1--1-- Br c n dae 11 1 1 11 1 1 11 1 1 --11-- 1--1--11--1--1-- o ----11 --- 1 1 -·11 -- 1 1 ------a i ------______r h ida ______Ce ap ron e______--

______Cynipidae ______J_ __ _ _ a ______D ______i r __ ph iidae _L a _ __ __ u o ______....L______l p e _L____ E hid _L _____ E ______urytomidae ______L ______i d ______ut l e ___ a l _L _ i _ M ______Platygasteridae______1__ - o da - - - - - t r m l ------P e i e ------c na -- -- e o id ------S li ae ------______E ______cyrt id e ______-- n a __ __ d ______e ae ______Eup lmi ___ --- o rm ic dae - - F ormicii nae _l_ _2' _..J.6. ..14 = _so_ -1.. -1... -35 ..22. _J_ -22.-2l1._10_� = _ __...2B...... _ _ ___a__']_=�______yrm n __ -52.. .12... ic e ______H i a 20 2 2_ i 2 _l._ _l ------Araneae -- -Z------r ------=

__ ======-- Labidognatha = = = __= __= ______= = = = = __ =__ = = = _ __= ______-- Hypochiloidea = _ _ __

Ne ______-- crib __ ll ae ____ o e at __ __ ------

c ______-- - - tyn dae ______-- -- D i i __ -- ______Lathys ______

_j______e o a ______------Ent l gyn e - -- -- Table Continued 7.

2 DATE : I 8-12-83 8-30-8 3 9-1 -83 9-30-83 10-16-8 3 10-3 1-8 3 11-16-83 11-30-83 HTOTALS SLOPE : l.JL.ftJLfLI .JL.llJLI l .JL.HJLIi j __w__ 11.lLj Lj .JL.lfLI ...§_.JL.lf.JLT-s I i llJLIi I .JL.UJL1i1.JL TAXA - _1 _ -- - -

-- --- u ------p -,------��i:��::: - ----1------

_ -1______t8- ______11 Brachycera _

______Do l ichopodidae______.....L ______.L ______!_ _ _a_ Emp ididae ______

_.l_ __ _ _2______,_J_ Rhag ioni dae� _ ____

______--1- ______S ratio ae______t my id _

- --- _L Therevidae ------

_ _z______Drosol1hi lidae___ �______..L -1.... _ -1.Q_ C l ______L ______L __ _ -1.. __ yc orrhapha _ _ _ Huscidae _a._ Pho e ======-L r ida = = == _ _ __ _J______L ______Hymenoptera______

______Symphyta ______S i r icidae Apocrita _ ======_ _ = ______Braconidae ______

______2______Ceraphronidae______

__ __ _ C ynipidae ______L -5.______D i aphr iidae ______-6______j______Eulophidae _ __ __ -2.._ ------��� ------__ --� ______Huti llidae ______

______L ______. ______Platygaste dae ___ _ _ ri _

______l_ _s_ Pteromalidae______---... ______-2.__ �______s_ Scelionidae ______

20 _J_ __ J______- -- Encyrtidae L _ ____ -- --

______-- Eupelmidae _ _ _!_ i Formicidae ------11 -

_ Formicinae ______- - --Z... -5...... L _,5.______j____ ...li.. .1L _ _ __-11 __ �__ _ Hyrmicinae .JL _2._ _j_ _il _ _ i _L -1. -1. _s_ -1.... _l_ Araneae 1

= __ __ =� d ______======______Labi ognatha_ _L _======______

_ Hypochi loidea______

Neoc bel latae ------ri ------Dic yn idae ------t -======�

��l ______E ______nte ogynae _ _ _ _ _ Continued Table 7.

� - DATE : 1 2-4-82 12-17-82 1-4 -83 1 -17-83 2 - 8-83 2 - 26-83 3-6 -83 3-27 8 3 _ SLOPE • __l ' ...L!....!L. 11..!!...l...Li...Lll..!!...l...Ll...Lll..!!... l 1 __ 1l 1 1 __IL.�l l _ ll T AXA ..!LL.!.. w ..!L i w .JL i w i w .JLIi Ii

_ _l l Trionycha _ Mi i d _ _ _ _ -- - - - met ae - - _ I - 11l 1I l 1,I -1-·1-1·-1-1-11-1-1-11-1-1-11- -1-1-11-1-1-11-1-1- Age l enidae == = - - = = -= -= - - - = Theridiidae ======_l_ = ______=______= ______= _ __ Linyph i idae______= ____ _.1_ Hahn iidae ======_ = = _L======l �!:ridae ..2._ ------Lyc si a ------o d e ------Pirata ------Dio ch ny a ------Gn ho i ap s dae ------r------Club ionidae - -- Thom i sidae - ======-1... ======_ = _ _ _ _.1______= _ _ = l t c i dae ______Sa i _ 1 ..L Annelida _L

��������:t�ernstrls __i_ ..1- ...5- -1.. 6 _l_ _i_ 13 14 44 2 16 a ------�:m ��:: __ _ e ______-1 - __ __-- _ _ yu l 2 - ______Chi lopoda ______--____ _j______-__ ___ --_____1 ______omorpha -- 1 ------1- Geoph i ------1 - - - __ _ Brachygeophilus ______- ______4 ______J_ _J_ i_ __§_ - ______- - S co l opendromorph ______Cryptops -1.. ______l______..2_____ Pauropoda ______

______Eurypauropod idae ______

______

Eurypauropus ______.1_ Po p uropo i ae _ __ ly a d d ..:J_ ..J_ --2,. ______-2_ _J______-1.. D iplopoda ______

______Lysiopetal id!!t______

______Abac ion ______1 _1._ _ _L ___ _ I sopoda ______------

______----- ______---- On i scoi dea ______-

______------� ------______------Porce llionidae__ ___ --

______--

------e ______------CheTracheione�ihpus a ra�hk i --- t id -- --

------Pha l ng i a __ ------a d ______- ---

� ______Opilon s ______e � ___ - -- Tab le 7. Cont inued

4-30-83 6-13-83 6-21-83 7-14-83 7-31-83 DATE : 4 - 2 - 83 5-8-83 5-31-8 3 1 N s N s w N s w SLOPE : N s w w -- - - _ _ .1L _§_ - .JL _§_ I - - - TAY.A _w .1L _§_ _w w w .1L _§_ - -

------Tr i onycha ' - - - Mimetidae - -- - - I ------

_ - - - Age len idae

-- - Theridiidae------_1_------

__ __ ------dae ------Linyphii ------_L _L - - - - Hahni idae _l_ ------....L ------PRania - - - - - isauridae _j_ _l_ _l_-- - -- ....L- _2,_-- - ...L- - L- -...L -- - _l_------Lycosidae ------Pirata -1... ------Dionycha ------..L ------Gnaphosidae ------Clubionidae - - - _1_ ------;------T Thomisidae - -2- - - T 2 1 1 Salticida --.1_ - - e ------..1- _j_-- - ...1...------Annelida ------Oligochaeta _ - -- _ - - - - Lumbr i cus terresttis _ - - 8 _2 -r - - _2._ .lL _z_ _l_--- � -1.. -- -- - Nematoda - - _i_ - --2- - _1_ _s__ ...1. --1.. - - - - Oxyuridae -2 - - 2 T -,- -r 2 - -- - - Chilopoda - -- ...l..- ...l..------Geophi.lomo rpha ------,- - Brach eo hilus ,-- ig e 2 1 _ =--L -- - Seo romorpha ..1.. _,l..- _L_-- -1...- -1... _J_ - ..L --L - - -- lopend - Cryptops 2 l T 1 1 1!L -- P _J_-- _l_ ...1...... L------auropoda - -....J_ - - ...L- - - --L- ..L- --1_ ------Eurypauropodidae ------Eurypauro2u.s - - -- - Polypauropodi dae _l_ ,-- - 1 T 27 2 J 0- T J -r- 9 -- i. - -- Diplopoda -- -- _&_.- - -1._-- 1L- _J_- -L------.1------Lysiopetalid!!L_ ------Abacion - - - - 1 1 T -- - -- Isopoda - -- ...1.------On iscoidea ------_ _ _ __ Porcellionidae ------___- r -r------Trac elipus rathke i ------h - 2 r ______=r= 1_ --r Chelonethida ------r - -- - _ _ _ i_ ------Phalangida ____ ------Opi lones I --. ___ _ - - · 1 - -- I - - Table Continued 7.

DAT 3 3 E: 8-12-83 8-30 -83 9-12-8 9- 0 -83 10-1 6 -83 10-31- 83 11-16-83 11-30-83 TOTALS SLOPE: J L..L l_w_ll-1Ll..Ll.JL.Jl-1Ll..!..l...LH-1LI..LI w_ _ ll.JLf f _w _HJLI H-1Ll..L1.JLll-1L ..!.. ..!..IH-1L ...L I 2...l.JL TAXA ______------1'r i onycha _ ·-- _ --· ______--

______Mime tidae _ _- ______' T ...l._ -- -- 1 ------..,,------Age lenidae -- -- 2 -- ___ ------� ------The r i d iid a e - -- --

__- __ -___ -_ __L ______- --_ -- -- ______1 _ _L - L i p dae _ _ __ _ 1 y hi __ n i __ - -- 10 _L ------__ ------_g_-- ---_L llahniidae ------1 -- -- r -- � - - �-- �b - � Pisauridae 1 - l ------T- - T-- -- - ....L ------ll-2-

l.ycosidae _ Pirata ======Dionycha ======2 - ======-- -- Gnaphos idae ------1 2 b - Clu ion idae ------L------b - 2 1 Tho misidae ------. ------,------J.5 Salticidae 2 ------_J_ ------l ------0 ------Anneli a ------d ------== ------Oli o ------g chaeta ------___,.. - -- II-ti'S ------� rres _ Lumb cus te t ______i _ _ _ ri ______1 -- i 6 11 78 ------�� - - r - oxyuridae ---.... -- �------1 _ 11.?.L - - _ -- _____ - _____ -- _ __ --______- - __ hi _ -- --__ --_ --- ___,. - C lopoda __ --

e ______- -- . ____,. ------G ophilomorpha _ ------

_ - llS ______,. �______J"" 2______T__ -- Brachygeophi lus T T ______.L _l. ______------� Scolopendromorpha _ __l_

______Cryptopa -1. _L -1...... 1.. _..2. _L_ _l_ _l_ ...L _J_ -1- ______-- -- Pauropoda ______

_ _ __ Eurypauropod idae __ _. ______

______. ______Rurypauropus 1______Pol ypauropodidae 11 ______JL _s. ______i___ _ !.2i. Dip lo oda ______44 --1._ __£__ ___..l__ __..l______lL _L_j_....L_'.L J._ � ___ -- L i l ______- -- ys opeta id!!L______

- -- Abacion ______---.... ______------

- -- .1______- ··- Isopoda ______- - --

_ -- - -- ______-- -- On iscoidea ______Porcellionidae ------

_ -__- _ Trachelipus rathk = = = tl ======1"" = = ______= ______T ______1_ Chelonethida _

1 ______- -- a ______j_ ___ _L _!_ _.§______j_ _!_ -1_ __ -- P a g id _ __ -- ha l n -- - Op i lones ______------2 1 3 1-17 -83 2 -8-83 3-6- 83 3 -27 -83 DATE : 12-4-82 . 12-17-82 -4-8 2-26-83 _ _ __ SLOPE: N _§__ _w .1L _§__ _w -1L _§__ -L .1L _§__ -L .lL _§__ .JL .lL _§__ w .1L _§__ .JL .lL _§__ TA.XA Symphyla Geophilellidae_ ------� T Mo l lusca ------Gas tropoda ------Ph ilomyc idae� T l Pallifera ------Zonitidae ------· ------Retinella ------· ------SO _ ------AcarAstigmidaeata ( ) ------Par i sitiformes or-_ .-12. _6._ .JL -1L -2.. -1. .w -9:_ --1. _jJ:._ -1.. _1_ Honogynaspides (SO ------Zerconidae :._ - 124 2 - 2 -- 1 1 2 Laelapidae _ 26 � . l -i l -0-2 - -t T 20 T 2 10 � -2.. 206 4 20 1 --r- 1 2 4 _l_ Ologamasidae_ · _R -1.. ...2._ -1.. lQ_ 1 2- 2 ,- 4 Uropodidae - - -- __ 1 ..£.. -1- _s__i_ 2 Act inedida ..L - _L 2 Promatides (SO)_ ------Eupodina ------Bde llidae ------Cunaxidae .JL - -- _.!t. ..L -L _l.. .li... -1.. _1__ --'- ...L 2 Eupodidae ' ------�4 - -- -1.. ..--1... _A_ ..L _L_ _J,_ � --- -- ..L 4 18 Rhagidiidae_ _!_ 18 () _£_ - 1 --- 2 4 - _l_ - � JL 10 2 10 8 4 2 Ty

__ ------Stigmaeidae ------· ------1 ---· - -- -1._ Tet ranychoidea_ ..L -- -- · -- Tetranychidae_ ------Trombic ulidae - -- ...2... -r- 1 + � Pachyna thoidea_ 1 1 -r- 2 -r ,- - ,- T _ _ __ ------======i.= Lordal ychidae _ . ------1 - 30 - 2 -83 -83 5 - 8-83 5-31-83 6-13-83 6-21-83 7-14-83 - 3 1 - 8 3 DATE: N 4 s w N 4 s w N N 7 s w - w ------SLOPE: -- -- _§__ ...lL .JL _§__ .JL _§__ w .1L 2- --w N s --w TA:XA

Symphyla ------Geophilellidae__ -1.. _L _2._ __.!! --1. � - - - __J_ - Mol lusca __.&_ -1.. i _J,_ __1._ _J_ � - - - - - _s_ ------Gastropoda ------Phi lomycidae -1.. ------� ------Pallif ------_1_ - - - - - era --. ------Zonitidae - _l ------Re tine lla ------1 ------. ------__ ------As t igmata Acaridae (SO) ::: 1 4 Parisitiformes -1. ---1.. _j --3 __&.,. 2 -1 .J. J... -1

__is o)_ �onogynaspides ------Zerconidae ------1 - - Laelapidae __ � - � _J_ i __,.&_ --3 . __L � ___5_ __!! -1. --1.. J --1. -1... _z_ __J :>logamasi dae__ � -1Q. 4 .ll. "-4 J -r -1- 2 1 2 Uropodidae __ ---1. 1 _j_ 2 � 1 1 2 1 _!_ 1 lctinedida (SO)_ .i. ..1_ Promatides ------Eupodina ------Bde llidae _1__ __1_ 46 18 � _u 64 i -1. ::unaxidae -1. - --2. _1. --2.. - - - _l._ _1t.. ...L - _J_ -2.. _J_ 2 - - _4_ Eupodidae 1 -- ---1.. __!.. ___!_ 1 2 -1. JJ_ -5.. --1.. 2 2 6 1 1 Rha idiidae____ 16 10 -1. g - 10 20 --i --1. T -2. "4"" T 18 8 ..!2.. .-2 14 10 14 .12.. _ll_ ryde idae -'- -5... _l!_ _l.. _l.. ---1 - - - _l_ - - - - - fleterostigmae_ - - 3 _1. ------J_- _1t..- __J_ ..1.. _s_ � -1 ------Eleutherengonina ------rarsonemo idea ------rarsonemidae ------

--__ ------a - - - Pygmephoroide ------_ ------'kutacaridae -- ..2... -- -2 --2,_ -1.. -- -- -3- -- -2- -4- __ - - Raphignathae ------_/!..------� - - _l.. ------Raphignathoidea_ ------__ - Stigmae idae ------1 -- __ ------2.. -2.. -- - - - retranychoidea ------� -- -- retranychidae ------_

-1. --1. --2- ..J._ _2_ rrombiculidae - - - _j_------1... _lL _£_ - _i_ -5. _L _z.. � _ _2.. _2-. -1- -L __ -1.. --2.. --l. -- _2.. - - - Pachynatho idea - - - _J,_ - - _2. - - - __..z ------__ ------______------___ Lo rdalychi dae -- _i _1.. DATE : 8-12-83 8-30 - 83 9-12-83 9-30-83 10-16-83 10-3 1-83 11-16-83 11-30-83 TOTALS _w_ _w_ _w_ _w_ _w_ SLOPE z N ..§._ .Ji_ ..§._ .Ji_ ..§._ .Ji_ ..§._ L ..L N s w .Ji_ ..L w .Ji_ ..L TAXA Symphyla ------Geophilellidae_ ------1 1 _2_ 1 81 M l J.._ .....s.. ol usca - - -.....s.. --1... - _J_ - - -1L ---1...... 1_- ....1__ ....L------

Gastropoda ------.J..

__ Phi lomyc idae ------11 Pallif -- 2 ------era Zonitidae ------1 Re tinella ------

------( = -- As ti ata S0)_ ------� 1 1 -r Acaridae 1 - -- - 2 Parisitiformes 2 1 -,- --r 2 2 - -- -1 - -- -2 11 215 (S_O)_ Monogynaspides.!------Zerconidae 4 1 --_ 4

__ � 1 ...£_ _J_ 2 J26 Laelapidae ._6..- -_6.. -1L l--2- -....1_ - J...- - _it_------2- ---1 - - -

__ Ologamas idae ------196 - _J_ Uropodidae ------Actin�dida ( SO)_ -5.. -2... _l_ _1._ � 1 _L _L -1.. -1 - 2 ,- 2 05 Promaddes _A. ....L _l_ _l_ ....1.. -1. -1.. T T 55

Eupod ina - - - - - Bdellidae ------..J.1... 1 0 14 18 12 14 20 JWL .2.2.. -.31. � 1 Cunaxidae 1-Jf --1... _J_ ...2.9 _J.9 __:n_ _JQ �2 _S1 __!2 _ _L _1_ 1 _ll_ ..J.1 .2.�108 Eupod i dae _..z...4 -1.. _..z..._ _l_ -2...21 10 _l_4 _j_2 463 -t _J_8 18 _!.._ ...£_ _2 _2._ _J__ _J_

__ 6 Rhagidiidae ll.. � 1 1 _2_ 10 .fil_ 22 17 T � 2 8 -a Tydeidae ...li .ll.. 16 _:]_ 4 ..L2 I4 i[ -2- lb lJ 2 2 1 95 Heterosti mae � r T +- T + -11m. g - - _J_ _s_ 2 L _1_ - - - - - _ ------

Eleutheren onina -- - - g ------

Tarsonemo idea ------

Tarsonemidae ------Pygmephoroidea_ ------

- Scutacaridae ------4 Raph __ _Ji. _A_ ..J_ __l_ ...2... -6.._ ..Ji...... 1_ ..2-. i i _z_ ...L ..L n ignathae ------_:n_

Raphignathoi dea_ ------

__ igmae idae ------· - ...L _Jl_ St 2 4 Tetranychoidea_ ..2-. � _J__ _J_ _l_ ..L Tet ranychidae_ _2_ _2 _ 1 __:&__ 4 Tromb i culidae_ ---1.. -1.. _2_ -1.. __1f_ __L __L 1 _J_ Pachynathoidea_ -1.. -1.. _l_ _£_ _! -1. -Lii _l___Jj______Lorda lychidae ______-- J_ _£_ ------Table Continued 7.

DATE : 12-4-82 JZ-17 -82 1-- 4-8- 3- 1 - 1 7 -83 2-8-83 2 - 26-83 3 - 6 - 83 3 - 2 7-8 3 LOP L S E: w w_ _w w tJLI __ _ JL 2- _ i1 IL1L 2- w JL 2- .JL JL 2- w .1L _§__ .1L _§__ .1L _§__ TAXA - ______------Pachygnathidae ------§ _ - Sphaerolichida _ ...... 1... -6_ - - - --2.... -1... -2.. ..2.!t_ .L ...L -1- _lt.. __l_ ....10 J..... ( SO) ______------1 -..L. ------Or ibatida -....!L --1L --1 -

_ _ - - - Hacropylides ______------____ - - - - Epi lohmannidae_ __ _ ------_ _ _ __ - - - Nothroidea _ - - ..L- - ---1� - -1... - ,_, __,, _ _,s. -S- _g_ ..11.6. -5- ...... 1... � C::am is.i.a -Z.L _a_ _J,_ _.2Q - - - Perlohmannioidea 11...... 1... ------�- - - - - 3- 3- J...... J______--1. ------Lohmannioidea _ - � ------

- _J.:l - - Orlbotrltia lL_ _lt.. - -'- _2__ _J_..JL -.1- Oppioidea .192. _.L _j,_ _J_ ...L. ..ll ..20. Ji_ JL � .2.Q6 .l.Q9 ... 11- 122 32- Or ibutuloidea_ � i.JL i -59 ....91. _j,_ _g_ -Z2. ..2fJ; � ...160.. .li. _ill .J5S 81. 22-111-- W 1*.58... 232'-1J � -9.6 2.:L ..:z_ L2Q. w. � � ...31.5_ .32... JlQ._ 1.5.1._ _ __ -2.6_ li.1LlQOZ31. Trhypochthonius ....!!.� _i.. , J Table Continued 7.

4-12-83 4-30-83 5-8-83 5-31-83 6-1 3-83 6-21-83 7-14-83 7-31-83 DATE : SL0PE : .1L ..L .1L ..L _ w _w .1L ..L .JL .JL ..L .JL .JL ..L w ..1L ..L w .JL -s --w -N -s -w TAXA Pachygnathidae__ Sphaerolichidae_ ""2 - 8 - - - - _ 6 - - - - 4 -'- ...L Orlbatida {SO � 4 ....L -1... _j_ _l_ _!_ _2 .i _L _J_ .Ji_ .JL

Hacropylides____ ) -- Ep ilohmann idae ------__ ------__ 1 1_ Nothroidea · 1 -1 _1. -· - -1. - - -- _jt_ - -1. -2.. - ..2.. -- -:/- ...... 6 _u._ -1.. ...2..... -- Camisia � Perlohmannioidea. 1 6 ...D -1J8 �'- -SI- � 2 2 l 1 Lohmannioidea_ ..&.. _J_ _Jf ------_J, ------Oribotritia Oppioidea ..Jt. -.Jf:. -1.... .11. -1._.J._-5..-i _9,_ _a ....9... -2- -1- ....s... --2...... 1-.-3--3--.JL...---1.�-10.-2.CI..-2- Ot:ibutuloidea_ � ..1tL .12.0- iU JJ::L ..JL . ..JQ:i .liB ..2AtL w 122 ...1'92 l4- .5S-...sa...205 328 l.09- 288. I ..lir.: � � 622..i _, • �" ..sz ; '-'! 1 -:· 9Z- � -61_ 1.QQ � 1.86. � -12+ � _ 11aa. 1t ll$ ______! ..1.0; ..69-- I� I� 2.sa... .251_ .21.6 l.39-. l:ch;JPQchthon11111_ _i_ ...L - - ..L - - - -'- -2.. - - - .,2___ Table Continued 7.

-83 9 -12 -83 9-30-83 10-16-8 3 11-16-83 11-30-83 8-12-83 8 - 30 nTOTALS DATE : I 10-31-83 lL ..L ..L __ N ..L. SU>PE : w .JL __§_ w .JL ..L w .JL ..L _w_ .JL w ..L w .JL ..L w .JL

TAXA -· ------Pachy nathidae ------g _ __ 8 Sphaerolichidae__ - -- - - � -'- -1- ...... !&... -1,_ -1,_ ( O)_ _ -2...- _l_------12!.. S j_ -6...- -1 Or ibatida _L -_a_ ...L ...2- - - -_z.. - -l.. .L- _L- -2..- -- __ ------__ - Hacropylides ------

------Epiloh81annidae_ _ 1 1 2 -r 2 _L_2_ _ u_ Nothroidea T 2 1 rr /;" T 2 T --1 -1. -1.. - 2)1 - - - 06 - - - __l - - _i_- 1 Camisia ---- 14 1 ..L 41l _l_ - -- - i2a- ...l6.- -2a �- M------2.... l nn d - JL- - - _J2.. _2_ _2_ Pe o i ea _i_ -----� - - _ � r hma oi -- _L _ . ...l.. ------1. ----- _L Lohmannioidea ,._ 2 _2_ ����--�-ii����� �l������� _J_ OppOr ibotritioidea ia 2 1 T s2a 121 16� 62 66 O r ibutuloidea 121.A_ 1�..fil. 1�&1���..!J2... m226 llL� �lli��� � .:ru §1.Q_ .Mmo�� mm� 312.�� 36L � �� _ ..6il.�� JS � -- � 217__ _ - i- - _ Trhypochthoniu. L L-�-�1� -- �------2L