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This is to certify that the

thesis entitled

EXAMINATION OF THE ARTHROPOD COMMUNITY ON A
GOLF COURSE FAIRWAY AND ROUGH

presented by

Breana Lee Simmons

has been accepted towards fulfillment
of the requirements for

M.S. Entomology

degree in

 

 

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Date 01‘ laa’o\

 

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6/01 c:/CIRC/DateDue.p85-p. 15

EXAMINATION OF THE ARTHROPOD COMMUNITY ON A GOLF COURSE
FAIRWAY AND ROUGH

By

Breana Lee Simmons

A THESIS

Submitted to
Michigan State University
In partial fulfilhnent of the requirements
For the degree of

MASTER OF SCIENCE
Entomology

2001

ABSTRACT

EXAMINATION OF THE ARTHROPOD COMMUNITY STRUCTURE ON A GOLF
COURSE FAIRWAY AND ROUGH

By

Breana Lee Simmons

The arthropod community structure was studied in the fairway and rough on the eighth
hole at Groesbeck Golf Course in Lansing, Michigan. Pitfall traps (32ml) were used to
study surface arthr0pod activity. Tullgren-type funnels were used for heat extraction of
soil samples in order to determine the abundance and distribution of soil-dwelling
arthropods on a golf course fairway and rough. In both 1999 and 2000, Staphylinid
adults were 3-fold more active in the rough compared with the fairway. Collembola and
Acari were also 2 to 3-fold more abundant in the rough compared with the fairway in
both years(P<0.05). Carabidae were significantly but only slightly more active in the
rough in 1999, and significantly but only slightly more active in the fairway in 2000.
Activity of Formicidae was greater in the fairway than in the rough, in both 1999 and
2000. The number of all species recovered decreased in 2000 compared with 1999. High
levels of Collembola and Acari in the rough compared with the fairway may create a food

chain for supporting more generalist predators such as Staphylinidae, in the rough.

For mom, dad, andy and steph, thanks.

iii

ACKNOWLEDGMENTS

First and foremost, I would like to thank my major professor Dr. David
Smitley, and my committee: Dr. Maria Davis, Dr. Deb McCullough, and Dr. Richard
Snider, for all their guidance and support during my project. Thank you for having
faith in me.

Also, a big thanks to Terry Davis for keeping me on track and out of trouble.

I appreciate you! Thank you to the Landscape Entomology crew: Abby, Keri,
Rhonda, Mark, Jeremy, Marija, Lynette, Mike, Julia, Leah and Darren - you make me
laugh! I will never forget watching Marija run away from rampaging irrigation
sprinklers, or strapping Rhonda to a golf cart, and I will always remember to “wear
my big head.” Also, I would like to extend my gratitude to Dr. Roy Norton, Dr.
Richard Snider, Dr. Jim Zablotney, Gary Parsons, and Brian McComack for their
arthropod identifications.

Credit should go to my family for keeping me inspired. I have never had to
seek out their approval, it was always there. I would like to express my appreciation
for my fiiends that have been so supportive along the way, and were always ready to
listen to me rant and rave about anything. Meghan, Jen, Chanda, Natalia, Alicia,
Brian, Chuck, Aram, Jana, and John — thanks for the memories! To the Montreal
Sock Posse (St. Anne, The Walk, and QK), you guys are going to make a crazy bunch
of entomologists one day. Thanks to those of you who worked with me in the Bug

House, especially Barb and Laura, it was a great experience.

A special thanks to John Johnson and his terrific crew out at Groesbeck Golf
Course. It’s not often you find someone willing to let you rip up his fairway in search

of soil insects he can’t even see.
Support for this project was provided in part by the Michigan Turfgrass
Foundation and the Hutson Student Grant. It’s been grand, thank you to everyone in

the entomology department for looking out for me.

TABLE OF CONTENTS

LIST OF TABLES

LIST OF FIGURES
LITERATURE REVIEW
INTRODUCTION

MATERIALS AND METHODS
Study Site
Sampling for subsurface arthropods
Pitfall trapping
Arthropod identification
Sampling for A. spretulus, A. granarius, and P. japonica
Soil tests
Heat extraction efficiency
Statistics

RESULTS
Arthropod identification
Surface and subsurface sampling
Sampling for A. Spretulus, A. granarius, and P. japonica
Simple regression tests
Soil tests
Heat extraction efficiency

DISCUSSION
Arthropod identification
Surface and subsurface sampling
Sampling for A. Spretulus, A. granarius, and P. japom'ca
Heat extraction efficiency
Mowing height regimes

LITERATURE CITED

APPENDIX 1 Record of Deposition of Voucher Specimens

vi

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LIST OF TABLES

Table 1. Activity of adult A. spretulus, A. granarius, Carabidae, Staphylinidae,
Formicidae and others at a Golf Course in Lansing, Michigan.

Table 2. Numbers of Mites, Collembola, Formicidae, and others extracted from soil
cores taken from Groesbeck Golf Course in Lansing, Michigan.

Table 3. Efficiency of pitfall traps for capturing F ormicidae. Data are the ratios of ants
caught in pitfall traps to ants found in soil cores, grouped by block.

Table 4. Larvae recovered fromsoil cores taken from 25 May to 20 July 2000 at
Groesbeck Golf Course in Lansing, Michigan.

Table 5. Relationship among the major groups of arthropods found within 2m of the
border between the fairway and the rough on a single hole at Groesbeck Golf Course in
Lansing, Michigan. Data are the seasonal distributions after loglo transformations.

Table 6. Relationship of potential adult predators caught in pitfall traps to the
distribution of Japanese beetle larvae (y) collected in May and early June 2000.
Regression statistics are the seasonal distributions after loglo transformations of the data.

Table 7. Data from soil moisture tests using the Aquaterr ® soil moisture meter. Data
are the means i SD per ten probes per block per treatment over 3 sampling dates in 2000.

Table 8. Results of the heat extraction efficiency for selected groups of arthropods based
on the total number of specimens obtained per treatment. %Efficiency = [(N individuals
obtained by heat extraction)/(N total individuals obtained by heat extraction plus soil
floatation)] x 100; F and R = Fairway and Rough.

vii

LIST OF FIGURES
Figure 1A. Experimental design on new #8 at Groesbeck Golf Course in Lansing, MI.
Figure 1B. Example of a single block of the experiment at Groesbeck Golf Course.

Figure 2. Results of surface sampling for arthropods on Groesbeck Golf Course. Bars
represent seasonal distributions of the arthropods caught in pitfall traps between May and
July 1999.

Figure 3. Border effect shown in staphylinids caught in pitfall traps between 25 May and
20 July 1999 at Groesbeck Golf Course in Lansing, Michigan. Data are the total trap
catch for each pitfall trap. Grey columns are the pitfall traps at 0.5m to 2.0m from the
turf border'in the fairway, while black bars represent pitfall traps at 0.5m to 2.0m from
the turf border in the rough.

Figure 4. Results of surface sampling for arthropods on Groesbeck Golf Course. Bars
represent seasonal distributions of the arthropods caught in pitfall traps between May and
July 2000.

Figure 5. Results of subsurface sampling for arthropods at Groesbeck Golf Course. Bars
represent seasonal distributions of the arthropods heat extracted from soil cores between
May and July 1999.

Figure 6. Results of subsurface sampling for arthropods at Groesbeck Golf Course. Bars
represent seasonal distributions of the arthropods heat extracted from soil cores between
May and July 2000.

Figure 7. Relationship between Collembola extracted from a soil core and the
staphylinids caught in the 2 adjacent pitfall traps at Groesbeck Golf Course in 1999.
Equation for regression statistic can be found in Table 5.

Figure 8. Relationship between mites extracted from a soil core and the staphylinids
caught in the 2 adjacent pitfall traps at Groesbeck Golf Course in 1999. Equation for
regression statistic can be found in Table 5.

Figure 9. Precipitation data from East Lansing, Michigan for April to July 1999 and
2000.

Figure 10. Temperature data in degrees F from East Lansing, Michigan for April to July
1999 and 2000.

Figure 11. Accumulated degree-days (base 50 degrees F) for East Lansing, Michigan for

both field seasons. Data was taken from a weather station at the Horticultural Gardens at
Michigan State University.

viii

LITERATURE REVIEW

Grasses cover a larger portion of the world’s land surface than any other
vegetation type (Tscharntke and Greiler 1995). The grasses, Gramineae, appeared in the
fossil record during the late Cretaceous period, evolving with grazing mammals (Beard
1973). Temperate grasslands are dominated by insect herbivores and tropical grasslands
are dominated by large ungulate herbivores (Tscharntke and Greiler 1995). It can be
inferred that since grasses are adapted to grazing by herbivores, and compensate for
herbivory with rapid regrowth, then they also tolerate being mowed. Young plants are
usually more nutritious for herbivores, but may actually deter certain insects due to high
concentrations of secondary compounds (Tscharntke 1988). Most grasses, however, are
simply structured and do not have toxic defenses; therefore they do not deter herbivory
(Bemays 1990, Bernays and Barbehenn 1987).

Lawns and golf course turf is similar in many ways to native grasslands.
Turfgrasses are herbaceous plants that can be annual or perennial. There are 40 species of
turfgrasses in the Poaceae family that are utilized for various purposes throughout the
world (V ittum et al. 1999). While turf is considered mainly ornamental or recreational, it
is also of economic importance, not only for control of wind and water erosion of soil,
but also for climate control. Turfs reduce glare, noise, air pollution, and heat buildup in
urban areas (Beard 1973). For recreational purposes, well maintained turf is not only
aesthetically pleasing, but in many cases it is necessary for functional purposes, such as
tight, short grass for putting. Turfgrass health and performance is essential to the

recreation industry as well as to homeowners and landscapers.

Contrary to pOpular opinion, grasses harbor the heterogeneity necessary for the
evolution of a diverse insect community (Tscharntke and Greiler 1995). While studies
have shown that sown seed, such as that found on a golf course, will not sustain the large
and diverse populations of insects that are found in natural grasses (Harper 1977), the
diversity of insect species found in monoculctures of cereal or fodder grasses is
surprisingly high (Altieri 1991). Root feeders and other soil invertebrates are usually
more abundant on grazed or mowed grasses than ungrazed or unmowed grasses (Grieler
and Tscharntke 1993). Damage caused by root feeders may increase the abundance of
foliar feeders due to the increased vulnerability of the plant (Masters et a1. 1993). Long
grasses support more predacious and saprophagous beetle families than does mowed
grass. Mowing reduces seed and foliage feeders, which also reduces predators (Morris
and Rispin 1987). As the number of mowings per year increases, permanent turfgrass
communities, both plant and animal, become less diverse (Grieler and Tscharntke 1993).
Cutting decreases species richness of insects, and cessation of mowing results in changes
in insect populations, sometimes resulting in increased species richness (Morris 1981,
Morris and Plant 1983). As the cutting height is lowered, turf plants exhibit decreased
carbohydrate synthesis and storage, increased shoot growth, and decreased root growth
(Beard 1973), which may explain changes in insect diversity as well.

Insect outbreaks in natural grasses are common and many times are not
threatening to grass health. When a particularly large outbreak occurs it can cause cycles
of damage resulting in substantial loss of biomass (Henderson 1978). However, on well-
maintained turf, such as on golf course fairways, greens, and tees, even small outbreaks

are almost immediately problematic (Beard 1973). Although grasses show low

2

susceptibility to grazing pressure due to their rapid regrowth, root feeding may be more
problematic, as it weakens the plant from the more susceptible underside. According to
Stanton (1988) soil organisms are the limiting factor to grassland productivity, as
microbial grazers may serve as regulators of plant growth.

Of the herbivores, white grubs (Coleoptera: Scarabaiedae) may be the most
damaging to turf because they feed on the more susceptible roots (Potter 1998). Popillia
japonica (N euman), for example, is an introduced species that is considered the single
most important scarab turf pest in the United States (V ittum et al.1999). The larvae of P.
japom'ca cause significant damage to turfgrass in eastern North America by chewing the
roots. The adults are also a major pest, feeding on foliage or flowers of nearly 300
species of plants (T ahsiro 198%. P. japonz'ca overwinters as larvae below the frost line,
moving to the surface to pupate in May and June. Adults are usually present in June and
July, and larvae are found again in August and September, feeding on turfgrass roots until
cold weather forces them further down into the soil (Potter 1998).

Outbreaks of grubs, especially invasive pest species, are devastating to turf, and
we know very little about why outbreaks occur. To understand why outbreaks of many
different species of grubs occur in lawns and golf courses, we need to have a better
understanding of the arthropod community structure in the soil, where scarabs spend
three-quarters or more of their life as eggs and larvae.

In the soil, Stanton (1988) allows for two major trophic levels: herbivory and
decomposition. However, in studying insects it becomes clear that predation is also a
major component of the grasslands system. Of course, prey availability is the most

important factor in determining the diet of any predatory arthropod (Pollet and Desender
3

1987), and soil fauna are typically formd in numbers 2-10 times higher than their
aboveground counterparts (Lavigne et a1. 1972). If there is an abundance of prey
underground, then therefore there may be an abundance of generalist predators that may
also feed on scarab eggs and larvae.

In the soil, soil protozoans and nematodes are the most abundant animals,
followed by the microarthropods (Leetham and Milchunas 1985). These organisms are
important decomposers, but certain microarthropods also may be important predators in
the soil. Many mites in the suborder Mesostigmata may be important predators of insect
eggs, nematodes, Collembola and other small arthropods (Wallwork 1976). In leaf litter,
mites are generally the most important predators of Collembola, followed by
psuedoscorpions (Arachnida: Chelonithi), and members of the families staphylinidae and
carabidae (Coleoptera) (Christiansen 1964). Larvae of Staphylinidae and Carabidae are
important predators of Collembola and mites below the soil surface (Wallwork 1976
based on Luff 1966). Carabid larvae are especially adept at catching Collembola, which
are typically trapped in soil cavities where they cannot jump. If Collembola are
encountered in a larger arena, the carabid larvae still have the advantage, as their touch is
too gentle for the flight stimulus of the Collembola (Bauer 1978). The habitat of the
larvae of any carabid species is determined by the position of the egg, and as larvae are
not able to move long distances they must survive where the female oviposited (Lovei
and Sunderland 1996).

According to Theile (1977) carabid activity in temperate zones is based on their
reproductive habits, whether they are summer or autumn breeders, larval or adult

hibernators, and if there are dormancy periods in the adults. In a study spanning 10 years

4

in Western Europe den Boer and den Boer-Daajne (1990) reported the reproductive
cycles of the 68 most abundant carabid species of Drenthe (The Netherlands) including
Amara aenea (Degeer) and Pterostichus melanarius (Illiger) which are also found in
Michigan. A. aenea is a spring breeder, reproductive between April and June with larvae
present between July and October, and overvvintering as adults (den Boer and Den Boer-
Daajne 1990, Kegel 1990). P. melanarius is an autumn breeder, reproducing between
June and September with larvae present between January and May (den Boer and Den
Boer 1990).

Collembola are relatively abundant above and below the soil surface, where they
feed on a variety of materials such as plant material, fimgi, and bacteria (Butcher et a1.
1971). The abundance of these food items are of major importance to the distribution of
soil Collembola. Collembola are rapid reproducers that may become resistant to
pesticides (Christiansen 1964). They are bionomically divided into three major life
forms, epigious, hemiedaphic, and euedaphic. Epigeous Collembola are those with well-
developed eyes, antennae, pigmentation and have a long furcula. These are the surface
Collembola. Hemiedaphic Collembola have moderately long antennae, pigmentation, and
well-developed eyes. These are found in ground litter, moss and bark. Euedaphic
Collembola are true soil dwellers, with reduced eyes, short antennae and little pigment.
They inhabit the deep soil, as well as caves and soil cavities (Butcher et a1 1971). Some
soil dwelling Collembola, unable to jump within the soil cavities, are capable of secreting
noxious fluids as a defense against predators Willani et al, 1999).

After six years of studying 2 species of small grubs (Ataenius spretulus and

Aphodius granarius) and generalist predators on golf course fairways and roughs, it is

5

still not understood why Staphylinids (a generalist predator) are much more abundant in
the roughs, while scarab larvae are much more abundant in the fairway (Smitley et a1
1998, Rothwell and Smitley 1999).

I believe that a broader examination of the entire arthropod community in
pesticide-free fairways and roughs may reveal a relationship within the food chain that
will help explain outbreaks of A. spretulus and A granarius on golf course fairways. If
we examine these scarabs as part of a larger system, rather than as an isolated pest of
turfgrass, it may be possible to predict, treat, and possibly prevent outbreaks more

effectively than our current knowledge of them allows.

INTRODUCTION

The black turfgrass ataenius, Ataem'us spretulus (Haldeman), and a similar
looking beetle, Aphodius granarius (L.) (Coleoptera: Scarabaeidae), are pests of turfgrass
(Potter 1998). Larvae of both species consume turfgrass roots, causing patchy damage to
turf. A. spretulus is native to North America, and is an important pest of golf course
fairways, greens and tees in the eastern and mid-western United States and some
provinces of Canada (N iemczyk and Dunbar 1976). The first report of damage caused by
A. spretulus came from a golf course in Minnesota in 1927. However, A. spretulus was
not well documented as a turf pest until after 1970, when the amount of reported damage
rose sharply (Tashiro 1987). This increase was apparently due to development of
resistance to insecticides (Weaver and Hacker 1970, Niemczyk and Wegner 1982). All
reports of turf damage by A. spretulus have been limited to golf course fairways, greens
and tees (N iemczyk and Wegner 1982).

A. spretulus completes two generations a year in regions south of central Ohio and
has one generation a year in northern Ohio and Michigan (Wegner and Niemczyk 1981,
Smitley 1994). In southern Ohio, overwintering adults deposit eggs in May, larval
densities reach their peak in June, and a new generation of adults emerges in July. Eggs
are laid again in July developing into larvae in August (Potter 1998). In Michigan, eggs
are laid in June and larvae are present in July (Jo 2001).

A. granarius adults are very similar in appearance to A. Spretulus adults.
European in origin, A. granarius is a pest of golf courses in Ontario, Michigan, Colorado,
and Ohio, where it was originally mistaken for A. spret'ulus (Tashiro 1987). Adults of A.

granarius are distinguished from A. spretulus adults by the presence of transverse

7

carinae on the hind tibia, while the larvae of each can be identified by the rastal pattern
(Tashiro 1987). A. granarius larvae damage turf in the fairway in a pattern similar to
that of A. spretulus. A. granarius was originally believed to complete 2 generations in
Ontario and Ohio due to the presence of adults in May and then again in August (Tashiro
1987). However only one generation of A. granarius larvae apparently occurs in New
Jersey (Wilson 1932) and Michigan (Smitley 1994). In Michigan, A. granarz'us adults
overwinter and become active again in May. The larvae are present in June, about four
weeks earlier than those of A. spretulus (Smitley et al. 1998).

In a recent study, A. spretulus and A. granarius larvae were found to be most
abundant in the short turf of the fairway (1-2 cm); while surface predators (Carabidae and
Staphylinidae) were uncommon in short turf, but abundant in the adjacent rough (5-7 cm)
(Smitley et al. 1998). The activity of A. Spretulus and A. granarius may be inversely
proportional to the number of insect predators on the ground surface. When mowing
practices were changed, A. spretulus and A. granarius numbers increased in the shorter
fairway while surface predator numbers increased in the longer rough (Rothwell and
Smitley 1998). These differences existed in the absence of pesticides, although
differences may be even greater when pesticides are used (Smitley et al. 1998). The
differential distribution of these predator and prey species in different habitats suggests
predation affects the abundance of A. spretulus and A. granarius in turfgrass.

The activity of Carabids and Staphylinids was shown to be lower in the fairway
compared with the rough (Rothwell 1998) and we suspected their primary source of food
is also different in the fairway and rough. One potential source of prey for Staphylinid

and Carabid adults and larvae are several species of Collembola and mites (Pollet and

8

Desender 1987). The microarthropod fauna in the soil may be especially important to
larvae of Carabids and Staphylinids because these carabid and staphylinid larvae reside
entirely in the soil. Both the Collembola and mites can be found in all layers of soil,
including the thatch layer of turfgrass. If populations of generalist predators are
dependant on small arthropods in the soil, as well as surface prey, then outbreaks of pest
species in the fairway may occur because the bottom of the food chain is too small to
support enough generalist predators to control invasions of turf pests.

The objectives of this research were to determine the abundance of Collembola
and mites in the soil of a golf course fairway and in the rough, and to determine the

relationships between the soil fauna, the activity of insect predators and the presence of

scarab larvae.

MATERIALS AND METHODS

Study Site. This study was done on a single hole at Groesbeck Golf Course in
Lansing, Michigan. A sprinkler head in the center of the fairway provided irrigation
coverage for both the fairway and the rough. The soil is generally loamy, and the turf is
at least 25 years old, and is a mixture of bentgrass (Agrotis Spp. L.) and annual bluegrass
(Poa annua L.), with Kentucky bluegrass (Poa pratensis L.) and perennial ryegrass
(Lolium perenne L.) moving in. In 2000, fine fescue (Festuca capillata Lam.) was found
to have invaded the inner circle of the fairway in large patches. On this particular hole,
the grass has not been reseeded or disturbed since 1927 (John Johnson, Superintendent,
Groesbeck Golf Course, personal communication). Currently, the rough seems to be in
transition back to P. pratensis, having been predominantly L. perenne. Pesticides are
sparingly used on this course, for economic reasons. The greens are treated regularly
with herbicides, and the fairways and roughs are spot treated for fungal outbreaks if
necessary. No insecticides were used on this area of the golf course in the two years prior
to this study.

Arthropod communities in the fairway and rough were compared with a
randomized complete block split-plot design. Six blocks were set up around the outside
of an oval shaped fairway (Fig 1A). Each block (5m x 2m) was centered on the border
between the fairway and rough (Fig 1A). Blocks were divided length-wise for taking soil
samples and for pitfall trapping.

Sampling for subsurface arthropods. Once a week between 25 May and 20
July in 1999, 4 standard golf course cup-cutter samples 12 cm in diameter and 12-13 cm

deep were pulled from each block and placed in sealed plastic bags. They were replaced

10

by pre-cut soil cores from the same turf type so as not to disturb the turf environment.
Soil cores were always taken at 1m and 2m away from the border but at different
distances (0-1.0m) away from the medial line to minimize disturbance to the area, and to
avoid collecting from the same spot twice.

The plastic bags containing the soil cores were placed in a large cooler for
transportatiOn to the field station at Michigan State University. The sampling took
approximately 1-2 hours between the hours of 9 am and 11am. The samples were gently
crumbled by hand inside plastic bags and searched for visible arthropods at the field
station. The turf at the top of the soil core was also separated into large pieces to allow
arthropods to more easily escape from the samples during heat extraction. Any
arthropods found at this time were placed in 70% ethanol and transported to the
Landscape Entomology lab at Michigan State University for identification.

After visual inspection, the contents of the plastic bag were poured onto a screen
in Tullgren-type funnels to collect arthropods too small to observe without magnification.
Before the study began, the 24 Tullgren-type funnels were set up in 2 large racks. The
lips of the funnels were sprayed with Teflon ® to keep contaminating species out.
During the experiment, the funnels were heated with 40-watt bulbs connected to a
rheostat. Each day the heat was slowly increased over a period of 7 d, or until the sample
was dry. Specimens extracted from the soil cores were collected for counting and
identification by Dr. R.J. Snider (MSU), Breana Simmons (MSU), Brian McComack
(MSU), Gary Parsons (MSU) and Dr. Roy Norton (Syracuse, NY). Those that were
unidentified were categorized by morphotype and placed in an “other” category. This

experiment was repeated exactly between 22 May and 20 July 2000.
11

Pitfall trapping. Small glass vials (32 ml) filled with ethylene glycol were used
to sample surface insects. Eight pitfall traps were placed 0.5 m apart in a line
perpendicular to the border between fairway and rough in each block. Pitfall traps were
collected and the insects were sorted weekly (Fig 1b). The pitfall traps remained at the
same location throughout the study. In 1999 the traps were set between 25 May and 20
July, coinciding with the soil sampling. In 2000, the traps were set earlier, on 10 April, to
detect adult activity of Aphodius granarius. Specimens of the families Carabidae were
identified by Brian McComack (MSU), Staphylinidae by Breana Simmons (MSU) and
Curculionidae by Gary Parsons, (MSU). As in the soil sampling, those that were
unidentified were placed in the “other” category.

Arthropod identification. From each pitfall trap, the first individuals counted
(up to five) in each family were kept for identification to the species level. From each
heat extraction sample, the first five mites and collembola counted were collected for
further identification. Also collected were any unknown individuals, so that they could be
identified and categorized by family if necessary.

Sampling for A. spretulus, A. granarius, and P. japonica. To determine the
distribution of A. spretulus and A. granarius larvae, soil cores were collected during the
peak incidence of third instars, when grub visibility is highest. For A. granarius, this is
usually in the first 2 weeks in June and A. spretulus larval densities peak 4-6 weeks later.
Eight cup-cutter samples were taken from the fairway and rough of each of the 6 blocks
and dissected for grubs on site. The grubs were collected for species identification and
the soil cores placed back into the ground. On 19 June 2000, we also sampled for

P. japonica larvae in this manner. P. japom'ca was sampled because the population at

12

Groesbeck was predicted to rise to pest proportions, much like A. granarius had in
previous years. For analysis, these were added to any larvae recovered from the soil
cores that were subjected to heat extraction.

Soil tests. The Aquaterr ® soil moisture/ temperature probe (Aquaterr
Instruments Inc. 1990) was used to measure soil moisture in each block to ensure that the
blocks were homogeneous. The probe was calibrated in a bucket of water before each use
to ensure accuracy. Ten soil probes per block in both the fairway and the rough were
recorded for 3 consecutive weeks between July 17 and July 31, 2000. The equipment
was not available before this time period, and was not used past 31 July because the time
period of the study had already past. Temperatures and moisture readings were averaged
by fairway and rough and analyzed for statistical significance. Also, weather data was
obtained from the Michigan Automated Weather Network Interactive Data Access
website (2000) for East Lansing Michigan, which was as close to the research site as
possible.

Heat extraction efficiency test. According to Snider et al. (1997) a heat
extraction efficiency test must be run on field samples at least once to determine the
actual percentage of individuals recovered from the extraction process. Heat does not
extract all individuals from a soil sample. Some individuals are dead before placement in
the funnels, and some die before they make it all the way through the sample and into the
alcohol. Therefore, in order to determine the actual number of individuals in the sample,
it is necessary to float out any remaining specimens after the heat extraction has been
completed. Floatation is more accurate then heat extraction, but is time consuming and

labor intensive. Heat extraction is a standard method for sampling soil invertebrates, and

13

is an acceptable means of collection, as long as an efficiency is determined (Snider et al.
1997). In 1999, 24 soil samples were taken in August and placed in the Tullgren funnels
for 7days. Each sample was retrieved from the fimnel and placed in a 0.946 liter jar. The
jar was filled with a saturated sugar solution and shaken. The contents of the jar were
allowed to settle for 2 hours to allow the organic matter to rise to the surface. The
solution was decanted through a ZOO-mesh sieve into a plastic container. Care was taken
to ensure that none of the silt that had settled to the bottom was poured onto the screen.
Any organic material on the mesh screen was placed in a 0.473 liter jar and preserved
with 95% ethanol. The sugar solution was poured from the bowl back into the jar
containing the sediment, shaken again, and allowed to sit for 2 hours. This process was
repeated twice more (Snider and Snider 1997). Invertebrates collected from the original
heat extraction were counted and compared with the number of invertebrates found in the
sugar floatation experiment.

Statistics. Due to the initial design of the project, a split plot analysis was run
using SAS ® (SAS Institute,l990). The whole plot factor was the treatment (fairway v
rough), and the sub-plot factor was the distance from the fairway/rough border. An
ANOVA was created for each individual group of arthropods to determine if the
treatment was significant within the group. Due to concerns about the non-parametric
nature of the data, another test was run using a log regression model in SAS ®. In this
model, a regression was created for each group of arthropods, which could determine
significant differences between blocks, treatments (whole plot and sub plot), and weeks.
In 2000, only the pitfall trap data that corresponded in time with the 1999 data was used

for analysis. Data from April and early May were not included, because sampling at that

14

time was only intended for trapping A. granarius. Also not included in the analysis was
count data collected from week #6, which was considered to be inaccurate due to
exceptionally heavy rains, which, combined with sprinkler irrigation system installed on
the eighth hole, flooded the pitfall traps. In addition to these statistical procedures,
simple regressions were run using SUPERANOVA ® (Abacus 1991) to show

correlations between the distribution of predatory arthropods and potential prey items.

15

 

RESULTS

Arthropod identification. Carbidae recovered from pitfall traps in 1999 and
2000 consisted mainly of Amara anea (Degeer). Of 135 specimens kept for identification
126 were A. aenea. Other species of Carabidae recovered included Scarites quadriceps
(Chd), Stenolaphus comma (F.), Anisodactylus rusticus (Say), and Pterostichus
melanarus (Illiger). Most of the mites extracted from soil cores were found to be of the
suborders Mesostigmata and Prostigmata (Families: Eupodidae, Tetranychidae,
Pygmephoridae). Only 2 individuals were found to be of the family Oribatidae:
Epilohmannia elongata (Banks) and Graptoppia italica (Bernini). Of the suborder
Astigmata, members of the genus Tyrophygus were identified. All Staphylinidae kept for
identification were identified as Philonthus cognatus (Stephens) which are known
predators of turf pests (J o 2000). All Curculionidae kept for identification were
identified as Sphenophorus minimas (Hart) which are pests of turfgrasses (V ittum et al.
1999). The most abundant collembola at Groesbeck Golf Course are Psuedosinella rolfsi
(Mills), and Isotoma notabilis (Folsom). P. rolfivi is a common collembola found in
grassy areas. P. rolfsi has a well-developed furcula for jumping and is found in the top
layers of soil. 1. notobilis is another common species frequently found in lower Michigan
(Snider 1967).

Surface and subsurface sampling. In 1999 there were differences in Carabidae
activity between the fairway and the rough as determined by both parametric and non-
parametric statistical tests (Fig 2). The activity of Carabids in the rough
(6.9/10traps/week) was significantly higher (DF 1,7, F =5.75, P=0.02) in the rough than in

the fairway (4.8/10traps/week). In 2000, the activity of Carabidae was significantly
16

higher (DF 1,7, F=6.13, P=0.01) in the fairway (8.9/10 traps/week) than in the rough
(6.7/10 traps/week), just the opposite of what was observed in 1999 (Fig 4). Total trap
catch for Carabidae was similar in both years (208 individuals caught in 1999 and 212 in
2000)

Staphylinid activity was significantly greater (DF1,7, F =65 .76,P=0.0001) in the
rough than in the fairway (T able 1). In 2000, Staphylinidae were again significantly (DF
1,7,F=52.41,P=0.0001) more active in the rough than in the fairway (Fig 4). Total trap
catch for Staphylinidae decreased from 407 individuals caught in 1999 to 213 individuals
caught in 2000.

Due to the way the experiment was designed, it was also possible to examine the
data for a border effect. A border effect is a phenomenon in which the abundance of an
insect changes as you approach the border of a different habitat. If it is beneficial to
exploit both the fairway and the rough, then the individuals will remain near the border
between both habitats, and there will be a build up of activity nearest the border on both
sides. In the rough, the number of Staphylinidae caught in pitfall traps was constant
between 1.0m and 2.0m of the border but nearly doubles within 0.5m of the border with
the fairway (Fig 3). A slight increase in Staphylinid activity on the fairway side of the
border suggests some movement from the rough into the first 1.0m of the fairway,
however, this may simply be spillover. There were significantly more Staphylinids
caught at 0.5 m from the border in the rough than in any other pitfall trap in 1999. This
may indicate that the Staphylinids are not utilizing both habitats, but are stopping before
they crossover into the fairway. However, no other significant differences were found

between pitfall traps in the same grass length.

17

In 1999, number of Collembola extracted from soil samples was 2-fold higher
(DF1,3,F=43.7,P=0.0001)in the rough than in the fairway (Table 2, Fig 5). Similarly, the
number of mites extracted from soil samples was 3-fold higher (DF1,3,F=58.5,P=0.0001)
in the rough than in the fairway. In 2000, the total number of Collembola and mites
recovered from soil samples was reduced compared with 1999. In 1999, a total of 9234
Collembola and mites were extracted from soil cores, whereas in 2000, only 2722
individuals were extracted. However, despite the reduction in abundance, the numbers of
both Collembola (DF1,3,F=19.7,P=0.0001) and mites (DF1,3,F=19.5,P=0.0001) were
again 2 to 3-fold higher in the rough than in the fairway (Table 2, Fig 6). No significant
border effect was found in either of these groups in either 1999 or 2000.

The unidentified insects included members of the orders Diptera, Coleoptera
(Elateridae, Tenebrionidae, Curculionidae), and Homoptera. The total number of
unidentified insects found in both surface and subsurface sampling was also higher in the
rough than in the fairway in 1999 (Tables 1 and 2). In 2000, a separate category for
Curculionidae was added to the pitfall trap data because we began to catch them in high
numbers and thought they could be important members of the arthropod community, as
many curculionids are pests of turfgrass. For this reason we also added a category for
Homoptera (later classified Aphididae) in the soil data. In 2000, Curculionidae and
Aphididae were found in higher numbers in the rough (Table 2). In 2000, unidentified
insects from subsurface samples were higher in the rough (Fig 6). Data were collected on
spiders during both 1999 and 2000, but not presented because only 12 were caught in

pitfall traps in 1999 and only 17 were caught in pitfall traps in 2000.

18

More ants were caught in pitfall traps in the fairway than in the rough (Table 1),
while ants found in the soil were slightly more abundant in the rough compared with the
fairway in 1999 (Table 2). Because large numbers of ants were collected from soil cores
and pitfall traps, and the species composition was the same in the fairway and rough, we
could use the soil core data to compare pitfall trap efficacies in the fairway and rough.
The ratio of ants in pitfall traps to ants in soil cores was not different between fairway and
rough samples in 1999 (P=0.4) or 2000 (P=0.7), suggesting that the efficacy of pitfall
traps was similar in the fairway and rough (Table 3).

Sampling for A. spretulus, A. granarius, and P. japonica. A. granarius was
abundant in previous years at Groesbeck Golf Course, causing extensive damage to some
greens, but populations appeared to be very low in 1999 and 2000. In 1999 no A.
granarius larvae were found. In 2000, only 19 larvae were found, not enough to evaluate
differences between the fairway and rough. In 1999 only 10 A. spretulus larvae were
found, and in 2000, only 20 larvae were found. This amount of larvae was not enough to
evaluate differences between the fairway and the rough. No differences were discovered
for the adults of A. spretulus or A. granarius in 1999 or 2000 (Figs 2 and 4). While A.
granarius and A. spretulus were found in low numbers, numbers of Pjaponica adults
and larvae were on the increase, allowing us to obtain good data in 2000. More P.
japonica larvae were found in the fairway than in the rough (Table 4). The adults of P.
japom'ca were also found in higher numbers in the fairway than in the rough (Table 1).

Simple regression tests. Positive linear relationships were found for the 1999
data, and for combined 1999 and 2000 data, between the abundance of Staphylinidae and

the abundance of both mites and Collembola. In 1999, as the abundance of mites

19

increased, so too did the abundance of Staphylinidae (P=0.55) (Fig 7). Also, as the
number of Collembola present in the soil cores increased, so did the number of
Staphylinidae caught in adjacent traps (r2: 0.37) (F ig 8). No other significant
relationships were found between other major arthropod groups (Table 5).

For P. japonica, relationships between grub distribution and predator abundance
were varied (Table 6). No significant relationship was found between Carabidae caught
in 1999 or 2000 and P. japonica larvae found in soil cores in 2000. For Staphylinidae, in
both 1999 and 2000, significant negative linear relationships were discovered between
adult Staphylinids caught in pitfall traps and Pjaponica larvae found in adjacent soil
cores (r2=0.29, r2=0.45). Relationships between the distribution and abundance of ants
caught in both 1999 and 2000 and the larvae of P. japonica found in adjacent soil cores in
2000 were positive and minimal, although 1999 was significant to the p=0.05 level
(r2=0.2, r2=0.15)

Soil tests. No significant differences in moisture content or temperature were
found between the fairway and rough within blocks, nor were significant differences
recorded between blocks. We did not expect any differences in soil temperature or
moisture between the fairway and the rough, but we were testing for homogeneity.
Readings from the Aquaterr® meter registered different soil temperatures depending on
the ambient temperature data, and moisture levels were dependant on the time elapsed
since the last irrigation. However, despite these dependencies on surface conditions, the
readings were uniform throughout the plots on each sampling date. On a moisture scale
of 0-100, the fairway averaged 84.5 over the three sampling dates, while the rough

averaged 83.9 (Table 7).
20

Heat extraction efficiency test. As expected, the floatation experiment (T able 8)
resulted in more Collembola and mites than the previous heat extraction methods.
Collembola were heat extracted at a 40% efficiency level in the rough and a 45%
efficiency level in the fairway. Using both the floatation efficiency percentages and heat
extraction data (volume of soil core/mean number individuals found per core/percent
effiency) we can estimate that there were approximately 30,000 Collembola per m3 in the
fairway and 80,000 Collembola per m3 in the rough. Mites were extracted at a 52%
efficiency in the rough and a 44% efficiency in the fairway. As with the Collembola, this
means that there were approximately 29,000 mites per m3 in the fairway and 70,000 mites
per m3 in the rough.

Other insects collected included several types of unidentified Dipteran larvae and
Homoptera nymphs, which were categorized as “others” for statistical analysis in the heat
extraction experiments, thus they were classified in the same way for this test. No other
arthropods were found in the experiment. This does not mean that heat extraction of
other arthropods was 100% efficient, but that at that time, in those soil cores, no other

individuals were present.

21

DISCUSSION

Arthropod identification. The identification of the mites and Carabidae put a
new spin on the interpretation of the research results. First, Amara aenea are generally
phytophagous carabids, and are not considered predatory. As they made up the bulk of
the trap catch, this may explain the weak relationship between Carabids and prey items.
However, Pollet and Desender (1987) found Collembola in the gut contents of adult A.
aenea, suggesting that the species is not entirely phytophagous, and may be a predator of
Collembola, but probably not of larger arthropods. In the same study mites were
identified as a minor food source for Carabidae, due mainly to their subterranean
lifestyle, although the authors speculated that mites may be an important source of food
for carabid larvae. Although feeding data on this particular genus of carabids is not
available, some carabid larvae are known predators of Collembola, which they trap in soil
cavities (Bauer 1979). In laboratory studies, A. aenea was found to consume small
arthropods, but was presumed too small to consume scarabaeid grubs (Hagley et al 1982).
In feeding tests involving adult A. aenea and larval A. spretulus, in petri dishes, A. aenea
did consume this small third instar grub (Jo 2000). It is not known if the larvae of this
species are predators, but if they are, A. spretulus and other small scarab larvae as well as
insect eggs may be impacted.

In laboratory feeding studies, Philonthus adults were found to readily consume
corn rootworm (substituted for A. spretulus) eggs but were not as likely to eat A.
Spretulus larvae (Jo 2000). Because adult Staphylinidae are surface insects, they may not
come into contact with the eggs or larvae of scarab beetles very often. Philonthus larvae

should be investigated as potential predators of scarab larvae because of their high

22

density in home lawns and golf course roughs (Potter 1998). In laboratory studies
Philonthus larvae were the best predators of A. spretulus larvae (100% consumption) of
any potential predator tested, which included adult carabids A. aenea, Harpalus afiim’s
(Schrank), Stenopholus ochropezus (Say) and adult staphylinids Apocellus sphaericollis
(Say) Philonthus carbonarius (Gravenhorst) and Philonthus cognatus (Stephens). (J o
2000). i

The mites recovered form the heat extraction samples consisted of high numbers
of mites in the group Mesostigmata. A large portion of this group are considered
predatory on nematodes and other mites, while others feed exclusively on Collembola
(Wallwork 1976). This does not mean that they cannot be considered prey items, but that
they may also be important predators in the turf system.

In 2000 we added a category for Curculionidae, as they are widely studied as
pests of turfgrasses and we were finding them at our course in large numbers. The
curculionid was identified by Gary Parsons (MSU) as Sphenophorus minimas (Hart). S.
minimas is a small weevil closely related to the bluegrass billbug, S. parvalus (Gyllenhal)
(Vittum et a1 1999). Discovery of S. minimus at a high enough density to cause turf
injury in this study indicates a need to study billbug pests of turf more carefully in
Michigan, where S. parvalus was the only species of Sphenophorous previously believed
to be a pest.

However, it is not possible for us to extrapolate species diversity and richness
from this data, as not all individuals form the samples were identified. Because only five
individuals per family were kept for identification from each sample, information on the

total number of species inhabiting Groesbeck Golf Course can not be ascertained. For

23

example, although A. aenea made up the bulk of our samples, without more precise data
on how many of this particular species were caught, it is not clear whether A. aenea are
actually more abundant than other carabids, or if they simply are more likely to fall in the
pitfall traps.

Surface and subsurface sampling. In 1999, the insect distribution data collected
fiom one hole on Groesbeck golf course seemed consistent with the data from previous
studies of predatory arthropods on four other golf courses in Michigan (Smitley et a1
1998), which reported a sharp difference in predator abundance in the rough compared
with the adjacent fairway. In previous studies, samples were taken from 0.75 m to 10 m
from the turf border, whereas in this study, all sampling was done within 2.0 m of the
border, due to the desire to study a potential border effect. In 2000, staphylinids were
again more abundant in the rough, but carabids were slightly more abundant in the
fairway, differing from the distribution reported in the previous studies (Smitley et al.
1998). In both years, ants were more abundant in the fairway, which is the opposite of the
results from previous studies done at Franklin Hills Golf Course in Oakland County, MI
and Oakland Hills in Oakland County, MI in 1994 (Smitley et al. 1998).

Staphylinid activity was highly correlated with mite and Collembola abundance
for both years. The greater abundance of Collembola and mites in the rough compared
with the fairway supports our hypothesis that Staphylinids residing in the rough did not
need to travel to find food. If Collembola and mites are found in 3-fold more abundance
in the rough, and staphylinids are preying on them, then this might explain the high
correlation between them. A larger prey base may keep the predators from moving out of

the rough, even when high prey populations (such as outbreaks of pest species) are

24

present in the fairway. This is the first account of how the specific abundance of soil
Collembola and mites relates to the abundance of Staphylinids in the turf. Adult female
Staphylinids may select the rough for ovipostion over the fairway due to the larger
amount of potential prey for their larvae.

However, other environmental factors, such as soil compaction, mowing
frequency, soil nutrition and heat build-up may also play a role in the distribution of both
soil animals and surface predators. At Groesbeck, the fairway was mowed three times as
often as the rough, but was less likely to be driven on by golfers. Also, the fairway
experiences high temperatures during the day, and the risk of desiccation may be greater
in the fairway than the rough. There could be differences in evapotranspiration rates
between the fairway and rough. Evapotraspiration rates in turf are dependent on solar
radiation (Beard 1973) which may be higher in the fairway. Most likely, it is a
combination of factors which contributes to the high numbers of predators caught in the
rough compared to the fairway. More research is needed to establish this food chain
connection between staphylinids (Philonthus) found in turf and Collembola and mites.

Another concern of this study was the ability of the pitfall trapping to accurately
portray the abundance of surface predators in relation to the abundance of soil animals.
Pitfall traps are not a measure of abundance, but rather an indication of activity (Snider
and Snider 1986). And while most insects will fall into a pitfall trap, it is not known if
certain species are capable of avoiding this sampling method. Also, the vials used in this
study may not have been effective for all arthropods, especially large ones, such as the

carabid Scarites quadriceps, which is almost too large to fit in the mouth of the vial.

25

In 2000, Carabidae and Staphylinidae were found in much lower numbers. In
fact, abundance of all arthropod groups, surface and subsurface, was dramatically reduced
in 2000. Collembola and mites were found at about 1/4 of the numbers recovered in the
previous year. This may be due to unusually heavy precipitation levels in May and June
of 2000 (Fig 9) as well as a cooler spring, between May and June of 2000 (Fig 10).
However, accumulated degree-day totals (Fig 11) reveal that there were actually more
degree-days accumulated between May and June of 2000 than in the previous year,
therefore it is unlikely that the cool weather had an effect on the insect activity.

Despite the higher number of degree-days, it is still possible that wet weather
played a major role in arthropod activity. During several sampling trips, the turf was
soggy, and during sampling week number six, the plots were under water for 48 hours,
causing the pitfall traps to overflow. The vials measure 8.5cm deep, and were already
filled with over 4 cm of ethylene glycol. More than 4.0 cm rain in one week caused them
to overflow, and the trapped arthropods are washed away.

High levels of precipitation also affect soil samples. Mold can form on the
samples while in the heat extraction units, hindering the drying process and possibly
causing death. Also, arthropods from wet soils may not be as affected by the heat, and
therefore, do not move into the alcohol as quickly as those found in drier soils. Soil
arthropods such as Collembola and mites do not tolerate high moisture content very well
(Wallwork 1976). A study by Weis-Fogh (1948) showed a preference of 20 -30% water
content/dried soil (g) for most soil microfauna, with the exception of certain species of

Cryptostigmata. This is true for Carabidae as well, which decrease in abundance as soil

26

moisture increases from moist to saturated (Wallwork 1976). Thus, the wet weather may
have played a role in the low abundance of soil arthropods at Groesbeck in 2000.

Relatively few A. spretulus or A. granarius were found during the course of this
study. The adults were present in moderate levels in pitfall traps, but the larvae were
scarce. Groesbeck did not have turf damage from A. spretulus or A. granarius in 1999 or
2000. They have had damage from A. granarius in previous years. However, larvae of
Japanese beetle and European chafer caused turf damage in 1999 and 2000, mostly in the
roughs. Therefore, no clear information can be ascertained from Groesbeck regarding A.
spretulus and A. granarius, except that perhaps they are heavily preyed upon, and
therefore do not cause problematic turf damage.

Formicidae presented a particular problem for analysis, as they were highly
abundant in some samples, and absent in others. The resulting trend for ants in the
fairway may be misleading. Ants are highly aggregated, and using the soil sampling
methods employed in this study, it was possible to sample an entire colony of ants at a
single time, or miss them entirely. If the ant colony itself was missed, ants would be
absent from the sample after heat extraction. We did not intentionally choose sites that
contained high numbers of colonies. In fact, the colonies were not visible in the turf,
therefore, we didn’t know that we had sampled one until the soil was placed in the plastic
bag. In samples containing an entire ant colony, other types of arthropods were scarce.
Ants are notoriously protective of their colonies, and may actively prey on or remove
other arthropods residing near their colonies. Also, when using pitfall traps to capture
surface dwelling ants, their habit of laying down a scent trail causes the ants to be

captured in clumps, creating more variability. However, our efficacy test shows that,

27

while these methods of monitoring ants are highly variable, ants have an equal chance of
falling in a pitfall trap in the fairway or the rough.

Sampling for A. spretulus, A granarius, and P. japonica. Groesbeck Golf
Course does not currently have a problem with infestations of A. spretulus or A.
granarius larvae, on new #8 or any other fairway on the course (John Johnson, Groesbeck
Golf Course Superintendent, personal communication). In 2000, the presence of P.
japom'ca allowed us to examine the relationships between potential predators and P.
japonica larvae. P. japom'ca was found to be more abundant in the fairway, consistent
with the observations made by Rothwell and Smitley (1998) on A. spretulus larvae. More
P. japam'ca adults were caught in pitfall traps in the fairway than in the rough, either
because more adults emerged in the fairway, or adults are attracted to the fairway (Table
1). From our data we cannot tell if more P. japonica larvae were found in the fairway
because more eggs were deposited there or because more larvae survived there compared
with the rough.

Heat Extraction Efficiency. The results of the heat extraction efficiency test
show that the number of individuals obtained by heat extraction during this study were
significantly lower than the actual total number of animals present in the soil core. Thus,
it appears that the numbers of Collembola and Acari found per soil core during this study
were approximately only 50% of the true number of individuals in the soil (Table 8).
While the floatation method is more accurate, it is labor intensive and time consuming.
Heat extraction is a simpler and acceptable method for providing data on soil animals,

provided that a true estimate of the efficiency of the method is obtained (Snider and

28

Snider 1997). The efficiency of heat extraction of samples taken from the fairway
compared with those taken from the rough was not significantly different.

Mowing height regimes. The fairway at Groesbeck Golf Course was mowed to a
height of 15 mm 3 times per week. Previous research at Spring Lake Country Club in
Ottawa County, MI and the Cattails Golf Course in Oakland County, MI by Smitley et al.
(1998) was conducted on fairways mowed to heights between 10 and 16mm, and mowed
between 2 and 4 times per week. Populations of A. spretulus and A. granarius larvae
were higher at those courses than at Groesbeck. However, Staphylinids were slightly
more abundant at Groesbeck than at Cattails, and less abundant than at Spring Lake,
which were sampled in 1995 and 1996. This may indicate a slightly higher level of
potential predation by Staphylinids at Groesbeck, than at the Cattails. The abundance of
insects found at Groesbeck fell between the previous studies. There was not as large a
difference between the number of Staphylinids caught in pitfall traps in the fairway and
rough at Groesbeck as there was at Spring Lake. Groesbeck had larger differences
between fairway and rough than did the Cattails, although the Cattails had a similar
mowing regime. This suggests that the distribution of insects on golf course fairways and
roughs is affected by factors other than mowing height. This should be expected, as
frequent mowing translates into higher disturbance of the turf. A large mower could also
lead to increased soil compaction in the fairway, but golf cart activity in the rough may be
equally disturbing and have similar effects on soil compaction.

A limitation of this study is that the experiment was not replicated at other sites.
Because only one fairway and one surrounding rough were studied, it is not appropriate

to make generalizations about the distribution and abundance of soil animals on all golf

29

courses. Replicating this study at other field sites will reveal more information about the
dynamics of the arthropod food chain in turfgrass. This information will be helpful in
understanding the role of generalist insect predators as biocontrol agents for invasive pest

species, such as A. spretulus and A. granarius.

Most previous research done on turf arthropods focused on bionomics, control,
and treatment of arthropod pests (Potter 1998, Tashiro 1987, Vittum et al.1999). This
research, with a focus on examining and understanding the turf arthropods as an
ecological community, will hopefully prove helpful for further studies on the ecology turf
and the biocontrol of turf pests. However, further research into this system must also
include investigation of the micro site variability of the turf, and environmental
conditions such as heat, plant density, soil compaction and disturbance as other factors in
the distribution of predators in turfgrass, and consequently, the outbreaks of pest species

on golf course fairway.

30

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3
3

Table 4. Larvae recovered from soil cores taken from 25 May to 20 July
2000 at Groesbeck Golf Course in Lansing, Michigan

lnsect n Fairway Rog—gh‘

A. spretulus 24‘ 2.5 i 1.0 3.0 1 0.8
A. granan’us 24 1.2 1: 0.4 1.4 1 0.5
P. japonica 24 9.7 1; 1.0* 4.2 i 1.0

 

 

Data are the mean number of individuals found per 10 soil cores (0.1m2)1 SD.
Fairway means 1 SD followed by an asterisk are different from corresponding rough

means at the P = 0.05 level.
Zero A. granarius grubs were found in 1999. P. japonica were not sampled in 1999.

34

 

 

  
 
 

   

    
 

 

 

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36

Table 7. Data from soil moisture tests using the Aquaterr ® soil moisture
meter. Data are the means per ten probes per block per treatment over

8 sampling dates in 2000.

 

 

Date Fair Rough
17-Jul 84.9 i 1.2 82.8 i 1.9
24-Jul 86.7 i 2.3 87.4 i 1.6
31-Jul 81.9 i 1.7 81.5 i 2.0
84.5* 83.9*

 

* Cumulative means data for all 3 sampling dates

37

Table 8. Results of the heat extraction efficiency for selected arthropods
based on the total number of specimens obtained per treatment.
%Efficiency = [(N individuals obtained by heat extraction)! (N total
obtained by heat plus floatation)] x 100; F and R = Fairway and Rough.

 

 

@ecies n Treatment Heat extraction* Floatation“ % Efficiency
Collembolla 12 F 241 291 45.30

12 R 566 856 39.80
Mites 12 F 125 153 44.96

12 R 762 694 52.34
Others 12 F 72 49 59.50

12 R 104 55 65.41

 

* Number of individuals extracted by heat
“Number of individuals obtained from sugar floatation, after heat extraction.
n is the number of soil cores used in this experiment, data were pooled for each turf type.

Figure 1A. Experimental design on new #8 at Groesbeck Golf Course in Lansing, MI.

III

Rough

 

 

 

IV

 

Fairway ] V

 

 

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Rough

Figure 1B. Example of a single block of the experiment at Groesbeck Golf Course.

Fairway

2m 1m

Rough

 

. Soil Cores

EB Pitfall traps

39

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caught in the 2 adjacent pitfall traps at Gmbeck Golf Course in 1999. Equation for regression
and regression statistics can be found in Table 5.

Regression Plot
Dependent: staphlog
. I . ‘1

 

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.h.
I

 

 

 

 

2 2.2 2.4 2.6 2.8 3 3.2 3.4

45

 

Figure 8. Relationship between mites extracted from a soil core and the staphylinid adults caught
in the 2 adjacent pitfall traps at Groesbeck Golf Course in 1999. Equation for regression and
regression statistics can be found in Table 5.

Regression Plot
Dependent: staphlog

2.4 J I 4 I n n n I
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49

LITERATURE CITED
Abacus Concepts. 1991. SUPERANOVA statistical analysis program.

Altieri MA. 1991. The agro-ecology of temperate cereal fields: entomological
implications. From The ecology of temperate cereal fields. Oxford: Blackwell Scientific,

pp. 235-58.

Bauer T. 1979. 1979. The behavioral strategy used by irnago and larva of Notiophilus
biggatus F. (Coleoptera, Carabidae) in hunting Collembola. Misc. Pap. Agric. Univ.
Wageningen 18:133-42.

Beard J. 1973. Turfgrass: science and culture. Prentice Hall, Englewood Cliffs, NJ.
658pp.

Bernays EA. 1990. Plant secondary compounds deterrent but not toxic to the grass
specialist acridid Locusta migratoria: implications for the evolution of graminivory.
Entomol. Exp. AppI. 54:53-56.

Bernays EA, Barbehenn R. 1987. Nutritional ecology of grass foliage-chewing insects.
From Nutritional ecology of insects, mites, spider, and related invertebrates. New

York:Wiley, pp. 147-175.

Butcher J, Snider R, Snider RJ. 1971. Bioecology of edaphic Collembola and Acarina.
Ann. Rev. Entomol. 161249-88.

Christiansen K. 1964. Bionomic of Collembola. Ann. Rev. Entomol. 9: 147-78.
den Boer PJ, den Boer-Daajne W. 1990. On life history tactics in carabid beetles: are
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ground beetles in ecological and environmental studies. Intercept, Andover.

Hagley EA, Holliday NJ, Barber DR. 1982. Laboratory studies of the food preferences
of some orchard carabids (ColeopterazCarabidae). Canadian Entomologist. 114:431-37.

Harper JL. 1977. P0pulation biology of plants. London, Academic.

Henderson IF. 1978. Assessing the effects of invertebrates on grassland productivity.
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J o Y-K. 2000. Predation, soil moisture and mowing height as potential factors in the
skewed distribution of Ataenius spretulus (Coleoptera:Scarabaeidae) on golf course

50

 

 

fairways and roughs. M.S. Thesis, Dept. of Entomology, Michigan State Univ., East
Lansing, Mich.

Kegel, B. 1990. Diurnal activity of carabid beetles living on arable land. In N.E. Stork
[ed.], The role of ground beetles in ecological and environmental studies. Intercept,
Andover.

Lavigne RJ, Kumar R, Leetham J, Keith V. 1972. Population densities and biomasss
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Univ., Ft. Collins. 203 pp.

Leetham JW. Milchunas DG. 1985. The composition and distribution of soil
microarthropods in the shortgrass steppe in relation to soil water, root biomass, and
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Lovei G, Sunderland K. 1996. Ecology and behavior of ground beetle (Coleoptera:
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Niemczyk HD, Wegner GS. 1982. Life history and control of the black turfgrass
ataenius (Coleoptera-Scarabaeidae), pp. 113-117. In HD Niemczyk and BG Joyner (eds.),
Advances in turfgrass entomology. Hammer Graphics, Piqua, Ohio. 150 pp.

51

 

Pollet M, Desender K. 1987. Feeding ecology of grassland-inhabiting carabid beetles
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366.

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Smitley DR. 1994. Entomology Research. 64th Annual Michigan Turfgrass Conference
Proceedings. 18—20 January 1994, Lansing, MI.

Smitley DR, Davis TW, Rothwell NR. 1998. Spatial distribution of Ataenius spretulus,
Aphodius granarius (Coleoptera: Scarabaeidae) and predaceous insects across golf course
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Stanton N. 1988. The underground in grasslands. Ann. Rev. Ecol. Syst. 191573-89.

Tashiro, H. 1987. Turfgrass Insects of the United States and Canada. Cornell University
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Entomol. 40:535-58.

Villani M, Allee L, Diaz A, Robbins P. 1999. Adaptive strategies of edaphic arthropods.
Annu. Rev. Entomol. 44:233-56.

Vittum P, Villani M, Tashiro H. 1999. Turfgrass insects of the united States and
Canada. Cornell University Press, Ithica, NY. 422pp.

52

Wallwork J. 1970. Ecology of soil animals. McGraw-Hill Pub. Co. Ltd., Maidenhead,
Bekshire, Eng. 25 8pp.

Wallwork J. 1976. The distribution and diversity of soil fauna. Academic Press Inc.,
London, Eng. 355pp.

Weaver JE, Hacker JD. 1978. Bionomical observations and control of Atanius spretulus
in West Virginia. W.V. Univ. Agric. Exp. Stn. Curr. Rep. 72.

Wegner GS, N iemczyk HI). 1981. Bionomics and Phenology of Ataenius spretulus.
Ann. Entomol. Soc.Am. 74: 374-3 84.

Weis-Fogh T. 1948. Ecologocal investigations on mites and collemboles in the soil.
Natura Jutl. 1:135-270.

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New York Entomol. Soc. 40:77-93.

53

Appendix 1
Record of Deposition of Voucher Specimens“

The specimens listed on the following sheet(s) have been deposited in the named
museum(s) as samples of those species or other taxa, which were used in this research.

Voucher recognition labels bearing the Voucher No. have been attached or included in
fluid-preserved specimens.

Voucher No.: 2001-02
Title of thesis or dissertation (or other research projects):

EXAMINATION OF THE ARTHROPOD COMMUNITY ON A GOLF COURSE
FAIRWAY AND ROUGH

Museum(s) where deposited and abbreviations for table on following sheets:

Entomology Museum, Michigan State University (MSU)

Other Museums:

Investigator’s Name(s) (typed)
Breana Lee Simmons

 

 

Date 02/14/01
*Reference: Yoshimoto, C. M. 1978. Voucher Specimens for Entomology in North

America.
Bull. Entomol. Soc. Amer. 24: 141-42.

Deposit as follows:
Original: Include as Appendix 1 in ribbon copy of thesis or dissertation.

Copies: Include as Appendix 1 in copies of thesis or dissertation.

Museum(s) files.
Research project files.

This form is available from and the Voucher No. is assigned by the Curator, Michigan
State University Entomology Museum.

54

 

 

 

  

 

 

 

 

 

 

Appendix 1.1
Page 1 of 2 Pages

Voucher Specimen Data

 

 

 

 

 

 

 

 

 

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00.02.
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m M 10 0+ 8
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m .m .m W M M on” m. m m 0 00.00.60 208.0000 .0. 0.00 .0004 .
.00 000802

 

 

 

 

55

Appendix 1.1

Voucher Specimen Data

 

Pagej of 2 Pages

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