... VVvvv-VV'V‘" v—v- n-v-v vv'-----—-n-.c.~l'— ~¢gf.o.ri vsanvv‘OVpOOOIOOOOOOoo‘Q?OWMoclat-(ooogq'ovo¥-«mr.~.”moghu\hqy.,.’nww . , _. u ‘ ‘:v‘ . . ~ I '

''''''''' """"

1 O - Q-I O

.0 .t. . IA, _ D ‘ ' I" 7-. _

53'" THE SOIL ARTHRGPODS or A woonan LOT' :_ ,5: ; IN SOUTHWESTERN MICHIGAN - ~ f ;*;;j_]}_;v;:3;;§§;‘=3

Thesis 'foI The Degree 0‘ M 5. J; __ ' MICHIGAN STATE UNIVERSITY \ - _ _. . . . f Judithuir Aitkenw ' ‘ - - ;:'. ' "9 I f "19.6.4." " . "L' -g“ V 4 ‘ vv‘ 400¢3N.¢..fi.‘.‘ _'1 " "°‘ '4- ‘fl” ' '}:

rd

N

I

'

' 0..."

Ilflmfllfilllmflmfltflmflfllfifl IL 31293 00093 5993

LIBRARY Michigan State University

ABSTRACT

THE SOIL OF A WOODED LOT

IN SOUTHWESTERN MICHIGAN

by Judith Muir Aitken

Arthropods found in a wooded lot in Kalamazoo County,

Michigan revealed little variation among the taxa found in the various micro-habitats.

Polvxenus §E,, a of millepede not recorded for the area, was found in the area studied.

The amount of precipitation, position on the hill, drainage, temperature, pH, and phosphorous content of the soil contributed to the importance of the distribution of the . Plant and associations play an impor- tant part in the balanced soil community. THE SOIL ARTHROPODS OF A WOODED LOT

IN SOUTHWESTERN MICHIGAN

BY

Judith Muir Aitken

A THESIS

Submitted to Michigan State University in partial fulfillment of the requirements for the degree of-

MASTER OF SCIENCE

Department of Zoology

1964 ACKNOWLEDGMENTS

I am greatly indebted to Dr. T. Wayne Porter for his encouragement, direction, and constructive criticism. His initial suggestions and observations in Invertebrate

Zoology were the inception of my interest in the field.

Sincere thanks are also expressed to Dr. Ivan F.

Schneider, John G. Eaton, Charles S. Scarborough, and

Richard J. Snider who have all contributed to the many phases of this research.

Helpful suggestions and opinions were given by Drs.

Jane E. Smith, John A. King, Wilbert E. Wade, and Roland

L. Fischer. They were greatly appreciated.

Gratitude is also expressed to Mrs.IL P. Henderson for the services she rendered, and to Mr. G. Noordam who aided in many ways at W. K. Kellogg Biological Station.

ii TABLE OF CONTENTS

Page

ACKNOWLEDGMENTS ...... ii

INTRODUCTION ...... Ia

DESCRIPTION OF THE AREA . . . .

-b Location ...... In

Geological History and Surface Features a

Study Area ...... u:

METHODS AND MATERIALS . . . .'. . . as

Collections ...... ox

Preservation ...... ~4

Temperature Data . . . . .‘. . . (n

Chemical Data ...... (n

Identification and Study . . . . (n STUDY AREA AND SELECTION OF SAMPLES 0

ECOLOGICAL DISCUSSION ......

H

o Physical Factors ......

F‘

P4

Chemical Factors ...... Lu

P‘

Biological Associations . . . . .5

F4

Botanical ...... 9

IA Zoological ...... '. . .

Ia U1

SUMMARY ......

Lo

Id

LIST OF ARTHROPODS FOUND ...... 9 Io

BIBLIOGRAPHY ......

UI

w

iii LIST OF TABLES

Table Page

Distribution record for macroarthropods

collected in the sampling area . . . 20

List of Collembola ...... 30

List of Acarina families found . . . . 31

Number of organisms per sample . . . . 32

iv LIST OF FIGURES

Figure Page

1. Sampling plots and vegetation in the

wooded lot ...... 33

2. Position of lot ...... 35

3. Amount of rainfall . .i...... 37

4. Position on the hill ...... 39

5. Temperature ...... 41

6. Months sampled ...... 43

7. pH ...... 45

8. Available phosphorous ...... 47 INT RODUCTI ON

Arthropods are represented in every type of soil en- vironment. In most soils they are the most numerous of micro- and macrofauna present. In spite of their abundance in the soil, they have not been intensively studied as a group.

Published European studies have been more extensive than those in the United States. Among the more notable are those of Bornebusch (1930), Kevan (1955, 1962), and

Kuhnelt (1955, 1961). Bornebusch's work was not as thorough as that of the workers that followed him, but it stimulated a wave of studies of the soil fauna. Articles written in the past twenty years show the influence of Bornebusch's work and established the basis for subsequent ecological and taxonomic studies (Dowdy, 1947, and Lawrence, 1953). With a refinement of equipment and techniques, and, with the grow— ing interest in the subject, there tended to be a division of pure phylogenetic investigations into that of certain orders or classes. Studies of specific taxa appeared to become prevalent. Kuhnelt (1955) and Kevan (1962) have attempted to organize the workdone on the various orders and classes, but the vast expanse of information available has made it difficult to publish a complete synopsis that will adequately cover a study of the phylum. Progress in different geographical areas has also advanced at different rates. Recent ecological investigations by Kevan (1962), however, indicate reassemblage of the phylum and a more comparative analysis.

Investigation of soil arthropods in the United States has not been intensively pursued. Hairston and Byers (1959) acknowledged the great expanse of material that could be obtained from a study of the phylum, but in their research, they identified the arthropods only to order. Families, genera, and species were referred to by letter and number.

Engelmann (1961) listed only the Acarina or mites and again, only to suborder and lesser taxa were referred to by letter and number. Gasdorf and Goodnight (1963) identified only soil arachnids to genus.

Michigan is one of the few states in which investiga- tions have been made of soil arthropods. Hairston and Byer

(1959), and Engelmann (1961) studied in Michigan, however, these were made in southeastern Michigan, and the arthropods were not identified other than to order. Engelmann was concerned with an old field community. Hairston and Byer discovered that little correlation or agreement was found between populations in pastured and forested lands.

The present research is an attempt to investigate the arthropods of a wooded community in southwestern Michigan and to indicate the diversity of arthropods living within a small area. DESCRIPTION OF THE AREA

Location

The area investigated is part of Michigan State Univer- sity property on Gull Lake, Kalamazoo County, approximately twelve and one—half miles northeast of Kalamazoo, Michigan.

The area is on the northeastern side of the lake (T15, R9,

W56).

Geological History and Surface Features

The land in the vicinity of the study area is pitted outwash plain connected with the Kalamazoo Morainic system of the Lake Michigan and Saginaw Lobes of the Wisconsin glacial stage.

The basin occupied by Gull Lake suggests the persistence of a mass of stagnant ice along the course of the glacial drainage system during the deposition of outwash gravel.

This area is southwest of the morainic belt built by the

Saginaw lobe where knob and kettle topography is present.

The area investigated is in the outwash plain.

The drift of this morainic system is Bellefontaine sandy loam. Numerous lakes, swamps, and marshes are found surrounding the studied area. Little soil development has occurred, and sand and gravel are prominent immediately under the shallow organic layer.

4 Study7Area

The area studied is in a wooded site adjacent to the

W. K. Kellogg Biological Station of Michigan State Univer- sity. One side of the woodlot is bounded by a field and on the opposite side by a marsh. It was logged over approxi- mately 60 years ago and a more recently used logging road runs through the area. Sheep grazed the land in 1960.

The woodlot is located on a slope of approximately 30 degrees which runs the length of the area between the field and the marsh. The slope faces northeast, the direction of drainage. The study plot is 50 meters across the top, bot- tom, and along one side, the fourth side is 60 meters long, the greater length of this side accorded by the contours of the marsh at the bottom of the plot (see Figure 2). METHODS AND MATERIALS

Collections

Samples were taken randomly from the area. The plot was divided in 27 squares each 100 square meters by driving steel stakes into the ground. The stakes were painted white in order to facilitate their location. Using the above squares, a stratified random sampling procedure, each of the squares was divided into 100 smaller squares, one meter on a side and numbered consecutively from left to right when looking north. By randomly picking one of these numbers, a one square meter area inside each larger square was selected.

These one square meter areas were then divided into 16 equal parts, each 25 centimeters on a side. By randomly choosing one of these sixteen parts in each one square area, 27 sampling areas were obtained and distributed within the sample plot. Two subsequent samplings were taken as follows: a) the first, to the right or north side of the original; and b) directly below or to the east of the second sampling in each area (Figure 1).

Soil samples were collected to a depth of one inch plus leaf litter and debris, and were 25 centimeters square in area. These samples were carefully placed in plastic bags and carried to the laboratory; there, sorted by hand for

macroarthropods. If time prevented immediate sorting, then

they were refrigerated temporarily. This did not appear to

have a detrimental effect on the animals.

Four separate core samples were taken to a depth of six

inches. These were used in evaluation of the vertical

distribution of the organisms and not included in the com-

puted data.

The sorted refuse from the samples used in the researCh

was then placed in a modified Tullgren funnel so that the

effect of the heat and light would drive the microarthropods

down and out of the soil. The animals were then collected

in specimen jars filled with 70% alcohol.

Preservation

The jars of 70% alcohol were stored until identification

could be accomplished. The annelids and molluscs were then

removed and the arthropods sorted. Acarina and some of the

Collembola were mounted on slides in Hoyer's mounting medium.

Some of the pseudoscorpions and polyxenids were also mounted. The remaining Collembola were placed in vials with

90% alcohol and sent to the Entomology Museum at Midhigan

State University for species determination. Arthropods remaining were labeled with date, locality, and collector,

and placed in vials of 90% alcohol for future reference.

Temperature Data

A Weston Testing Thermometer was used to take air and

soil temperature readings at the study site. Further air

temperature readings and daily precipitation rates were

obtained from the United States Weather Bureau at Kellogg

Research Farm, approximately one and one—half miles away.

Chemical Data

Samples from each ten meter square area were taken back

to the Michigan State University campus in East Lansing,

and Dr. Ivan Schneider and members of the Soil Science

Department ran tests to determine the soil pH and the avail- able phosphorous (pounds per acre) in the soil.

Identification and Study .— Three kinds of microscopes were used to identify the :‘“'.‘ .‘ arthropods. They were as follows: a) Bausch and Lomb

v' 3‘! dissecting microscope with 3X, 6X, and 10X powers; b)

Bausch and Lomb compound microscope with 10X and 45X powers;

and c) A. 0. Spencer Phase Contrast microscope.

STUDY AREA AND SELECTION OF SAMPLES

Distribution of the collecting sites is shown in Fig-

ure 1. Three samples were taken from each ten square meter

area. The first sampling sites were randomly chosen within

‘ten meter square areas and the two subsequent samplings

were as follows: the second was to the right or north of

the original site and the third site was directly adjacent

and east of the second in each site. The sites sampled were

distributed through the woodlot so the two areas that were

closest were about four meters apart. Most of the areas

were at least six or seven meters apart. A total of 81

samples were taken from the studied area.

The distribution of predominant vegetation is indicated

in Figure l by symbol. Oak covers much of the area and is

mixed with sumac or hickory in the plot. However, a stand

of pine does occupy about one-fourth of the area, and the

litter in this portion is almost all pine needles. About

one—ninth of the area is inhabited by willow and this area

is bordering the marsh at the bottom of the hill.

The material for this study was collected during a

year's time starting October 21, 1961 and ending Octdber

21, 1962.

ECOLOGICAL DISCUSSION

On the whole, the populations within the

studied area do not appear to vary. The presence of some

species appeared to be consistent in almost every studied

'area and those taxa that were less abundant were scattered

in a random pattern. Eighty—eight families of arthropods

were discovered in the woodlot. These families came from

five classes or 21 orders of the phylum.

No one of the taxa was found in all the studied sites.

A few were found in more than 80% of the sampled sites.

These were Porcellio §E,, Neobisiidae, Clubionidae,

Geophilus £22: Julus £23: Parajulus £23: Compodeidae,

Staphylinidae, Nitidulidae, Curculionidae, Chalcidoidea,

and Formicidae. The species of Collembola were uniform

throughout the study area. The number of individuals of

Acarina were uniform throughout the study area.

A millepede genus, Polyhenus £23: not previously re-

ported for Midhigan, was collected during this investiga-

tion. Occurrences of this animal are recorded in Texas and

Georgia, and it has been found in southern Illinois.

10 11

Physical Factors

The amount of water in the soil is affected by such diverse factors as slope of surface, nature of organic constituents, soil texture and structure, and amount and type of precipitation. Soils with somewhat sandy surfaces, such as in the studied area, allow rapid penetration and such soils do not retain moisture as well as the heavier clay soils. Precipitation was measured for each seven day period preceding each sampling day. The precipitation dur— ing this period appeared to have little effect on the arthropods collected in this study. However, if there had been no precipitation for seven days previous to sampling, the number of organisms declined in the top inch of soil.

Core samples taken indicated a vertical migration of the animals to a depth of at least one inch. Most of the ani- mals were found in the top three and one-half inches of soil (Figure 3).

When there_was an increase in rainfall from .00 inches to .50 inches over a seven day period, there appeared to be a slight increase in the number of organisms in the upper layer. Previous work done by this author (unpublished), however, revealed that a precipitation of over three-fourths of an inch within a 24 hour period caused a decrease in the number of macroarthropods. 12

Samples at the top of the slope where there was little

litter and ground coverage contained fewer organisms than

those on or at the base of the slope. Those with larger

numbers of organisms tended to contain predominately animals

of the same taxa (Figure 4). This is illustrated by sam-

ples taken from A-l, D-2, D—1, and E-l. In A-l, of the 104

animals taken, 55 were Formicidae, and 15 were Thripidae.

Of the 28 animals in sample D—2, 12 were Porcellio s2, and

seven were Phalanqium sp, In D-l, of the 61 animals studied,

31 were Formicidae and 11 were Nitidulidae. Twenty—two

Thripidae and nine Formicidae were the greater portion of

the 42 macroarthropods found in sample E-l.

Although a few samples at the bottom of the hill indi-

cated the same trends, many of the samples showed a more

rounded distribution. Sample D-5 contained 36 individuals

of 15 taxa. The twenty individuals in sample C-3 were from

11 taxa. Sample D-3 had 21 individuals divided into 14 taxa.

Numerical variations in arthropods resulted from a dif-

ferential response to temperature. The optimum temperature

for the largest number of arthropods in a twenty-four hour

‘period was around 600 F. There was a marked decline at 450

F and another at 700 F. Within the range of 45° F to 700 F,

there appeared to be a fairly uniform distribution with a 13 slight tendency toward the maximum temperature. At tem- peratures ranging from 550 F to 600 F, fifteen samples were found to contain an average of 45 individuals. Thirty-four samples were found in temperatures ranging from 650 F to

700 F. These samples contained, on the average, 55 indi- viduals per sample (Figure 5).

Another factor affecting the number of arthropods was the mean monthly temperature. Samples taken during late

July and early August contained many more organisms than those of earlier or later months. The number tapered off in October and November, further declined during the winter months and then started to climb again in April (Figure 6).

Chemical Factors

Recorded values for soil pH lie between 2.2 and 9.6 inclusive, and values below 4.5 and above 8.5 are unusual

(Allee, Emerson, Park, Park, and Schmidt, 1949). The pH of soils tends to be a limiting factor for the number of indi- viduals and variety of taxa in an area.

Arthropods collected in the sampled area were in soils ranging in pH from 4.4 to 6.8. In most cases the thof the habitats in which the organisms were living was between 4.5 and 5.9, the average being 5.1. Organisms increased where 14 the pH was around 6.5, but this is due to a greater number of individuals within certain families rather than within the arthropods as a whole (Figure 7).

Phosphorous is essential in the separation of decom— posing protoplasm. It is resolved into phosphoric acid by soil bacteria and is stored in the soil in the form of phosphates of aluminum, calcium, iron, and magnesium. Such phosphates are essential in metabolism and making of proteins.

Arthropods were found in the area studied in soils containing from nine pounds phosphorous per acre to 120 pounds per acre.

Most organisms in the sampled area were found in soil con— taining from 40 pounds per acre to 80 pounds per acre. The soils containing between 40 pounds of phosphorous per acre and 55 pounds per acre contained the highest number of organisms (Figure 8).

Biological Associations

Botanical

The area studied had a variety of plants that are indi- cated in Figure 1. In the study area, oak and hickory formed the most prominent association, but it also contained both a pine and a willow association. The following trees and larger shrubs were most abundant: bitternut hickory, pignut 15

hickory, shagbark hickory, white oak, black oak, red pine,

American elm, weeping willow, flowering dogwood, sassafras,

basswood, hawthorn, buckthorn, black raspberry, sumac,

burning bush, wild honeysuckle, summer grape, and juniper.

'Where two or more species of plants formed thickets growing

close to the ground, they were recorded in Figure 1.

None of the arthropods found in the woodlot appeared to

be in association with only one type of plant. Although

Polyxenus s2, has been described by others as being asso—

ciated with pine, and was found during this study in pine

litter, several specimens were discovered in a willow asso-

ciation. Since keys for determining species of Polyxenus

are not published and specific determination has not been

made, statements as to the relationship of habitat appear

to not be a factor limiting Polyxenus to pine. Twenty-eight

animals of the genus were found in the willows, so apparently

it was not a chance association. The Polyxenus s2, occurring

in the pine sites occurred in more or less a random asso—

ciation with the vegetation.

Zoological

The most numerous of all microfauna in the soil are the

soil mites (Acarina) and (Collembola) which are 16 confined to the soil. The other arthropods designated as macrofauna migrate out of the soil into the litter, plants, and other ecological associations such as fields, and marsh areas, and are not those Kevan (1955) considers true soil arthropods because they spend most of their time in the layer of litter.

The distribution of the macroarthropods and Collembola appeared to show no consistent pattern in the sampled area.

For the most part, each taxa of macroarthropods was present at least once in 55% of the collected samples. Five classes and 21 orders were present in the study area. Eighty-eight families were identified (Table l).

The acarina also appeared to have no pattern to their distribution when referring to them by family as would be expected, but when examined by number of organisms present, oribatid mites were discovered to be more prevalent at the top of the hill where it was dry and sandy. Near the marsh at the bottom of the slope, the mesostigmatids were more numerous.

Other invertebrates associated with the arthropods were nematodes, earthworms (Lumbricidae), snails, and slugs. A rainfall of .50 inches brought these animals to the surface and into the leaf litter. After a rain, sample D-3 yielded

17

57 annelids, and five or six snails. Only 29 arthropods were found in conjunction with 46 annelids in sample A-5 and 22 of these arthropods were Acarina. In sample D-3, 38 arthropods occurred and 32 of these were Acarina.

In addition to the above, species of Polygyra §E,,

Anguispira §p,, and Succinea s2, also occurred. A para-

sitized Succinea gp, was located in one of the samples.

During a warm, humid day the Succinea s2, was in the leaf litter. When the snail was taken back to the laboratory, the tenacles appeared to pulsate and swell. They looked very much like two annelids burrowing in the soil. The animal was killed and upon dissection, the parasitic fluke redia Leocochloridium paradoxum, which resembles a small annelid when it pulsates, was discovered. This movement attracts birds and aids in transmission of the fluke.

The microenvironment is a close-knit community with the necessary components. Predator-prey relationships among the insects, spiders, Collembola and Acarina are evident.

Mesostigmatid mites such as Parasitidae, Veigaiaidae and

Macrochelidae are predators and.have chelicerae that are adapted to this purpose. Collembola, small insects, and oribatid mites such as Belbidae, Camisiidae, and Cara- bodidae serve as the prey and also as primary consumers l8

and scavengers in the community. The predators, in turn, become victims to the larger predators. Dead animals are processed by carrion feeders. The activity of all these

animals leads to the production of humus which aids the

development of plants which adsorb the inorganic nutrients

liberated by bacteria. The earthworms, in turn, mix the humus and inorganic soil elements.

Investigations made by others into soil arthropods are many, varied, and equally interesting. They include all phases of active research such as physiology, ecology, and genetics. Limitations in the scope of this paper prevent further elaboration, but I hope that the subject has been discussed sufficiently to assist the reader toward an understanding and appreciation of the field of soil ecology.

SUMMARY

Arthropods of a woodlot in Kalamazoo County, Michigan, were sampled. The distribution of the arthropods from

81 samples was determined and placed on a chart.

Acarina and Collembola were listed on separate tables.

Polyxenus £22: a genus of millipede not previously re- corded in Michigan, occurred in the woodlot.

The amount of precipitation, the position on the hill, drainage, and the temperature variation all appeared to have an effect on the number of arthropods.

A pH of 5.1 appeared to be an optimum for the soil arthropods and phosphorous is essential for them, with most organisms occurring in soil containing from 40 pounds of phosphorous per acre to 80 pounds per acre.

The plant associations within the studied area did not markedly affect the distribution of particular taxa.

Nematodes, earthworms, and gastropods were found in close association with the arthropods.

19 20

Table 1

List of the Taxa of Macroarthropods Found in the Sample Area

A + indicates the presence of the a animal in that particular sample

A-l A-2 A-3 A-4 A-S A-6

Porcellio sp. + + + + + + + Neobisiidae + + + + + + + + + + Liobunum sp. Phalanqium sp. Argiopidae + Attidae + + + + Clubionidae + + Lycosidae Gnaphosidae Thomisidae + Theriidae ,1 Salticidae ' Geophilus sp, + + + + + + + + +

Scolopendra s2, ' Lithobius £2, + + + Polyxenus s2: £2195 §E, + + + + + + + + + + + + + Parajulus gp, + + + + + + + + Polydesmus s2, + + Fontaria s2, Acerentomidae Campodeidae + + + + + + + + + Tetrigidae ' + Gryllidae Blattidae Psyllipsocidae + Phloethripidae . Thripidae + + + + Anthocoridae Miridae Reduviidae Nabidae + + Lygaeidae + Pentatomidae Cercopidae + + +

21

A—5

Cicadellidae Hemiptera Nymphs Homoptera Nymphs Cicindelidae Carabidae Ptiliidae Staphylinidae Elateridae Nitidulidae Murmidiidae Silphidae Cerambycidae Coccinellidae Anthicidae Mordellidae Bostrichidae Scarabaeidae Chrysomelidae

Curculionidae +

Scolytidae +

‘Coleoptera Larvae + Hydroptilidae Lepidoptera Larvae Noctuidae Diptera Larvae Drosophilidae Asilidae Chironomidae Diapriidae Pipunculidae Ichneumonidae Braconidae Chalcidoidea Formicidae + + + Hymenoptera Larvae

22

B~4

Porcellio g2,

Neobisiidae Liobunum g2, Phalanqium g3.

Argiopidae Attidae Clubionidae Lycosidae Gnaphosidae Thomisidae Theriidae Salticidae Geophilus s2,

Scolopendra s2, Lithobius s2,

Polyxenus g2,

Julus g2, Parajulus sp, Polydesmus sp.

Fontaria §p, Acerentomidae Campodeidae Tetrigidae Gryllidae Blattidae Psyllipsocidae Phloethripidae Thripidae Anthocoridae Miridae Reduviidae Nabidae Lygaeidae Pentatomidae Cercopidae Cicadellidae Hemiptera Nymphs Homoptera Nymphs Cicindelidae Carabidae Ptiliidae

23

B-4

Staphylinidae +

Elateridae +

Nitidulidae +

Murmidiidae + Silphidae Cerambycidae Coccinellidae Anthicidae Mordellidae Bostrichidae Scarabaeidae Chrysomelidae Curculionidae Scolytidae Coleoptera Larvae Hydroptilidae Lepidoptera Larvae Noctuidae Diptera Larvae Drosophilidae Asilidae Chironomidae Diapriidae Pipunculidae Ichneumonidae Braconidae Chalcidoidea + + + Formicidae + Hymenoptera Larvae

24

C-4

Porcellio g2,

Neobisiidae Liobunum sp. Phalanqium §E, Argiopidae Attidae Clubionidae Lycosidae Gnaphosidae Thomisidae Theriidae Salticidae

Geophilus s2, Scolopendra g2, Lithobius §E, Polyxenus g3.

Julus §p, Parajulus gp, Polydesmus sp.

Fontaria gp, Acerentomidae Campodeidae Tetrigidae Gryllidae Blattidae Psyllipsocidae Phloethripidae Thripidae Anthocoridae Miridae Reduviidae Nabidae Lygaeidae Pentatomidae Cercopidae Cicadellidae Hemiptera Nymphs Homoptera Nymphs Cicindelidae Carabidae

25

c-4

Ptiliidae Staphylinidae Elateridae Nitidulidae Murmidiidae Silphidae Cerambycidae Coccinellidae Anthicidae

~Mordellidae + Bostrichidae Scarabaeidae Chrysomelidae Curculionidae Scolytidae Coleoptera Larvae Hydroptilidae Lepidoptera Larvae Noctuidae Diptera Larvae Drosophilidae Asilidae Chironomidae Diapriidae Pipunculidae Ichneumonidae Braconidae Chalcidoidea + + + Formicidae + + Hymenoptera Larvae

26

D—4

Porcellio g9, Neobisiidae Liobunum sp, Phalanqium s2, Argiopidae Attidae Clubionidae Lycosidae Gnaphosidae Thomisidae Theriidae Salticidae Geophilus g9, Scolopendra sp.

Lithobius sp,

Polyxenus §p§ +

+ +

Julus s2, + + + Parajulus s2, + +

+

+ + Polydesmus s2, +

+ Fontaria s2, + Acerentomidae

Campodeidae + Tetrigidae Gryllidae Blattidae Psyllipsocidae Phloethripidae Thripidae Anthocoridae Miridae Reduviidae Nabidae Lygacidae Pentatomidae Cercopidae Cicadellidae Hemiptera Nymphs Homoptera Nymphs Cicindelidae I) ‘\ Carabidae

27

13—4

Ptiliidae Staphylinidae Elateridae Nitidulidae Murmidiidae Silphidae Cerambycidae Coccinellidae Anthicidae Mordellidae Bostrichidae

Scarabaeidae + Chrysomelidae

Curculionidae +

Scolytidae +

Coleoptera Larvae +

Hydroptilidae + Lepidoptera Larvae Noctuidae Diptera Larvae Drosophilidae Asilidae Chironomidae Diapriidae Pipunculidae Ichneumonidae Braconidae Chalcidoidea Formicidae + + + Hymenoptera Larvae

28

E—4

Porcellio sp, Neobisiidae Liobunum sp, Phalanqium §p,

Argiopidae Attidae Clubionidae + + + Lycosidae Gnaphosidae Thomisidae Theriidae Salticidae Geophilus §p, Scolopendra g2, Lithobius ER: Polyxenus s2,

Julus EB: +

Parajulus s2, +

Polydesmus sp. Fontaria sp, Acerentomidae Campodeidae Tetrigidae Gryllidae Blattidae Psyllipsocidae Phloethripidae Thripidae .+ + Anthocoridae Miridae Reduviidae Nabidae Lygaeidae Pentatomidae Cercopidae Cicadellidae Hemiptera Nymphs Homoptera Nymphs Cicindelidae Carabidae

29

E-4

Ptiliidae Staphylinidae Elateridae Nitidulidae Murmidiidae Silphidae Cerambycidae Coccinellidae Anthicidae Mordellidae Bostrichidae Scarabaeidae Chrysomelidae Curculionidae Scolytidae Coleoptera Larvae + + + Hydroptilidae Lepidoptera Larvae Noctuidae Diptera Larvae Drosophilidae Asilidae Chironomidae Diapriidae Pipunculidae Ichneumonidae Braconidae Chalcidoidea + + + Formicidae + + Hymenoptera Larvae

30

Table 2

List of Collembola Collected

Order Collembola Suborder Arthropleona Family Poduridae Hypoqastrura s2, Neanura muscorum Templeton

Family Onychiuridae Onychiurus subtenius Folsom Family Isotomidae ‘ Folsomia fimetaria Linnaeus

Proisotoma s2,

Isotoma trispinata MacGillivray

Isotoma viridis Bourlet Isotoma albella Packard Family Mydontidae Mydonius multifasciata Tullberg Tomocerus vulqaris Tullberg Tomocerus flavescens Tullberg Pseudosinella pettersini Borner Pseudosinella pettersini f. rolfsi Mills Lepidocyrtus curvecollis f. achromatis Maynard

Lepidocyrtus cyaneus Tullberg Lepidocvrtus lanuginosus f. albicans Reuter Orchesella ainslici Folsom Suborder Family_Sminthuridae s9, Family Neelidae Neelus minutus Folsom 31

Table 3

List of Acarina Found

Order Acarina Suborder Mesostigmata Parasitidae Veigaiaidae Macrochelidae Laelaptidae Zerconidae Uropodidae Circocyllibanidae

Suborder Trombidiformes Scutacaridae Pyemotidae Rhagidiidae Bdellidae Trombidiidae Trombiculidae

Suborder.Acaridiae Saproglyphidae

Suborder Oribatei Carabodidae Ephilophnammidae Galunmidae Camisiidae Eremaeidae Belbidae Liacaridae Ceratozitidae Phthiracaridae 32

Table 4 List of the Sampling Plots, and the Number of Macroarthropods in Each Sample

I ' II III

A-l 26 104 10 A-2 57 73 58 A-3 3o ‘ 23 21 A-4 48 16 29 A-S 19 7 22 A-6 19 15 19

B-l 22 23 11 B-2 12 41 11 B-3 26 96 35 B—4 10 w 17 37 8-5 30 81 24 B-6 14 26 197 c-1 15 42 16 c-2 4o 7 8 c~3 20 17 50 c-4 18 23 566 c-5 22 42 52

D-l 57 19 11 D—2 12 23 50 D-3 6 37 21 D-4 25 16 52 D-5 54 36 88

E-l 33 50 44 E—2 10 57 10 E-3 16 26 21 E-4 111 26 56 E-5 7 115 41 33

Figure 1.

Sampling plots and vegetation in the wooded lot.

The symbols represent the following vegetation:

Oak

Sumac

Hickory

Pine

Juniper

actual studied area is indicated in red.

35

Figure 2.

Position of the wooded lot with respect to the field and marsh that border it. The contours of the land and the position of the logging road are also indicated. I Bela / ’ ’ ’-/,

/\/r / z ’ / A 8 , , ’ c d E / / I/ Thicket ///

\ /"/ 1 0% \A‘{\‘\

/’ \A / “-\ / . I / “LC ke‘t 1 '\ 3 \ 5 \\ \\ \7 [FL /I

/‘ 4 I d? Hm

I” ‘1 %V

/ Rea / ‘\ ’\ \o x’Buming \ 5 5 L0 // BUSH / ,/ “flicked: / \‘\ / /

=’- 10 meters

37

Figure 3.

The amount of rainfall in inches influenced the number of organisms in a sample. Each dot represents a sample taken after the indicated amount of rain had fallen during a period of seven days.

O . .4 9

1.5

TO. 0

\.0 (inches)

«I 0 o 0

O O O

Rainfall

o‘F

0.5

O O O

O. 0.... Amount

O 0 0.0 000 .. ° .'. on. .0. ’ O 0 j m 1- 2:;

0 a 3 9 8 N Number of Macroarthropods /sample 39

Figure 4.

The position on the hill affected the number of organisms in a sample to a certain degree.

The number of the sample is the same as the plot number. This is plotted against the number of organisms in the sample.

>Topa? Hill

UPPer‘ Part 0? Siope

Lower Parr): OF Siope

Base oi" Hi”

C L

25 50 100 200 400 Number or Macmarihr‘opods / sample

41

Figure_5.

The temperature had an effect on the number of organisms in the sample. Each dot represents the number of organisms found at that temperature.

75°

o. T ‘ o o 70"

(°F)

‘ ‘J u. u 60°

55° Temperatune

50°

, .. .L . .

45°

o 8 3 3 d .....

Number of Macroarthropods / sample 43

Figure 6.

Months sampled was plotted against the number of organisms in the sample to show an increase in the later summer months. The red line is a hypothetical line to show the curve in

distribution.

——

i“ ‘A

a; Oct,

[Sepbnber

LA 3 1.. . W TAugust

_U

0 O. .9

0019 o. l x, g .0 m g \ c (n ‘3 D Ln 5’ f2 \ as 5 \ 5: Z

:2 O. 0.. O O. < o—

Winter Months , " _.__. ———————— (we

1 06+...le 3 8 9 6 Number of Macroarthropods / Sample 45

Figure 7.

The number of macroarthropods plotted against the pH of the sample. The average pH was around 5.1. Each dot represents a sample taken. The green circles represent samples in which the number of organisms is large because of the increase in number of individuals of one taxon rather than to the arthropods as a whole.

GI 8

sample

/ 8

S

croarthr0pods (9

Ma GI 8

0?

Number ZR

ILL

‘I‘ U

In0. W 4.5 In

6.0 65 pH Reading LIST OF ARTHROPODS FOUND

Phylum Arthropoda Class Eucrustacea Order Isopoda Porcellionidae Porcellio s2, -Class Arachnida Order Pseudoscorpiones Neobisiidae Microbisium s2, Order Opiliones Phalangiidae Liobunum sp, Phalanqium s2, Order Araneae Argiopidae Attidae Clubionidae 5”9* ~Lycosidae ' Gnaphosidae Thomisidae Theriidae Salticidae Order Acarina Suborder Mesostigmata Parasitidae Veigaiaidae Macrochelidae Laelaptidae Zerconidae Uropodidae Circocyllibanidae . Suborder Trombidifromes Scutacaridae Pyemotidae Rhagidiidae Bdellidae Trombidiidae Trombiculidae Suborder Acaridiae Saproglyphidae. ,

49 50

Suborder Oribatei Carabodidae Ephilophnammidae Galunmidae Camisiidae Eremaeidae Belbidae Ceratozitidae Phthiracaridae Class Insecta Order Protura L Acerentomidae Order Thysamura Campodeidae »eOrder Collembola Suborder Arthropleona Poduridae Hypogastrura s2, Neanura muscorum Templeton Onychiuridae Onychiurus subtenius Folsom Isotomidae Folsomia fimetaria Linnaeus Proisotoma _p, ‘ Isotoma trispinata MacGillivray Isotoma viridis Bourlet Isotoma albella Packard Mydontidae Mydonius multifasciata Tullberg Tomocerus vulqaris Tullberg Tomocerus flavescens Tullberg Pseudosinella pettersini Borner Pseudosinella pettersini f. rolfsi Mills Lepidocyrtus curvicollis f. achromatis Maynard Lepidocyrtus cyaneus Tullberg

Lepidocvrtus lanuginosus f. albicans Reuter Orchesella ainslici Folsom Suborder Symphypleona Sminthuridae Sminthurus s2, Neelidae Neelus minutus Folsom Order Orthoptera Tetrigidae Gryllidae «Blattidae 51

Order Psocoptera Psyllipsocidae Order Thysanoptera Phloeothripidae Thripidae -»Order Hemiptera Anthocoridae Reduviidae Nabidae -vLygaeidae Miridae <—Pentatomidae -.Order Homoptera Cercopidae Cicadellidae I~Order Coleoptera Lnrr* Cicindelidae :3 Carabidae!“ Ptiliidae zi‘f ~rStaphylinidae HM Elateridae ’V' ~Nitidulidae’fi CoccinellidaeIGI MordellidaeJW? Anthicidae/55* Bostrichidae222 Scarabaeidae 2:; CerambycidaeJLW Chrysomelidae272 «CurculionidaesI Scolytidae ?¥? Order Tricoptera4”“ HydrOptilidae Order Lepidoptera Noctuidae 'eOrder Diptera 25* .~» Chironomidae Culicidae Asilidae Drosophilidae Pipunculidae \ uoOrder Hymenoptera Ichneumonidae Braconidae 52

Chalcidoidea ~.«Formicidae pDiapriidae "Class Diplopoda Polyxenidae Polyxenus sp, Polydesmidae Fontaria s2, Polydesmus s2, Julidae Julus ER: Parajulus s2, , Class Chilopoda "Lithobiidae ' ~vLithobius s2, Geophilidae Geophilus s2, Scolopendridae Scolopendra gp, BIBLIOGRAPHY

Allee, C. W., K. Emerson, T. Park, 0. Park, and K. SChmidt, Principles 9f.Animal Ecology, W. B. Saunders Co., Philadelphia,

Pennsylvania, 1949, pp. 216—227, 461-466.

Bornebusch, C. H., The Fauna g§_Forest Soil, Neilsen and Lydiche, Copenhagen, 1930, 249 pages.

Dowdy, W. W., "The influence of temperature on vertical migration of invertebrates inhabiting different soil types," Ecolggy, 1947, 25, pp. 418—439.

Engelmann, M. D., "The role of soil arthropods in the energetics of an old field community," Ecological_Monographs, 1961, 31, pp. 221-38.

Gasdorf, Edgar C., and Clarence J. Goodnight, "Studies on the Ecology of Soil Arachnids," Ecolggy, Spring 1963, Vol. 44, No. 2.

Hairston, N. G. and G. W. Byers, "The soil arthropods of a field in southern Midhigan: a study in community ecology," Contribs. Lab. Vertebrate Biol. Univ. Michigan, 1954, 64, pp. 1-37. '

Kevan, D. K. McE., "A practical key to the orders and sub- orders of soil and litter inhabiting animals," Soil Zool. Proc. Unix, Nottingham, 1955, 1, pp. 452-88.

Kevan, D. K. McE., "Soil Zoology," Proceedings of the Uni- versity of Nottingham Second Easter School in Agricultural Science, 1955, Academic Press Inc., Pub., New York, 1955.

Kevan, D. K. McE., "Identification of soil and litter inhabiting animals," Soil Zool. Proc. Nottingham, 1955, 1, pp. 23-28.

Kuhnelt, W., Soil_Biology, Faber and Faber, London, 1961, 397 pages.

Kuhnelt, W., "An introduction to the study of soil animals," Soil Zool. Proc. Univ. Nottingham, 1955, 1, pp. 3-22.

54

Lawrence, R. F., The Biology 9; the Cryptic Fauna Q§_Forests with Special Reference §g_the Indigenous Forests 9; South Africa, A. A. Balkema, Cape Town-Amsterdam, 1953, 408 pages. a; 3'3 USE UIHLi

Ilia.“

o0

r

‘1

EL

9M)...-

4.1,.» ALL]

A31? 5....qu

U

1.,

.

.

.p.

4

I. .1.‘

IIIIIIIIIIIIIIII