Town Ants: The Beginning 

of John Moser’s Remarkable 

Search for Knowledge

James P. Barnett, Douglas A. Streett, and Stacy R. Blomquist

United States Department of Agriculture

Forest Service

Research & Development

Southern Research Station

General Technical Report SRS-182
September 2013

Southern Research Station

200 W. T. Weaver Blvd.

Asheville, NC 28804

www.srs.fs.usda.gov

Cover: Town ants (Texas leaf-cutting ants), Atta texana (Buckley), 

clipping needles from a loblolly pine (Pinus taeda) seedling. 

Photo credits: Unless otherwise noted, the photos are from the 

collections of John Moser and the U.S. Forest Service.

Conversions: 1 ha = 2.47 acres and 1 cm = 0.4 inches

Pesticide Precautionary Statement

This publication reports research involving pesticides. It does not contain 

recommendations for their use, nor does it imply that the uses discussed 

here have been registered. All uses of pesticides must be registered by 

appropriate State and Federal agencies before they can be recommended. 

CAUTION: Pesticides can be injurious to humans, domestic animals, 

desirable plants, and fish or other wildlife—if they are not handled or 

applied properly. Use all pesticides selectively and carefully. Follow 

recommended practices for the disposal of surplus pesticides and pesticide 

Product Disclaimer

The use of trade or firm names in this publication is for reader information 

and does not imply endorsement by the U.S. Department of Agriculture of 

any product or service.

The Authors:

James P. Barnett, retired Chief Silviculturist and 

Emeritus Scientist; Douglas Streett, Supervisory 

Research Entomologist; and Stacy Blomquist, 

Biological Science Technician, USDA Forest Service, 

Southern Research Station, Pineville, LA 71360.
Town Ants: The Beginning 

of John Moser’s Remarkable 

Search for Knowledge

James P. Barnett, Douglas A. Streett, and Stacy R. Blomquist

Abstract

John C. Moser’s career spans over 50 years, and his research has focused 

on understanding the biology of town ants (Atta texana) and phoretic 

mites and other associates of ants and pine bark beetles. His approach to 

developing methods for the control of these pests has been to understand 

more completely the biology of these organisms. This research approach has 

established Moser as an internationally recognized expert in leaf-cutting ant 

biology and as a premier authority on phoretic mites and other associates 

of pine bark beetles. John’s efforts have led to the largest collection (over 

30,000 specimens) of mites associated with forest insects in the world. 

This knowledge is leading entomologists, ecologists, and pathologists to 

consider the possible role of mites in insect-fungal symbioses and disease 

transmission in trees.

Keywords: Atta texana, Dendroctonus frontalis, Dutch elm disease, fungal 

ascospores, mites and other pine bark beetle associates, southern pine 

beetles, southern pines, Texas leaf-cutting ants.
John C. Moser is a long-term scientist who has conducted research for the Forest Service, U.S. 

Department of Agriculture, Southern Research Station at the Alexandria Forestry Center in Pineville, 

LA for over 50 years. Retired for over 20 years, he continues to work daily and on most weekends as an 

emeritus scientist. He is noted for being somewhat eccentric, mainly because he studies mites carried 

on forest insects, an area that is generally considered out of the mainstream of forest entomological 

research. Few employees at the Forestry Center, or even the Southern Research Station’s scientific 

staff, understand and appreciate the work that Moser is conducting in entomological research and its 

significance to the international entomological community.

Although Moser’s research assignment related to the control of the town ant (Atta texana) was officially 

terminated in 1962, 4 years after he was hired by the Forest Service, by determination he nevertheless 

continued studies on the biology and ecology of the ants for decades and became an international 

authority on leaf-cutting ants. His work with mites associated with bark beetles has also gained him 

international recognition and respect and made a huge impact on this field of study. 

Many aspects of his research published in the most prestigious international scientific journals (Science

in the United States, Nature in the United Kingdom, and Naturwissenschaften in Germany) have 

enhanced his international recognition. His research, too, is becoming critical to understanding the 

biology of how major tree diseases and tree-killing beetles are transmitted. 

Moser’s accomplishments are even more significant in light of the administrative difficulties he has faced 

in pursuing his scientific interests—as they were usually outside the typical research emphasis of the 

research work units to which he was assigned.

John Moser’s long career, his accomplishments, and their significance need to be understood and 

appreciated. The objective of this publication is to document these research contributions.

—J.P. Barnett 

Photo of John C. Moser taken late in his career.

PREFACE
iii

Contents

Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .iii

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

The nature of John Moser’s education . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

The first job—Understanding town ant biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Discovery of new inquiline species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Town ant control treatments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Assignment to southern pine beetle research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Continuation of town ant research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Initiation of research on mites related to southern pine beetles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

Initiation of research with Chinese scientists . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

Retirement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

Research after retirement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

Excavation and mapping of a town ant colony . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

Interrelationships among beetles, mites, fungi, and nematodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

International collaboration on species identification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

Collaboration with Mexican scientists . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Role of mites in disease transmission . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

Fire ants and associated mites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

Hackberry gall makers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Implications for future research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

Significance of Moser’s contributions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

Literature cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

Scientific breakthroughs can occur through a number of 

approaches, such as by teams of scientists working together 

with common goals, by serendipity or happenstance, or 

sometimes by the dogged determination of a scientist 

working outside the mainstream of conventional science. 

The latter approach best fits the accomplishments of John 

C. Moser, research entomologist with the Forest Service, 

U.S. Department of Agriculture Southern Research Station.

John Conrad Moser, born in 1929, grew up near Columbus, 

OH, and was by his own admission a poor student, most 

likely because he found little that challenged him. Even 

as a child, John, also called “Bobby” by his parents, 

demonstrated his willingness to take a different path. At 

age four, he became distracted while waiting for his mother 

while she shopped, and left her to walk the 4 miles to their 

home. Fortunately, he met a stranger and asked, “My mother 

is lost. Can you show me how to cross High Street without 

getting hit by the machines?” He was escorted to the police 

station, and they soon located his mother. To this day, he 

states, “I could have made it home without any problem.”1

Another example of Moser’s independent thinking was 

during his junior year in high school when he and a friend, 

who both had joined the Civil Air Patrol during World War 

II to have something to do, decided that it would be fun to 

fly to the East Coast. Since his parents were out of town, he 

obtained permission from the high school principal for he 

1

Barnett, J.P.; Streett, D.A.; Blomquist, S.R. 2012. Oral history interview 

of March 7, 2012, with John C. Moser. Cassette tape. Pineville, LA: U.S. 

Department of Agriculture, Forest Service, Southern Research Station, 

Insects, Diseases, and Invasive Plants Research.

and his friend, Eddy Cooper, to be absent from school for 

a couple of days and went to Columbus’ Lockbourne Field. 

There they convinced a pilot of the renowned Tuskegee 

Air Men’s squadron of the merit of their visit to the East 

Coast. The 477th Bombardment Squadron was assigned to 

Lockbourne Field to be deactivated following the end of 

World War II, and the pilot from the squadron agreed to 

fly them to Patuxent Naval Air Station in Maryland in a 

B-25 bomber. After an overnight stay—and many efforts to 

answer the repeated question of “What are you guys doing 

here?”—they were flown back to Columbus the next day 

(see footnote 1).

Moser graduated from high school in 1947 with a less than 

spectacular scholastic record, and he had no desire to go 

to college (Moser 2000). There were no jobs, however, 

because of the number of veterans coming home from 

World War II who were seeking employment. So Moser 

entered Ohio State University (OSU), which was only 

three miles from his home. Since tuition was only $45.00 

per quarter, he could make his tuition by mowing lawns 

in the summer, even when making only 40 cents per hour 

(see footnote 1). Thus began the education of one of the 

most distinguished and internationally recognized forest 

entomologists of our time. 

The objective of this publication is to document the 

scientific accomplishments of John C. Moser. Equally 

important is to understand the vision, determination, and 

commitment required to accomplish this status in light of 

the administrative and scientific obstacles he faced.

A B-25 Mitchell bomber such as this was used to fly Moser and his high school classmate to Patuxent Naval Air Station in Maryland. 

(Photo from www.vg-photo.com)

INTRODUCTION
2

It was not until his sophomore year at OSU that Moser 

found an area of science that captured his interest—when 

he took his first course in entomology. Not only was 

it interesting, but he began to see a career that had job 

potential. In addition to the interaction with several OSU 

professors who began to shape his career, Moser discovered 

that the leaves of a hackberry tree (Celtis occidentalis) in 

his front yard had many kinds of interesting insect galls. 

He discussed this with Professor Ralph Davidson, his 

undergraduate advisor, who encouraged him to observe 

and take notes regarding the different species of insects.

Moser was particularly intrigued by a tiny, metallic green 

wasp that was flitting around and inserting its ovipositor 

into certain galls. He collected some of these wasps and sent 

them to a scientist at the U.S. Department of Agriculture, 

Agricultural Research Service Systematic Entomology 

Laboratory at Beltsville, MD, who identified the wasp as a 

new species of Torymus. This was really exciting to John 

because it meant that as a lowly sophomore student, “I had 

observed something that no one else in the world had ever 

seen” (Moser 2000). This was a pivotal event that launched 

Moser’s career as a research entomologist, as well as a 

motivation throughout his scientific endeavors. 

During the summers of 1949 and 1950, Moser worked 

driving and assisting OSU Professor C.H. Kennedy on his 

insect collecting trips to Upper Michigan and the north 

coast of Lake Superior in Canada, where they “chased 

ants” (Kennedy’s term) (Moser 2000). Moser states that 

he was far from the first choice for this job, but he was 

selected because Professor Kennedy was neurotic and 

other graduate students were afraid to work for him. 

Kennedy was difficult to work for, but he was a great 

teacher and Moser learned the basics of ant behavior and 

systematics that were to play an important role in his future 

job selection in the Forest Service. Kennedy told Moser, 

“If you ever get a chance to work on Atta, take it because 

this is where the real opportunities lie in myrmecology 

research” (Moser 2000).

Moser commented that in 1951, “much to the surprise of 

myself and everyone else, I received my B.S. in entomology 

from OSU. The year 1951 was the apex of the Korean War, 

and I was facing the draft. However, I was informed that my 

B.S. in entomology was worth a commission, the drawback 

being that I must enlist for a minimum of 4 years, and that 

I would be sent to Korea to participate in a pest control 

unit. The alternative was to be drafted, serve only 2 years, 

undergo 16 weeks of basic, but then be sent to work as a tech 

at a Department of Defense (DOD) research facility, and 

forever remain a private. I chose the latter” (Moser 2000).

After basic training, Moser was assigned to the Army 

Chemical Center at Edgewood Arsenal near Baltimore, 

MD, where he served as a technician at the Army Medical 

Corps, Insect Physiology Laboratory. He was associated 

with several other early mentors, namely Leigh E. 

Chadwick, Detrich H.F.A. Bodenstein who was elected to 

membership in the National Academy of Science in 1958, 

and Bertram Sacktor, who were three of the more important 

insect biochemists at that time. Moser was assigned to Dr. 

Sacktor, who included him as a junior author on one of his 

papers published in the 1953 Biological Bulletin entitled 

“Dephosphorylation of ATP by tissues of the American 

cockroach.” This was cutting edge science and John Moser’s 

first publication (Sacktor and others 1953).

Left: two types of insect galls that were found on the hackberry 

tree in Moser’s yard. The wasp that Moser first identified as a 

new species was associated with the inverted cone-shaped gall.

Above: this is the tiny green-metallic colored wasp that Moser 

found and was identified as a new species of Torymus.

THE NATURE OF JOHN MOSER’S EDUCATION
3

While attached to the Army Chemical Center, Dr. Chadwick 

allowed Moser to attend his first Entomological Society of 

America (ESA) meeting that was held in Philadelphia, PA. 

More important, though, were the three-day passes that 

Moser received each month. Normally he would visit New 

York City and Washington, DC on alternate months. While 

in New York City, he would attend Broadway shows and, 

as a member of the military, was allowed free admission to 

show dress rehearsals (see footnote 1). 

During the alternate month’s visit to Washington, DC, he 

normally spent time at the Smithsonian National Museum 

of Natural History, where he would visit with Barnard D. 

“Barney” Burks, who was the curator of chalcidoid wasps 

at the Systematic Entomology Laboratory of the U.S. 

Department of Agriculture, Agricultural Research Service. 

Burks helped Moser to describe his new species of Torymus, 

in addition to several other new species of parasitoids that 

Moser had found associated with the hackberry galls in 

Columbus, OH (Moser 2000). Burks strongly encouraged 

Moser to apply to Cornell University as a Ph.D. candidate. 

In September 1953, Moser was released from the Army and 

received the G.I. Bill, which financed the remainder of his 

graduate education. He returned to his home in Columbus 

and entered OSU, where in 1954 he completed his Master’s 

degree, majoring in biological control under Professor 

Alvah Peterson, noted author and one-time President of the 

Entomological Society of America. His thesis on hackberry 

galls and description of the parasitoid, Torymus vesiculus,

was summarized in the 1956 Journal of the Kansas 

Entomological Society (Moser 1956).

A year later, in September 1954, Moser entered Cornell 

University, majoring in insect ecology and studying 

under Howard Evans, who taught courses in taxonomy of 

Hymenoptera and the minor insect classification orders, 

and who was awarded the Daniel Giraud Elliot Medal 

from the National Academy of Sciences in 1976. In 

addition, Moser shared an apartment with John G. “Jack” 

Franclemont, who taught Lepidoptera systematics and later 

donated his extensive collection of 350,000 moths and 

extensive library to Cornell University. These two mentors 

helped shape Moser’s views on the meaning and values of 

systematic studies to the study of insect interrelationships. 

At Cornell, he continued his research on the natural 

enemies of hackberry gall makers. This research was 

published in 1965 as Bulletin 402 of the New York State 

Museum and Science Service (Moser 1965). Of Moser’s 

nearly 200 publications, this was his longest, at 95 pages.

John Moser’s education was strengthened and his 

career focused by the entomologists who served as his 

mentors—these were the outstanding entomologists of 

the 20th century. They included individuals from OSU, the 

Army Chemical Center, Smithsonian’s Natural History 

Museum, and Cornell University. Moser freely admits that 

his acceptance into Cornell University’s Graduate School 

would have been difficult based solely on his academic 

achievement, but the recommendations from his mentors 

based on his research capability overcame any such 

weakness (see footnote 1).

Several months before Moser received his Ph.D. in 1958, 

the Forest Service’s Southern Forest Experiment Station 

sent Jack Coyne to Cornell University to earn a Master’s 

degree. Today, Coyne is known by his pioneering work 

of saving and planting scions of loblolly pines that had 

survived southern pine beetle attack. Coyne informed 

Moser of a GS-4 position in the Station’s Timber 

Management Research unit at Alexandria, LA. This 

position had been created for a person to solve pine 

regeneration problems caused by the town ant, Atta texana. 

John also had an offer of a GS-9 position (then the going 

rate for an entry level Ph.D.) from the Forest Service’s 

Northeastern Forest Experiment Station to study predators 

and parasitoids of the Gypsy moth (Lymantria dispar). 

Moser says, “Professor Kennedy’s words rang in my ears, 

and I took the GS-4 position. I arrived in Alexandria, LA 

on July 1, 1958, and was the first Ph.D. entomologist hired 

by the Southern Forest Experiment Station” (Moser 2000).

Right: publication 

documenting Moser’s Ph.D. 

research on gall-making 

insects. Below: another gall 

form found on the hackberry 

tree that was associated with a 

different parasitoid wasp.
4

When John Moser arrived in Alexandria, LA and reported 

to work for the Southern Forest Experiment Station’s 

Timber Management Research unit, he was given the 

assignment to “eradicate” the town ant, a project that 

“should last a couple of years.” Native of western Louisiana 

and eastern Texas, the town ant can be a significant 

problem in reforestation by causing pine seedling 

mortality that results from the clipping of the needles and 

terminal buds of seedlings. This material is carried into 

underground chambers, where it forms the fungal substrate 

each colony cultures for food. 

William F. “Bill” Mann, Jr. was the supervisor of 

the research unit and was noted for his 

authoritarian management style that often 

resulted in personal conflicts. Soon there was 

disagreement on the approach of the proposed 

research. Mann wanted Moser to perfect 

methods of killing the ants—the state-of-theart treatment at the time was to gas the colony 

with methyl bromide (Johnson 1944). Moser 

convinced Mann that real control could not 

be accomplished without knowing something 

about the biology of the insect.

In September 1958, Moser gave a seminar 

at the Louisiana Entomological Society 

meeting at Louisiana State University (LSU). 

There he met Murray S. Blum, an expert in 

insect chemical ecology, of the university’s 

entomology faculty and they collaborated for 

several years on the chemical ecology of town 

ant pheromones and the ants’ mating flights.

Mann’s insistence on a short-term effort 

to control the ant colonies conflicted with 

Moser’s long-term goal to develop a better 

understanding of the ant’s biology, and led to 

personal conflicts. Despite this, Moser soldiered 

on, collaborating with elite scientists, often 

in defiance of his project leader. One such 

collaboration involved Edward O. Wilson, a 

renowned ant biologist and winner of many 

international awards, including two Pulitzer 

Prizes. Wilson was interested in the chemical 

ecology and biology studies that Moser and 

Blum were conducting. During the 4 years that 

Moser was assigned to the Timber Management 

Research unit, Wilson visited Alexandria twice. 

Both times Moser brought his esteemed guest in 

through the back door to avoid paying a “verbal 

tribute” to the project leader (Moser 2000).

Town ants vary in size, ranging from 1/8- to 3⁄4-inch long. 

The queens, the largest, are rarely seen because they spend 

their entire lives underground, except for one short mating 

flight. Medium-sized worker ants do most of the foraging.

The worker ants cut and bring green leaf material into the 

nests for the “garden” on which a fungus grows—the only 

known food of the ant. Many smaller workers constantly 

care for the gardens, keeping them free of contaminating 

organisms. The need to understand what all went on 

belowground led to one of the most notable, and logistically 

and visually impressive, aspects of Moser’s town ant 

research, the excavation and mapping of a large colony. A 

Left: loblolly pine (Pinus taeda) seedling defoliated by town ants. Top right:

Continental Can Company provided men and equipment to help excavate and 

map the colony. Bottom right: structures of small, new nests were studied by 

filling them with watery cement and digging around the cast. In this photograph 

John Moser is observing the cemented nest which had a single gallery extending 

12 feet into the ground and had three small cavities; 

THE FIRST JOB—UNDERSTANDING TOWN ANT BIOLOGY
5

colony was excavated in 1960 using a bulldozer that cut a 

swath through it 25-feet wide, 100-feet long, and 12-feet 

deep (Moser 1963). Fungus-garden cavities were the heart 

of the nest and averaged a foot in diameter. Occasionally, 

they are much larger and a person can actually stand 

within the cavity. They may be as much as 8 feet below the 

surface and are connected by a series of tunnels used for 

movement to the surface. Material depleted of nutrients is 

deposited into dump or detritus cavities. Other cavities are 

packed with material such as ants, eggs, larvae, pupae, and 

sometimes green leaves or sand (Moser 1962). The function 

of these “dormancy” cavities is not clear.

This project, accomplished with significant support from 

forest industry, provided significant data on the functioning 

of the ants within the colony and resulted in a number 

of important publications. One of the first of these was 

“Probing the secrets of the town ant” (Moser 1962). 

Moser (1960) was also quick to defend other similar native 

ants, such as the harvester ant (Pogonomyrmex comanche) 

that does little damage to agricultural crops.

Discovery of New Inquiline Species

In the ant colonies, several undescribed specimens of 

inquiline insects (those who live in the nests of others 

without harm to them) were discovered by Moser. One of 

these insects described by Spangler (1962) was a minute 

scavenger beetle (1.3 mm long and 0.8 mm wide) that he 

named Oosternum attacomis. It is found in the detritus 

cavities of town ant nests. 

Another species found underground in the nests of the town 

ant is a fly in the lesser housefly genera (Fannia). Fly larvae 

scavenge in the detritus cavities of the ant colony. This species 

appears to move freely as adults through the nests of the ant 

below the ground surface (Chillcott 1965). Once the adults 

fully develop, they walk with wings unexpanded through 

tunnels to the surface where they rest until their wings 

expand. They then fly off to swarm, mate, and forage for 

food. Mature females return to the colony’s detritus cavities 

to lay eggs for another generation. This species was named 

Fannia moseri, in honor of John Moser (Chillcott 1965). 

Another beetle found in the nests is of the Euparixia

genera—these are scarab beetles that live as inquilines 

in detritus cavities. It was named Euparixia moseri by 

Woodruff and Cartwright (1967), also in honor of Moser.

Above: fungus garden cavity within a town ant nest. 

Right: this small beetle (Oosternum attacomis) (1.3 mm by 

0.8 mm) discovered by Moser lives in town ant detritus cavities.
6

Town Ant Control Treatments

Moser worked to refine and improve the methyl bromide 

treatments, but the excavation demonstrated that it is difficult 

to kill the entire colony by methyl bromide (Neelands 

1959). The effort to control the ant was expanded by the 

introduction and evaluation of a new chemical, Mirex®

(Bennett 1958). However, the application of Mirex® as 

a control agent was not finalized until after Moser was 

transferred from the Timber Management Research unit to a 

newly established Forest Insect Research unit.

With the transfer of Moser to the Forest Insect Research 

unit, Bill Mann recruited another entomologist to continue 

the work on the control of town ants. This scientist, 

William Echols, evaluated formulations of Mirex® designed 

especially for town ant control (Echols 1966). However, the 

development and widespread aerial application of a fire ant 

(Solenopsis invicta) formulation across the rural South by 

the U.S. Department of Agriculture significantly reduced 

the number of town ant colonies, and that line of research 

was terminated. 

Because of environmental problems, aerial applications 

of Mirex® were ended in 1975. Since Mixex® was more 

effective in eliminating native ant species than the 

nonnative fire ants, populations of fire ants quickly 

recovered, but it has taken decades for the town ants 

to begin reestablishing their earlier niche in the forest 

ecosystem (Barnett and others 2011). 

Moser’s research over a 4-year period with the Timber 

Management Research unit established his credentials as an 

expert in town ant biology and led to a career-long interest 

in its ecology and that of its interrelated species.

Injecting methyl bromide gas into a town ant colony was a longestablished treatment.
7

In 1962, an insect research project was established at 

Pineville under the leadership of William H. “Bill” 

Bennett, giving Moser an opportunity to transfer to a 

project much more to his liking. Southern Station Assistant 

Director L.W. “Les” Orr, who knew that Moser had a 

penchant for working with “small stuff”, asked if he 

wanted to work on the mites associated with bark beetles. 

This was an opportunity that Moser quickly embraced. Orr 

had a reason for suggesting the research on mites because 

he co-authored a paper with Leach and Christensen (Leach 

and others 1934) establishing that mites riding on Ips pini

fed on and disseminated the blue stain fungus in red pine 

(Pinus resinosa) (Moser 2000).

Les Orr wanted to know the identities of these mites and 

if the same thing was going on with southern pine beetles 

(SPB) (Dendroctonus frontalis), and he thought that Moser 

could find out all the information needed in about six 

months. Again, Moser encountered the short-term research 

attitude that conflicted with his long-term view requiring 

more thorough research to fully understand insect ecology.

Continuation of Town Ant Research

Fortunately, Moser was able to continue publishing earlier 

research related to the ecology of the town ant because he 

had collected significant data related to trail marking and 

mating flights that needed publication. 

Trail-marking substances—The work of Moser with 

Murray S. Blum of LSU’s Department of Entomology on 

trail-marking substances of the town ant created a lot of 

interest. An article published in Science established that 

potent chemicals produced in the ant’s poison gland are 

distributed by the stinger, which is not used for stinging 

(Moser and Blum 1963). Instead, this ant’s stinging 

defense mechanism has been adapted for the purpose 

of communication, and its powerful mandibles provide 

defense by biting (Hermann and others 1970).

The adapted stinging mechanism provided a chance for 

Moser to pull a prank on Bill Mann, his supervisor. One 

day when Mann and Moser were in the field observing town 

ants, they happened upon a huge population of harvester ants 

(Pogonomyrmex comanche) actively scurrying all around. 

Mann asked Moser, “Do they bite?” remembering how 

vicious a bite a town ant can inflict. Moser replied “no” and 

placed a few harvester ants in Mann’s hand. The harvester 

ants proceeded to sting Mann resulting in substantial pain. 

Mann responded, “You told me they wouldn’t bite!” Moser’s 

response was, “They don’t bite. They sting.” This did little to 

improve their strained relationship (see footnote 1). 

Studies were undertaken to determine if the trail 

substances of similar species were also produced in 

the poison gland. The results demonstrated that stingassociated glands were similar in closely related genera 

and species (Blum and others 1964). Related research 

showed that the inquiline roach (Attaphila fungicola), 

which inhabits fungus gardens of the town ant, also follows 

the ant’s odor trails (Moser 1964) and has been observed 

outside the nest, traveling along the trail used by foraging 

ant workers. Since this roach is wingless, it may use the 

odor trail to move from one nest to another (Moser 1967a).

Nest associates and enemies—In addition to the roach 

mentioned above, there are many species of insects 

associated with the town ant. Most of these are found in 

detritus cavities. Walter and others (1938) listed 44 species 

of arthropods found in the nests located in Texas, and 

Moser has found at least that many in Louisiana. He has 

identified and described many of these species associated 

with the town ant (Krantz and Moser 2012; Moser 1962, 

1963, 1967b; Waller and Moser 1990). 

Top: town ant biting the end of Moser’s finger. Bottom: worker 

town ant (left) with a winged queen ant showing the differences 

in size.

ASSIGNMENT TO SOUTHERN PINE BEETLE RESEARCH
8

Moser and Neff (1971) described a small fly (Pholeomyia 

comans) that is an associate of the town ant and lays eggs in 

the detritus cavities of nests. Three species of mites that are 

carried (phoretic) on the flies were identified and discussed.

Many genera of insects can be found associated with the 

ant nests. These include silverfish (Thysanura), small 

cockroaches (Blattodea), crickets (Orthoptera), cydnid bugs 

(Hemiptera), beetles (Coleoptera), flies (Diptera), and wasps 

(Hymenoptera) (Waller and Moser 1990).

In addition, at least 75 species of mites are associated with 

the town ant. The mites are phoretic and depend upon 

dispersal by riding on the winged ants during their mating 

flights (Moser 1983). 

Although birds and bats consume large quantities of the 

thousands of winged reproductive ants (alates) that take 

off from the nests (Warter and others 1962), the number of 

predators are limited. Minute flies (Myrmosicarius texana) 

(about 1 to 2 mm body length) hover over the ants at nest 

entrances and along trails and dart down to land on the 

ants’ heads, suggesting parasitism (Waller and Moser 1990). 

There is, however, little documentation of any significant 

adverse effects from any of the many associated species. 

Mating activities—Moser spent many years studying 

the mating flights of town ants and has documented many 

aspects of the environmental and physiological conditions 

when the flights occur (Moser 1967b). Winged ants develop 

in early February and March, but swarming generally does 

not occur until late April or May. The sex ratio is about 1:1. 

Before swarming will occur, nest surfaces must be soaked 

from rain, and flights will continue as long as the nests 

remain wet and temperatures exceed 16 oC. A circadian 

rhythm seems to trigger swarming at about 3:55 a.m., and 

most leave the nest surface within 2 minutes (Moser 1967b).

The flights take place in darkness, and little is known of 

the fate of the alates once they leave the nest. After mating, 

surviving females land, lose their wings, begin digging 

tunnels to establish new nests, and become the queens of 

new colonies (Moser 1983). Moser and Lewis (1981) found 

that some new colonies expand quickly because more than 

one queen is involved in its establishment. 

Pheromone studies—In 1966, David L. Wood of the 

University of California, Berkeley, an authority on western 

pine beetle (Dendroctonus ponderosae) ecology, visited 

Pineville to explain advances in chemical ecology of that 

species. He was fascinated by Moser’s chemical ecology 

studies of the town ant and suggested that he contact 

Robert M. “Milt” Silverstein of the State University of 

New York, for possible help in identifying the ant trail 

pheromone. Silverstein, an expert on the chemistry of 

insect communication and later in his career a member of 

the National Academy of Sciences, was eager to work on 

the town ant pheromone, but financial support was needed 

for the project (Moser 2000). 

The Forest Service was reluctant to continue research on 

the town ant because Mirex® had adequately controlled 

this and other leaf-cutting ants. Silverstein applied to 

the National Institute of Health and obtained a grant for 

$60,000 to conduct the research, but he had to visit the 

Southern Station in New Orleans, LA to convince them 

that Moser’s research should continue. They reluctantly 

agreed to the project. 

In general, pheromones trigger a number of functions such 

as mating, alarm, attraction, repulsion, and trail following 

(Moser 1970). The challenges are to identify chemicals 

eliciting response and determine their function. 

Moser’s initial task for this project was to collect 8 

pounds of town ants. He paid locals to collect the ants in 

coffee cans, paying them by the pound. Once collected, 

Silverstein and James H. Tumlinson (Silverstein’s postdoctorate graduate student now Ralph O. Mumma 

Professsor of Entomology, Penn State University) ground 

up and extracted 150 μg of pure trail substance. This and 

about 100 other allied compounds extracted from the ant 

poison sacs were sent to Moser. 

His task then was to prepare artificial trails from these 

compounds and let worker ants indicate which substance 

was the most active. This turned out to be a pyrrole (a 

heterocyclic aromatic organic compound) that was named 

“Attalure.” Attalure was the first pheromone demonstrated 

to elicit trail following for any ant (Tumlinson and others 

A soil chimney created by new workers about 2 months after the 

mating flight.
9

1971). Unlike most pheromones, which are highly volatile, 

this compound was a solid of low volatility. Low volatility 

is needed because the ant requires long-term sources of 

food that necessitate repeated use of trails. This substance 

is very powerful; only 330 μg is required to draw a 

detectable trail around the world (Moser 2000). These 

results were published in two papers in Nature (Moser and 

Silverstein 1967, Tumlinson and others 1971).

In a follow-up study, the major component of the volatile 

trail-marking pheromone was identified as methyl 

4-methylpyrrole-2-carboxylate. Another component also 

establishes a trail-following response. The purpose of this 

second substance was unknown, but it was proposed to 

allow insects of closely related species the capability to 

communicate (Tumlinson and others 1972). Sonnet and 

Moser (1972, 1973) prepared and bioassayed a number of 

synthetic analogs of the pheromones and determined their 

sensitivity by ant responsiveness. 

Additional tests demonstrated that 24 species of leaf-cutting 

ants followed the trail pheromone of the town ant. However, 

when ants of other species were given a choice of following 

one of two separate trails, they chose the trail of the most 

closely related species (Robinson and others 1974).

After the initial research on the trail pheromone, Moser 

suggested that efforts should begin to understand the 

volatile alarm-causing components of the head, specifically 

of the mandibular glands of the worker ants. Two 

components were identified that alarmed and repelled 

worker ants: 4-methyl-3-heptanone and 2-heptanone 

(Moser and others 1968).

The capability to split the 4-methyl-3-heptanone 

pheromone into two volatile components allowed 

Moser to bioassay which of these alarmed the ant. 

Although this procedure is now routine, it never had 

been accomplished for any bioactive substance, such as 

a pheromone. The bioassay was a breakthrough because 

it was the first procedure to allow concentrations of 

volatiles to be determined within minutes (Moser 1983). 

Silverstein’s graduate student, Robert G. Riley, isolated 

the two substances and sent them to Moser. The bioassay 

established that only the (+) enantiomer of the mirrorimaged compound was active. In 1974, these findings were 

published in both Science (Riley and others 1974a) and the 

Journal of Insect Physiology (Riley and others 1974b).

Andryszak and others (1990) determined antennal 

olfactory response of town ants to the trail and alarm 

pheromones. The results established the queen and male 

ants responded to lower levels of pheromones, while minor 

workers in the nest required higher levels to show response. 

Timber losses from a town ant colony—Although 

biologists and foresters have frequently documented the 

adverse effects of town ants on agricultural crops and small 

trees, the economic impact of this defoliation has seldom 

been determined. Moser (1986) used aerial photographs to 

locate and determine the size of a colony active in 1955 and 

made on-ground measurements 30 years later to estimate 

loss in tree stumpage value.

The colony occupied about 3⁄4 acre, and based on 1955 

dollars the stumpage value loss was $653. Cost of 

controlling the ants would have yielded a 15 percent rate of 

return during the 30-year period (Moser 1986).

The U.S. Department of Agriculture’s aerial application of 

Mirex® during the 1960s and early 1970s for fire ant control 

greatly reduced the number of town ant colonies, and it 

was not until the early 1980s that populations of ants again 

began to present a problem to reforestation (Weatherby 

cited in Moser 1983). 

Top: enlarged harvester ant showing the abdomen with the 

poison sac and stinger. Bottom: town ant mound with several 

entrances into tunnels that lead to underground cavities.
10

As the publication of much of the town ant research neared 

completion, Moser began to focus his research efforts on 

issues related to SPB.

Initiation of Research on Mites Related to 

Southern Pine Beetles

Moser was aided in his research on the mites associated 

with southern pine beetles (SPB) by the assistance of 

very talented technical support personnel. His early and 

long-term assistant, Lawrence M. “Lary” Roton, not 

only supported Moser’s work on the town ant, but led the 

effort to find and process the different species of mites 

on SPB. While Moser remained heavily involved in town 

ant studies, he directed Roton “to learn all you can about 

mites” (see footnote 1). Roton led the effort to process 

many of over 20,000 slides of mites in support of Moser’s 

assignment in 1963 to study mites. This effort provided 

answers to the questions of Assistant Director Les Orr 

about the array of mite species associated with SPB and the 

possible transmittal of the blue stain fungus (Ophiostoma 

minus). The identification of these mites was the issue that 

resulted in Moser beginning mite research.

Early in SPB studies, more than 90 species of insects were 

identified from bolts taken from loblolly pines (Pinus 

taeda) infested by SPB and Ips engraver beetles (Moser and 

others 1971). Many of these species were mites and became 

the object of Moser’s research. 

The number of new species found was so great that many 

of the slides processed by Roton and Moser were often sent 

to mite taxonomists active in the United States and Europe 

for identification. Included were specialists Bob Smiley and 

Ed Baker, U.S. Department of Agriculture Agricultural 

Research Service, Beltsville, MD; Jay Woodring and 

Bruce Boudreaux, Louisiana State University, Baton 

Rouge, LA; Evert Lindquist, Agriculture and Agri-food 

Canada, Ontario, Canada; Preston Hunter, University 

of Georgia, Athens, GA; Earle Cross, University of 

Alabama, Tuscaloosa, AL; Sandor Mahunka, Hungarian 

Natural History Museum, Budapest, Hungary; Jerry 

Krantz, Oregon State University, Corvallis, OR; and Jerzy 

Wiśniewski, Academy of Agriculture, Poznań, Poland. 

These identifications resulted in a flood of descriptive 

papers, many of which Moser co-authored with comments 

on the biology of the various new species. Many of these 

cooperators have since retired or deceased, leaving no one 

to replace their taxonomic expertise (Moser 2000).

To assist in technical training as well as provide quality 

control in slide processing, Moser established a rating 

system to evaluate the quality of mite slide development. 

After mites are mounted on slides by supporting 

technicians, the quality of the processing is evaluated and 

a ranking is noted on the slide. The ranking criteria are: 

very good (vg), good (g), fair (f), and poor (p). Soon any 

technician who assisted Moser worked diligently to achieve 

and maintain the highest ranking possible (see footnote 1). 

Moser developed the broad perspective that research on 

mites was an opportunity to understand their role in the 

transmittal of the blue stain and other fungal diseases, and 

to determine if mite species could provide a biological 

control of SPB. He found certain species that fed on the 

brood (i.e., pupae, larvae, and eggs) of SPB and suggested 

that they might be reared in sufficient quantities to slow the 

expansion of SPB populations.

Mites on SPB related to the blue stain fungus—The first 

paper that listed the array of mite species associated with 

the transfer of blue stain fungus by pine bark beetles was 

published in 1971 (Moser and Roton 1971). There were, 

Top: the blue stain fungus (Ophiostoma minus) is transmitted to 

southern pines by southern pine beetles and the mites they carry.

Bottom: Pyemotes barbara was found to be parasitic on the 

Douglas fir cone moth. Moser and Roton experimented with 

Pyemotes species on SPB to see if they would provide good 

biological control.
11

however, a number of earlier publications that documented 

some species of mites associated with the bark beetles 

(Hunter and Moser 1968, Smiley and Moser 1968, Smiley 

and Moser 1970, Woodring and Moser 1970). 

Moser (1976a) documented that SPB can carry a large 

number of phoretic mites. Based on the 20,000 slides 

Roton and Moser had processed since 1963, it appeared 

the tarsonemid mites were the primary players associated 

with blue stain (Moser 2000). In a related effort, Kinn 

(1976) then developed a key that helped identify the mites 

associated with SPB.

Ophiostoma minus is the blue stain fungus associated with 

SPB, and is introduced into trees by the attacking beetles 

(Bridges and Moser 1983). This fungus is a major factor 

causing mortality and/or degradation of wood quality. 

Colonization of the tree by the fungus is thought to aid 

development of the beetle in the tree by reducing the 

moisture content of the wood (Nelson 1934). The fungus 

can be isolated from the body of the bark beetle, but is not 

transported in the mycangium (cup-shaped pits in the thorax) 

as are some other symbiotic fungi (Barras and Perry 1972). 

Although the fungus was known to be transmitted by 

the beetle, there was no understanding of how the beetle 

becomes inoculated with fungal spores. The work by 

Bridges and Moser (1983) established that a possible source 

of inoculum was phoretic mites. Tarsonemid mites feed 

on the fungus and move from areas of fungus-colonized 

bark to the cracks in the outer bark where they attach 

to emerging mature adult beetles (Roton 1978). The 

association of mites carried by beetles and infestation of 

pines by the blue stain fungus provides an example of a 

three-way mutualism among insects, mites, and fungi 

(Bridges and Moser 1983).

Although beetles were known to carry mites, no specific 

information was available on where the mites were 

attached to the beetles. Research by Moser and Bridges 

(1986) determined that two species of Tarsonemus mites 

attach to different locations on the adult beetle, for one the 

location is beneath the thorax and for the other beneath the 

outer wings, but both species are capable of infesting trees 

with the fungus. 

Moser (1985) determined that certain female Tarsonemus

mites, like their host beetles, carry a specific fungus on 

which the mites feed inside special morphological, sporecarrying, sac-like structures called sporothecae. The 

sporothecae is similar to the mycangium described for bark 

beetles, but differs in that no gland cells are present. In 

one study, it was determined that over 85 percent of mites 

carried blue stain fungal spores, and each mite carried an 

average of 15 spores (Moser and Bridges 1986).

Bridges and Moser (1983) and Moser and Bridges (1983) 

found that beetles transporting tarsonemid mites carried O. 

minus disease more often than mite-free beetles. That study 

did not, however, compare mite population levels with the 

occurrence of blue stain in SPB infestations. Another study 

showed a significant relationship between the occurrence 

of phoretic tarsonemid mites and blue stain in SPB 

populations; the lowest incidences of mites and blue stain 

were associated with outbreak populations of SPB (Bridges 

and Moser 1986). Therefore, changes in mite populations 

could cause changes in blue stain abundance. 

Top: Tarsonemus mite slide showing fungal spores being carried 

in the mite’s sporotheca. Right: bucket trap for SPB with a 

synthetic pheromone bait.
12

Improvements in methodology—As late as the mid-1970s, 

there was no efficient way to trap live SPB for population 

studies. Sticky traps were still in common use (Moser 

1976b), but they were messy and trapped many nontarget 

organisms. Moreover, beetles became mired in the sticky 

material and were of little use for chemical investigations.

Moser and Browne (1978) devised a bucket trap baited with 

a synthetic pheromone which attracted large numbers of 

male beetles. When infected bolts were used as bait, roughly 

equal numbers of males and females were captured.

Another improvement in methodology was in the tagging 

of mites with paint for studies of Pyemotes females in 

life-span determinations. Moser and Roton (1970) found 

that aerosol paint sprayed at a distance of three feet 

satisfactorily marked the hairs on the mites’ bodies without 

overwhelming the mites with paint and made identification 

of live stages possible under a microscope.

The study of mite biology was difficult due to the small 

size of the mites. Lary Roton, Moser’s technician who was 

very creative, was asked to make a movie of the life of a 

mite; but Kodak®, the premier maker of camera equipment 

at the time, said that the technology did not exist to make 

such a movie. Roton located an old 16-mm movie camera, 

developed a connecting ring to a microscope, and filmed 

the life cycle of mites.2 Roton’s ingenuity underscores how 

Moser instilled into his technicians a desire to meet the 

scientific needs of the program regardless of the difficulty.

Obtaining a continuous supply of beetles for research 

purposes was difficult. So, Bridges and Moser (1984) 

developed a continuous mass-rearing technique for SPB. 

Such a colony was obtained from a natural infestation, and 

bolts from trees were cut and placed in a screened corner 

of a laboratory. New bolts were added periodically and the 

beetles moved into the uninfested bolts. Such a colony was 

maintained for over a year.

To observe the behavior and development of beetles in the 

pine’s phloem layer, phloem sandwiches were developed, 

in which a piece of bark and phloem is removed from a live 

tree and pressed between glass plates. Previous designs 

required that the insects be placed into this sandwich. 

Taylor and others (1992) modified the sandwich to allow 

insects to freely and naturally colonize the phloem 

sandwich by adding a piece of plywood with holes drilled 

through it to the bark side. This modification allowed 

natural infestations by SPB and associates. 

2

Barnett, J.P.; Streett, D.A.; Blomquist, S.R. 2012. Oral history interview 

of March 22, 2012, with Lawrence M. Roton. Cassette tape. Pineville, 

LA: U.S. Department of Agriculture Forest Service, Southern Research 

Station, Insects, Diseases, and Invasive Plants Research.

Beginning of research on Mexican beetles—In the 

early 1970s, Jack Coyne, Bill Rose, Ed Clark, and Robert 

Wilkinson sent Moser mite material collected from bark 

beetles in Mexico, Honduras, and Guatemala. In 1974, the 

results from these evaluations were published in Turrialba

as the first list of mites associated with pine bark beetles in 

Central America (Moser and others 1974). This information 

would provide rationale for preparing and submitting, in 

1997, a grant proposal with Jorge Macías to the Forest 

Service to conduct a survey of mites and insect associates 

of SPB in Mexico and Central America.

Temperature thresholds for SPB flight—An ongoing 

need in forestry is to be able to predict SPB activity. The 

primary approach to anticipate populations is the use of 

aerial detection. This survey method is good to quantify 

beetle population changes over large areas, but methods for 

determining beetle population changes over short periods 

and small areas are needed. Moser and Dell (1979) found 

that when SPB brood emerges, the populations of beetles 

that reach their destination and infect other trees increase 

as temperature increases and decrease with heavy rain 

during their period of flight.

Moser and Thompson (1986) plotted the range of flight 

temperatures for SPB and found the optimum flight 

temperature was about 27 oC. Observed minimum and 

maximum flight temperatures were 6.7 oC and 36.7 oC, 

respectively. The minimum temperature was a surprise 

because it is the lowest ever recorded for any flying beetle. 

Unlike its western cousins, SPB is active all winter and 

often flies at low temperatures. Flight temperature data are 

basic to the understanding of bark beetle dispersal, and this 

understanding is necessary for predicting and managing 

SPB epidemics.

Weather factors were also evaluated in relation to 

predicting flying populations of the SPB predator clerid 

Collecting adult SPB under the bark of Pinus oocarpa in Chiapas, 

Mexico.
13

beetle (Thanasimus dubius) (Moser and Dell 1980). 

Population trends of the flying clerid beetles were similar 

to those of SPB. Variations in ratios of predator to prey 

suggest that in endemic situations, T. dubius may suppress 

SPB populations.

Pyemotid mites associated with bark beetles—Moser, 

with Earl Cross of Northwestern State University 

at Natchitoches, LA and later of the Department of 

Entomology at the University of Alabama, published a 

number of studies on the systematics and biology of the 

pyemotid mites associated with bark beetles (Cross and 

Moser 1971, Moser and others 1971, Moser and Roton 

1972). This group of mites is important because some of the 

species are parasitoids, while other related species consume 

and may transmit the mycangial fungi of bark beetles 

(Moser 2000). 

Among the numerous mite species found in galleries 

of North American bark beetles are those of the genus 

Pyemotes. These mites are insect parasites that exhibit a 

wide variety of trophic diversity (i.e., they feed on a variety 

of things) as well as dispersal polymorphisms (i.e., they 

assume different forms). The genus naturally divides into 

two different subgroups. In the scolyti subgroup, all five 

species are phoretic on one or more species of bark beetles. 

In nature, all feed only on immature bark beetle broods 

(eggs, larvae, and pupae), but not on adults (Moser and 

others 1987). At least one female morph of each of the five 

species is a phoretomorph—a mite specialized with claws 

or suckers for clinging to insects (Moser and Cross 1975)—

and one expresses an extreme form of polymorphism 

known for mites (Cross and Moser 1975, Cross and others 

1981, Smiley and Moser 1984a).

In the ventricosus subgroup, its ten species are not known 

to be phoretic, but at least two species possess venom. This 

subgroup includes the straw itch mites, Pyemotes tritici, 

which produce a toxic rash on human skin, whereas P. 

scolyti and P. parviscolyti caused no observable symptoms 

(Moser 1975a). To bioassay toxicity to humans, Lary Roton 

volunteered to have female mites that had been feeding 

on SPB placed under a plastic cap taped to his arm. After 

four days the cap was removed and the arm appraised for 

itching, redness, and blisters. The tests were repeated for 

several species and replications. 

Eight species of Pyemotes are natural enemies of one or 

more forest insects. Six of these attack bark beetles, four 

attack beetles infesting stored timber, one attacks a needle 

sheath midge, and one attacks the pupae of the Douglas-fir 

(Pseudotsuga menziesii) cone moth (Moser and others 1987).

The pyemotid mites are unique because the females of 

many of the species have discrete forms that look so 

different that sometimes they have been placed in different 

classification families.

Potential for biological control—In 1975, Moser 

evaluated 51 species of mites that were collected from 

SPB galleries for their potential as natural control agents 

for reducing SPB populations. Although 32 of the species 

showed predation on one or more of the brood stages, 

four were primary candidates for use as biological control 

in reducing SPB infestations: Histiogaster arborsignis, 

Proctolaelaps dendroctoni, Macrocheles boudreauxi, and

Dendrolaelaps neodisetus. Four other species that showed 

good potential were: Eugamasus lyriformis, Dendrolaelaps 

neocornutus, D. isodentatus, and Proctolaelaps fiseri 

(Moser 1975b). Interestingly, adult beetles were never 

attacked, a fact that may be explained by the phoretic habits 

of the mites. Most mites depend upon riding on the adult 

beetles for dispersal to host material. These mites were 

evaluated in a controlled laboratory situation and are not 

known to be predatory to SPB under natural conditions.

Although many mite species are associated with SPB, only 

the genus Pyemotes is typically predatory to bark beetles. 

One reason for the success of SPB may be the lack of 

Pyemotes mites associated with it (Moser and others 1978). 

Only P. parviscolyti is associated with SPB and only in 

Right: female Pyemotes 

giganticus displaying 

an extreme form of 

polymorphism—note 

enlarged frontal legs.

Below: to bioassay the 

dermatology of Pyemotes

species, mites were held 

in the caps (9-mm inside 

diameter) against the skin 

for four days.
14

a minor way. Under the sponsorship of a PL-480 grant,3

populations of Pyemotes species in Poland were bioassayed 

for potential use in the biological control of SPB in the 

United States (Moser and others 1978).

Of the pyemotid mites located in Poland, Pyemotes dryas

was found to offer the greatest potential as a biological 

control agent because it is a natural enemy of bark beetles 

(the female of this mite species is morphologically identical 

to that of the North American native P. parviscolyti). 

Although the mite was predatory to SPB larvae and pupae, 

the species was precluded from being released in the 

United States for the control of SPB because there was no 

phoretic host for the mite. If it could be reprogrammed to 

ride SPB or one of its associates, it could become a major 

factor in biological control of SPB (Moser and others 1978).

In a continuing search for predatory mites, Moser 

discovered a new genus and two new species of mites that 

3

Public Law 480 (PL-480) grants resulted from a portion of the repayment from loans made to a developing country for agricultural food 

commodities. Some of the funds for repayment could be held in the 

receiving country and used to fund cooperative research between it and 

U.S. Department of Agriculture scientists.

Top: this drawing provides the descriptive information for Megophthrombium gracile, a newly documented 

parasitic mite species. Bottom: this drawing illustrates the five bark beetles that attack southern pines 

and shows the differences in size among the Ips beetles (top three), the southern pine beetle, and the black 

turpentine beetle. Also shown are the locations within the tree bole where the different beetles attack. (Drawing 

from the Southern Forest Research Institute, Boyce Thompson Institute for Plant Research, Forest Research 

Laboratory, Beaumont, TX.)
15

were discovered as parasites of flying SPB adults (Moser 

and Vercammen-Grandjean 1979). These mites, as well 

as parasitic mites of the Parasitengona family, are found 

on the outer bark and presumably attach to beetles during 

or shortly after emergence (Moser 1979). They are found 

infrequently and have little impact on beetle populations. 

Other efforts to develop biological controls included 

evaluations of the transfer of an egg parasite (Pyemotes 

gigananticus) that rides on Douglas-fir pole beetles 

(Pseudohylesinus nebulosus) to SPB. Since these mites are 

phoretic on a wide range of bark beetles, potential exists for 

their transfer to SPB. Tests showed, however, that although 

they readily ride on SPB, they are reluctant to attack 

immature stages of any other bark beetle (Moser 1981).

Although SPB has been the primary focus of efforts to find 

possible biological control agents, opportunities developed 

to evaluate control agents for other insects. Moser and 

others (1986) were first to report a species of Pyemotes

parasitic on pupae of the Douglas-fir cone moth (Barbara 

colfaxiana) native to the Western United States and 

Canada. This mite seems capable of explosive population 

buildups (Moser and others 1987) and has the potential to 

join the straw itch mite (Pyemotes tritici) as a commercial 

biological control agent (Bruce 1983), but seems to lack the 

amount of venom needed to be highly effective. 

Black turpentine beetles (Dendroctonus terebrans), another 

significant pest of southern pine forests, attack the lower 

boles of pines damaged by lightning, logging, or other 

management practices. Currently, control strategies to 

reduce bark beetle populations are chemical treatments, 

infested tree salvage, and preventative management 

of silvicultural practices (Moser 1989). Of potential 

biological controls, the exotic insect predator, a small beetle 

(Rhizophagus grandis), appears to have potential as a 

control agent (Miller and others 1987). R. grandis has been 

introduced and released for evaluations of potential beetle 

control in England, France, Belgium, and Canada.

During 1986 and 1987, three shipments of R. grandis, 

reared in Belgium, were shipped to the Alexandria 

Research Center to test methods of rearing the predators 

on both black turpentine and southern pine beetles. The 

rearing attempts failed due to a pathogen problem. They 

did, however, demonstrate that the predator could be reared 

on black turpentine beetles using semi-artificial methods. 

Additional R. grandis insects were obtained for field release 

trials (Moser 1989, Moser and Branham 1988).

Miller and others (1987) summarized the potential of 

various approaches for developing biological control 

agents for native North American bark beetles. A number 

of studies provided evidence that at least two predatory 

species possess some of the necessary ecological and 

behavioral characteristics to be considered as potential 

biological control agents. One involves a North American 

clerid beetle, Thanasimus undulatus, a predator of the 

Douglas-fir beetle, Dendroctonus pseudotsugae, and the 

other is Rhizophagus grandis, a beetle that is a specific 

predator of Dendroctonus micans, the Great spruce bark 

beetle. Much work remains, however, to finalize the 

development of such control agents. 

Cooperative research on European species—With a 

PL-480 grant with Poland obtained in 1974, Moser and his 

collaborators, Bohdan Kiełczewski and Jerzy Wiśniewski, 

began research to study the mites associated with Polish 

bark beetles. Moser was able to travel to Poland in 1975 

where he worked with his collaborators to write articles 

on the potential of Pyemotes dryas as a biological control 

agent (Moser and others 1978). They also reported on 

surveys of mite associates of bark beetles in Poland 

(Kiełczewski and others 1983). This latter article reported 

that 181 mite species were associated with 45 bark beetle 

species throughout Poland. 

Moser collaborated with several mite specialists to evaluate 

and categorize species associated with bark beetles in 

Large resin deposits on a southern pine resulting from black 

turpentine beetle attack. (Photo from Ronald F. Billings, Texas 

Forest Service)
16

Germany. Moser and cooperator Hermann Bogenschűtz of 

the Forest District of Freiburg sampled 800 beetles from a 

population of 4,725 adult Ips typographus collected from 

pheromone traps in the Black Forest region. From this sample, 

a new species, Scutacarus scolyti, was found (Mahunka and 

Moser 1980). The mites attached to the beetles using a large 

claw located at the end of the mite’s first leg. Another new 

species of mite, Pseudopygmephorus bogenschutzi, was 

described in Mahunka and Moser (1981-1982).

Moser and Bogenschűtz (1984) identified 32 species of 

mites associated with flying I. typographus captured in 

traps in South Germany. Fifteen of these were judged 

phoretic because they were attached to the beetles. The 

biology of most of the species was unknown, but three 

were thought to be potential predators.

From the mite species that Moser and Bogenschűtz (1984) 

evaluated that were captured with I. typographus, a 

new mite subfamily, genus, and species were identified 

(OConnor and Moser 1985). 

The Norway spruce forest of Scandinavia also had 

widespread outbreaks of I. typographus, particularly in 

Sweden. Moser and others (1989) identified 24 species of 

phoretic mites associated with I. typographus, bringing 

the total number of phoretic mites identified for this beetle 

species to 38. 

Identification of other new species—Moser has become 

the internationally recognized authority on the mites related 

to bark beetles and other forest insects. Because of this 

recognition, he has collaborated with numerous scientists 

interested in identifying species and understanding the role 

of mites in the biology of host insects. Due to the number 

of these new genera and species of mites examined, it 

has been impossible to study the biology and role of all of 

these insects. Much of this identification effort was carried 

out with collaboration of Robert L. Smiley of the U.S. 

Department of Agriculture Agricultural Research Service’s 

Systematic Entomology Laboratory in Beltsville, MD. 

A summation of some of these efforts is documented in 

the publication of Histiostoma species associated with 

various pine bark beetles from Honduras, Guatemala, and 

Louisiana (Woodring and Moser 1975), two new species of 

Ereynetes of the Southern United States (Hunter and others 

1989), a new species of Microdispodides associated with 

a western bark beetle (Smiley and Moser 1984b), and two 

new phoretomorphic Siteroptes from galleries of the SPB 

(Smiley and Moser 1976). Weatherby and others (1989) 

documented the biology of a pine needle sheath midge, 

Contarinia acuta, on a new host in Louisiana.

The mite Pyemotes johnmoseri was first located and 

described by Khaustov (1998) in Asia. In 2007, it was found 

to be a predator of the bark beetle Hypoborus ficus which 

attacks Turkish fig (Ficus carica) trees and causes tree 

decline and mortality (Aksit and others 2007). Pyemotes 

johnmoseri was determined to be a predator of the beetle, 

has potential to reduce populations of H. ficus, and should 

be further evaluated.

A major reevaluation of the nomenclature of the 

tarsonemid mites was published by Smiley and Moser 

(1974) where descriptions of new species of these mites 

are presented with additional taxonomical information 

for some taxa. Included are descriptions for one new 

species of Ununguitarsonemus, two new species of 

Heterotarsonemus, four new species of Tarsonemus, and 

the male of T. ips, as well as a redescription of the genus 

Pseudotarsonemoides. A special subgroup (lectoype) 

within the genus Pseudotarsonemoides was designated for 

P. eccoptogasteris and included three new species.

Smiley and Moser (1985) also summarized the existing 

information of the identification and biology of the mite 

genus Heterotarsonemus, as well as describing a new species 

thought to have a relationship to the Dutch elm disease, 

Ceratocystis ulmi (recently renamed Ophiostoma ulmi).

Adaptation of a fungus for attachment to mites—With 

Meredith Blackwell of the Department of Biological 

Sciences at Louisiana State University and Bob Bridges 

and Thelma Perry of the Forest Insect Research Unit, 

Moser documented the first case of a fungus with an 

attachment mechanism for phoresy on arthropods. 

Described in Science (Blackwell and others 1986a), the 

previously unknown method of fungal development with 

extreme specialization for dispersal was discovered. The 

minute ascomycete, Pyxidiophora kimbroughii, is a long, 

slender, two-celled ascospore found associated with the 

mites on SPB (Blackwell and others 1986b).

Robert L. Smiley of the U.S. Department of Agriculture 

Agricultural Research Service’s Systematic Laboratory in 

Beltsville, MD, led the effort to identify many mite species.
17

A darkened region of the ascospores develops into a 

holdfast for attachment to mites where the spores develop 

into a Thaxteriola anamorph (sexual form). Studies have 

found Pyxidiophora ascospores attached to 116 collections 

of 35 species of mites that are associated with beetle 

habitats in trees and wood (Blackwell and others 1989). 

Majewski and Wiśniewski (1978) identified a new species 

of this fungus on mites in Poland and named it Thaxteriola 

moseri in honor of John Moser.

While the effect of the fungus Pyxidiophera kimbroughii

on bark beetle-infested trees and the beetles themselves is 

unknown, it is possible that mites associated with the beetle 

and its habitat are necessary for the spread of such fungi 

and may be important in the biological complex of insects 

and fungi that drive tree infestations (Bridges 1983).

Initiation of Research With Chinese Scientists

In 1984, Yang Zhongqi, of Northwest College of Forestry 

in Beijing, China, arrived in Pineville, LA, to spend a year 

with Moser researching the biological control of the black 

turpentine beetle (Dendroctonus terebrans). With the aid 

of Jean-Claude Gregoire of the Laboratoire de Biologie 

Animale et Cellulaire, Bruxelles, Belgium, they imported 

Rhizophagus grandis, a predator beetle of the European 

turpentine beetle, Dendroctonus micans, to rear, and 

possibly introduce for control of the black turpentine beetle 

(Moser 2000).

Although the initial evaluations were positive, the research 

direction was changed, and this area of research was 

closed. But the experience gained by Yang Zhongqi has 

been put to good use. Yang, who is currently Deputy 

Director of the Institute of Forest Protection, Chinese 

Academy of Forestry, Beijing, has used the training to 

develop a program in China to import R. grandis in an 

effort to establish a biological control of the red turpentine 

beetle, Dendroctonus valens, which caused large losses of 

pines in China (Moser 2000). 

In 1990, Yang Zhongqi arranged for Moser to travel to China 

to see the bark beetle problems in Central, West, and Northeast 

China. Of particular interest to Moser was the observation of 

Dendroctonus armandi, which attacks only the species Pinus 

armandii in the Qinling Mountains of Central China. This 

interesting bark beetle is an object of study by Zhongqi and his 

students, and they are collecting flying beetles so that Moser 

can study the mites associated with the beetle.

Retirement

Moser’s research assignment was under scrutiny in mid1989, and he was being pressured to terminate his town 

ant and mite studies. Since he was not interested in the 

Station’s proposed alternative projects, Moser began to 

consider retirement; with his military service, he had 32 

years of Federal service and therefore could retire with an 

annuity of about one half of his salary. By retiring before 

December 31, 1989, Moser could withdraw all the funds he 

had paid into his retirement system and invest it. John and 

his wife, Martha, had invested aggressively, and his yearly 

annuities from investments totaled twice his salary, so they 

could live comfortably without his Forest Service salary 

(Moser 2000). 

Moser retired on December 31, 1989 and immediately 

signed on as a volunteer with the Forest Service. This status 

meant that he could retain his mite and ant studies and 

maintain complete control of his research. He comments 

that “the arrangement was a bit rocky for 7 years, but 

with the ascension of Kier Klepzig as project leader, the 

environment vastly improved” (Moser 2000).

Since his retirement, Moser’s primary focus has been on 

the relationship of the tarsonemid mites to SPB mycangial 

fungi; he also maintains an interest in the leaf-cutting ants 

and has renewed his first encounter with entomology with 

his study of gall wasps.

Top: Robert Bridges, John Moser, Meredith Blackwell, and 

Thelma Perry. Blackwell is a professor of Mycology at LSU and 

Bridges, Moser, and Perry were with the Station’s Forest Insect 

Research Unit. Blackwell is now a member of the prestigious 

American Academy of Arts and Sciences. Bottom: Yang Zhongqi 

and John Moser observing the Dendroctonus armandi bark beetle 

in Central China.
18

Excavation and Mapping of a Town Ant Colony

Although the excavation and mapping of the mediumsized town ant colony shown below was done many 

years earlier, specific information on the effort was 

not published until 2006. The ant colony located in 

northern Louisiana was excavated completely, and a 

three-dimensional model of its external and subterranean 

features was constructed. In total, 97 fungus gardens, 27 

dormancy cavities, and 45 detritus cavities were located 

(Moser 2006). The complete excavation of such a large 

colony is rare due to its size and the immense resources 

and manpower required.

At the center of this nest was a large central cavity (about 3 

feet in diameter and height), which in the winter functioned 

as a protective site for the colony and where the flying ants 

were reared. Vertical tunnels, possibly as deep as 100 feet, 

served as wells leading to the water table (Moser 2006). A 

number of insects associated with the ants lived in the nest; 

many of these have been described earlier in this document. 

Documentation of this colony excavation is the last major 

effort in Moser’s long interest in understanding town ants, 

but he continues to be an international resource for those 

interested in the biology of leaf-cutting ants. For example, 

Moser collaborated with a group of specialists interested in 

eye size and behavior of day- and night-flying leaf-cutting 

ant alates (Moser and others 1998, Moser and others 2004). 

He, too, worked with a German colleague to study the 

interaction of mites and leaf-cutting ants (Wirth and Moser 

2008). However, research on mites associated with bark 

beetles continues to be his primary interest.

Interrelationships Among Bark Beetles, Mites, 

Fungi, and Nematodes

An extensive collection of organisms, primarily arthropods 

and fungi, can rapidly colonize trees that are attacked 

and killed by bark beetles and pathogenic fungi. The 

composition and arrival sequence of this community are 

predictable. Interactions among the key elements in this 

complex of arthropod natural enemies and associated fungi 

RESEARCH AFTER RETIREMENT

Map of the excavated town ant colony with views from the top and side. The nest was excavated to a depth of 12 feet and a width of 25 feet.
19

can affect significantly the reproduction and survival of 

bark beetles and the pathogenic fungi. These interrelations 

have not been studied in sufficient detail to assess their role 

in regulating bark beetle infestations and tree mortality 

(Stephen and others 1993). Thus began a collaborative 

effort to clarify these interactions.

Early in the effort, Moser and others (1995) documented that 

two species of Tarsonemus phoretic mites on SPB not only 

carried the crescent-shaped ascospores of Ophiostoma minus

blue stain fungus in their sporothecae, but these mites also 

carried tadpole-shaped ascospores of one of the two SPB 

mycangial fungi, Ceratocystiopsis ranaculosus, a species 

described by Thelma Perry (Moser 2000). This discovery of 

a new means of ascospore dispersal provides an additional 

pathway for fungal infection in conifers and may result in 

another agent of mortality of beetle-infested trees.

A note about Thelma J. Perry is due here. She was trained 

as a medical technician in New Orleans and joined the 

Forest Insect Research Unit in Pineville in the late 1960s. 

She worked primarily to support entomologists interested 

in the fungi in the mycangia of SPB, but also collaborated 

with Moser on several projects. Perry was one of the 

earliest female African-American technicians in the 

Southern Station and is described by Moser as an “excellent 

technician.” 4 Tragically, she died of cancer in 1998.

Klepzig and others (2001a) began a significant effort 

studying insect-fungal complexes with a focus on bark 

beetles and their associated organisms, particularly the 

relationship between the southern pine beetle and its 

associates in coniferous trees in the Southern United States.

In a review article, Klepzig and others (2001b) summarized 

the complex, multi-level interactions among bark beetles 

and their fungal and mite associates. They reported that SPB 

attacks and kills southern pines, which, in turn, introduces 

fungi into the tree. Ophiostoma minus may initially aid 

beetles in killing trees, but later this blue stain fungus 

becomes an antagonist, competing with the beetle larvae for 

host phloem. Two additional fungi, Entomocorticium sp. and 

Ceratocystiopsis ranaculosus, are carried within specialized 

mycangium in the beetle and inoculate the phloem, where 

they are fed upon by the beetle larvae. The beetle also 

carries several species of mites which transport spores of 

Ophiostoma minus and Ceratocystiopsis ranaculosus; the 

mites complete their life cycle on these fungi.

4

Blomquist, S.R. 2012. Oral history interview of June 12, 2012, with 

John C. Moser. Unpublished document. Pineville, LA: U.S. Department 

of Agriculture, Forest Service, Southern Research Station, Insects, 

Diseases, and Invasive Plants Research.

These strong indirect interactions of mites and SPB were 

studied by Lombardero and others (2003). Studies were 

conducted to evaluate the effects of phoretic mites on 

SPB. Tarsonemus mites carry ascospores of Ophiostoma 

minus, which tend to out compete mutualistic fungi 

carried by SPB. Experimental additions and removals 

of mites from beetles demonstrated that Tarsonemus

propagate O. minus in beetle galleries. The results 

Top: Pineville forest insect research scientists Kier Klepzig 

and John Moser. Bottom: diagram of the life cycle of 

Ceratocystiopsis ranaculosus. Ascospores are released from 

the fungus on the inner bark where they enter the sporothecae 

of adult female mites. Some mites are phoretic on emerging 

SPB, which disperse not only the mite, but also the fungus (from 

Magowski and Moser 2003.)
20

indicate that the O. minus fungus is an antagonist of 

SPB: beetle larvae seldom survive in the presence of 

O. minus. Apparently, this is a result of O. minus being 

more aggressive than the two species of mycangial fungi 

carried by the beetle for nutrition (Hofstetter and others 

2006, Lombardero and others 2000). Thus, the population 

of Tarsonemus mites and the fungi they carry may 

contribute to the population dynamics of SPB. Noted, 

too, was that the reproduction rate of the mites was more 

temperature sensitive than for beetles (Lombardero 

and others 2003). Consequently, seasonal variations in 

temperature can be anticipated to produce variations in 

SPB populations.

Research on the spruce bark beetle, Dendroctonus 

rufipennis, of the Western United States and Canada 

documented symbiotic associations with a number of 

microorganisms, especially the fungus Leptographium 

abietinum (Cardoza and others 2008). Eight mite species 

were associated with the beetle, but the most prevalent 

was Histiogaster arborsignis. Seventy-five percent of the 

beetles examined carried nematodes, with six species 

represented. Histiostoma arborsignis mites showed strong 

feeding and oviposition preferences for Leptographium 

abietinum among the fungal species evaluated. 

Leptographium abietinum was the fungus which 

cultured the nematodes most successfully, and it plays 

an important role in the bark beetle, mite, and nematode 

interactions (Cardoza and others 2008).

Mite species associated with tree-feeding insects seldom 

affect humans; however, high incidences of itching and 

red, painful welts on people in the Midwestern United 

States during the summer of 2004 led to the discovery of 

an imported European species of mite, Pyemotes herfsi, 

preying on gall-making midge larvae on oak leaves (Broce 

and others 2006). This study also revealed these mites were 

capable of being dispersed by wind, which made them 

difficult to control or avoid.

International Collaboration on Species 

Identification

A major aspect of Moser’s collaboration with international 

specialists is in developing knowledge of species important 

in understanding biological diversity, and this knowledge 

may facilitate developing potential control of insect pests. 

Little progress can be made in biodiversity unless there is 

information of the species involved in biological complexes. 

Moser is involved in numerous international collaborations 

to identify mites and other bark beetle associates. 

Moser is one of only a few forest insect taxonomists in the 

United States; therefore, identification of the many new 

species associated with bark beetles now being discovered 

requires the involvement of international specialists. 

Such collaboration began with Polish scientists Bohdan 

Kiełczewski and Jerzy Wiśniewski in 1974. Later, Wojciech 

L. Magowski worked with Moser to describe three new 

species of the genus Acanthomastix (Magowski and Moser 

1993), describe a new genus and species of mite associated 

with the bark beetle Pseudopityophthorus (Magowski, and 

others 2005), and redescribe the species-group Tarsonemus 

minimax with a description of two new species (Magowski 

and Moser 2003).

Ips typographus, the European bark beetle pest that 

ranges across much of Europe, parts of Asia, and even in 

Japan, carries numerous species of mites that also carry 

fungal ascospores. Ten morphologically distinct types of 

ascospores were found on bodies of 17 species of mites 

associated with I. typographus collected in Sweden (Moser 

and others 1989). Ophiostoma polonicum, the pathogenic 

blue stain fungus, was noted on nine mite species. 

Studies of I. typographus European bark beetles in 

Germany and Georgia (country) were investigated for 

protozoa, fungi, mites, and nematodes. Of all these 

possible antagonists, none were capable of reducing beetle 

populations (Burjanadze and others 2008). In addition, 

Levieux and others (1989) identified a number of mites 

associated with I. sexdentatus in Germany. In Bulgaria, 818 

specimens of I. typographus were tested for the presence of 

phoretic mites; five genera of mites were collected, making 

this the first report of phoretic mites on I. typographus in 

Bulgaria (Takov and others 2009).

Investigation of flying I. typographus japonicus from Japan 

found 12 species of phoretic mites, three of which were not 

previously recorded in Europe (Moser and others 1997). 

On the mites were found seven distinct species of fungal 

spores; Ophiostoma bicolor was the most common species, 

with the pathogenic Ceratocystis polonicum present in 

small numbers.

A mite, Schizosthetus simulatrix, common on bark 

beetles ranging throughout Europe and Asia, was 

studied to understand its morphology and ecology 

(Kaluz and others 2003), and the genera of commonly 

Wojciech L. Magowski of A. 

Mickiewicz University in Poznań, 

Poland, was a collaborator 

studying tarsonemid mites. 
21

encountered associate species were redescribed (AlAtawi and others 2002). Hofstetter, Moser, and McGuire 

(2009) observed S. lyriformis preying on eggs and larvae 

of I. pini, verifying that it is a predator of bark beetles in 

the Western United States.

The species composition and abundance of phoretic 

mites of bark beetles (Pityokteines spp.) on Silver fir 

(Abies alba) were investigated in Croatia. Ten mite 

species were documented for the first time (Pernek and 

others 2008). None of the species appeared to have the 

potential to be used for biological control of the beetles. 

In a later study, new associates of the beetles were 

recovered (Pernek and others 2012).

Collaboration with Mexican Scientists 

The international effort with scientists in Mexico has a 

broader context. Although cooperation began in the 1970s 

when Triplehorn and Moser (1970) identified two new 

species of the bark beetle Corticeus from Mexico and 

Honduras, it was not until Moser met Jorge Macías of 

Chiapas, Mexico, while attending the Western Forest Insect 

Work Conference in the 1990s that collaboration began in 

earnest. Macías was pursuing a Ph.D. under John Borden at 

Simon Fraser University and planned to return to Chiapas 

to study population ecology of SPB. Moser mentioned 

that he had a draft proposal to study the arthropod natural 

enemies of SPB; the proposal had been previously rejected 

as “pie in the sky” by the Forest Service.

Conditions changed, however, following a visit in 1996 by 

President W.J. “Bill” Clinton to the President of Mexico. 

One of the results of this visit was the release of some 

research money for cooperative forest research in Mexico. 

Moser added Jorge Macías’ name and resubmitted the 

proposal to the Forest Service staff at the Washington 

Office (Moser 2000).

The proposal was judged as second in all the proposals 

submitted, and as a result the Forest Insect Research Work 

Unit in Pineville, LA, received $15,000 in “seed money.” The 

timing was great: it arrived late in 1997, at the time when 

Macías received his Ph.D. and was returning to Tapachula, 

Chiapas, Mexico, to join El Colegio de la Frontera Sur 

(ECOSUR). Macías immediately began his studies on the 

SPB problem in Southern Mexico and neighboring countries. 

Since his initial budget for research was nearly zero, the 

grant money and the subsequent visit by Moser and the 

Pineville research unit staff provided a real boost for him. 

Since, there have been frequent communications, as well as 

trips by the Pineville staff to Chiapas, and at least one trip to 

Pineville by Macías to coordinate the joint research on mite 

and insect associates, mycangial fungi, and the impact of 

clerid predators (Moser 2000).

At the 2000 Entomological Society of America meeting in 

Montreal, Macías and Moser presented their initial findings. 

In the paper, they stated reasons why they believe SPB 

invaded the United States from Mexico only about 5,000 

years ago—a fact that has important implications for SPB 

control and future research directions (Moser and MacíasSámano 2000). For example, John Reeve and others have 

shown in the United States that natural enemies have an 

impact on SPB populations, with the most predominate 

candidate being the clerid, Thanasimus dubius (Turchin and 

others 1999). But in Mexico, T. dubius is absent and another 

clerid, Enoclerus ablusus, fills this ecological niche and 

seems to be regulating SPB in Mexico and Central America.

Although the Tarsonemus mites associated with SPB 

mycangial fungi in Chiapas seem to be the same as those in 

the Southern United States, the relative number of ascospores 

of different species that they carry are not. Preliminary 

indications are that in Chiapas, these mites carry lots of 

Ceratocystiopsis ascospores, but few of the Ophiostoma

ascospores—a situation that is reversed in the Southern 

United States. One possible explanation for this difference is 

that the species of these two SPB-associated fungi may differ 

in the Southern United States and Mexico (Moser 2000).

Left: Jorge Macías of Colegio de 

la Frontera Sur in Tapachula, 

Chiapas, Mexico. (Photo courtesy 

of Brian Sullivan) Below: Jane 

Hayes (center) and Kier Klepzig 

(right) with host at Teotihuacan, 

Mexico.
22

Supplementing this research is the recent discovery of 

the Mexican bark beetle, Dendroctonus mexicanus, in the 

Southwestern United States. Both the Mexican bark beetle 

and SPB were found infesting the logs of Chihuahua pine 

(Pinus leiophylla var. chihuahuana) in the Chiricahua 

Mountains of southeastern Arizona (Moser and others 

2005a). It is not known whether this is a recent introduction 

or if the two similar species have been co-occurring in this 

area for some time.

A beetle species new to science was captured in the 

same trapping surveys for bark beetles in Arizona that 

discovered the presence of the Mexican bark beetle in the 

United States. This beetle (Lascontus fitzgibbonae) seems 

to be an associate of one or more of the bark beetle species 

as a predator (Kingsolver and others 2006).

Role of Mites in Disease Transmission

Since the appearance of Dutch elm disease (DED) in the 

early 20th century, elm (Ulmus spp.) trees in Europe, North 

America, and parts of Asia have been seriously affected 

by this destructive vascular wilt disease that is carried by 

two related species of Ophiostoma. The well-known tree 

pathogens, O. ulmi (Buism.) Nannf. and O. novo-ulmi 

Braiser, are vectored by insects, in this case the elm bark 

beetles in the genus Scolytus and also in North America by 

Hylurgopinus rufipes (Lanier and Peacock 1981). 

Scolytus multistriatus and S. pygmaeus are primary 

beetle species infesting elm trees in Europe and in the 20th

century have been introduced and established in North 

America, Australia, and New Zealand. Although both 

species are regarded as secondary bark beetles that breed in 

stressed elm trees, logs, and branches, they serve as vectors 

of DED pathogens from diseased to healthy trees. These 

beetle species do not possess a mycangium, so ascospores 

are carried on their body surface (Moser and others 

2005b). Considering the importance of Scolytus species in 

the transmittal of the DED pathogens, control measures 

against the insects form an essential part of the integrated 

management strategies against DED. 

Although a great deal of research has been devoted to the 

bark beetles attacking elm species and the pathogenic fungi 

that the beetles carry, there has been little emphasis on the 

mites known to be carried on the beetles. An intriguing role 

the mites play on conifers is the relation to the transmission 

of the pathogens, mycangial symbionts, and fungal 

antagonists of bark beetles (Bridges and Moser 1983; 

Moser and others 1989; Moser and others 1995; Moser 

and others 1997; Klepzig and others 2001a, b). Mite taxa 

associated with bark beetles attacking elm trees may differ, 

but their roles should be similar to the mite associates of 

conifer bark beetles.

An investigation was conducted to record and quantify the 

fungal ascospores of Ophiostoma novo-ulmi on phoretic 

mites of three elm bark beetles species common in central 

Europe. Spores were found on 4 of the 10 mite species 

phoretic on Scolytus spp., and all 4 had spores attached 

externally to their body surfaces (Moser and others 2010). 

However, the mite Tarsonemus crassus carried many 

spores within its sporothecae, which are two paired pocketlike structures adapted for fungal transmission. Numerous 

spores were also found in the gut of Proctolaelaps scolyti. 

Top: the southern pine beetle (left) and the Mexican pine beetle 

(right) are closely related species and are difficult to distinguish.

Right: Proctolaelaps scolyti showing Dutch elm disease spores 

in the gut. These spores may remain viable even after digested, 

which reinforces the conclusion that mites significantly aid in 

transmission of the disease.
23

The beetle Scolytus scolytus, which is the most efficient 

vector of O. novo-ulmi in Europe, carried high numbers of 

T. crassus and P. scolyti. Therefore, the high efficiency of 

S. scolytus in spreading Dutch elm disease may be partly 

due to its association with these two mites and the spores of 

O. novo-ulmi they carry (Moser and others 2010). The fact 

remains, however, that these two mites alone are capable of 

transmitting Dutch elm disease.

Fire Ants 

The red imported fire ant, Solenopsis invicta, accidently 

introduced into the United States between 1933 and 1945, 

is an invasive species that has caused great negative 

economic and societal effects across the entire South. 

This small insect, originally from Brazil, has sparked 

much consternation and research. The biology, ecology, 

and control measures of fire ants have been extensively 

studied, but few studies have been conducted on the mites 

associated with the red imported fire ant, specifically on the 

monogyne (i.e., single queen) colonies found in Louisiana 

(Moser and Blomquist 2011).

Why would John Moser become interested in studying 

fire ants, an insect outside the usual range of interest for a 

forest entomologist? For a number of years, Moser’s wife, 

Martha, had a serious medical condition, and John went 

home each day at noon to assist her during lunch. During 

these trips, he observed fire ant mounds in his lawn and as 

an ant expert, he began studying them instead of applying 

ant bait to them. Of special interest was the presence of 

mites and their role in the biology of the ants.

Twenty-two nests were marked and checked daily for a 

16-month period. Some nests died during the 2 years and 

others moved. So, not all 22 nests were sampled on each 

date. The winged reproductive ants or alates were found 

to fly between the hours of 11:30 a.m. and 3:00 p.m.; daily 

flights were triggered only after a suitable amount of rain 

and with acceptable minimum flight temperatures (Moser 

and Blomquist 2011). Duration of flight was about 2.5 

hours, and once alates from a single nest began to fly, the 

period of flight for that nest was 15-30 minutes. Alates 

were plucked from nest surfaces using forceps as the 

winged females and males prepared to fly, usually from 

the tips of grass leaves.

The first efforts dealt with identifying phoretic mites 

associated with the fire ant workers and alates. Four species 

of Scutacarus and one of Imparipes were documented as 

phoretic, and of these, three were new species: Imparipes 

louisiana, Scutacarus nanus, and S. tertius (Ebermann and 

Moser 2008). Khaustov and Moser (2008) identified two 

additional new species of mites associated with the fire ant: 

Petalomium hofstetteri and Caesarodispus klepzigi.

Walter and Moser (2010) described another new mite 

species phoretic on fire ants. Named Gaeolaelaps 

invictianus, this and other species of this genus are 

generally predators, and it is assumed that this species may 

act as a generalist predator in the ant colony.

Wirth and Moser (2010) documented a new species, 

Histiostoma blomquisti, associated with the fire ant. It 

was dedicated to Stacy Blomquist who has contributed 

significantly to the mite research effort. A deutonymph 

Top: John Moser shown collecting fire ants from a nest in his 

yard. Bottom: this mite, Caesarodispus klepzigi, is named in 

honor of Kier Klepzig.
24

(a mite larva stage) was attached to ant queens in large 

numbers, some with more than 200 deutonymphs of H. 

blomquisti. The presence of these on the flying ants’ bodies 

stimulates the cleaning behavior of worker ants, but the 

deutonymphs were not removed in this process. It was 

speculated that the deutonymphs may produce chemical 

components that repel cleaning workers, and they remain 

strongly adhered to the carrier.

In central Louisiana, large numbers of phoretic mite 

species seem to be an important component of S. invicta

nests. They occupy many ecological niches in the nests, 

such as regulating the nest soil fungi and acting as 

predators, parasitoids, or both of each other and the ants. 

Also, the many microorganism species phoretic on the 

mites may be essential to nest biology, as well as possibly 

affecting the vigor of both the ants and the mites (Moser 

and Blomquist 2011). 

Hackberry Gall Makers

Moser has had a long-term interest in hackberry tree 

(Celtis spp.) galls; his earliest research had as its focus 

understanding the insects responsible for the galls on a 

hackberry tree in his front yard (Moser 1956, 1965). In 

1997, Moser obtained galled leaves of Celtis tournefortii

from Kornik, Poland, and found the galls resulted from 

a new species of the midge genus Celticecis (Gagné and 

Moser 1997).

Moser and Gagné (in press) have currently completed 

a comprehensive document of the North American gall 

midges of hackberries. Before this publication, The 

North American gall midges (Diptera: Cecidomyiidae) of 

hackberries (Cannabaceae: Celtis spp.), little was known 

about hackberry gall midges, and only 11 North American 

species had been named and only sketchily described. The 

extensive monograph details 23 species of gall midges 

found in North America north of Mexico and provides keys 

to the galls and larvae (second and third instars) of these 

Nearctic gall midges.

Fire ant alate preparing to fly from a grass. 
25

John Moser continues his efforts to understand the 

relationships among insects and their associates for major 

entomological pests such as the southern pine beetle. His 

research on the mites and other associates of these primary 

insects has opened a new opportunity for increasing 

our understanding of the transmission of tree diseases 

within forest ecosystems. He, too, has put together one 

of the world’s largest collection of slides of mites that are 

associated with major insect pests. 

One aspect of Moser’s legacy is provision in the wills of 

John and his wife, Martha, to fund the establishment of 

two chairs in entomological systematics—one at Cornell 

University and another at Ohio State University. John has 

been an aggressive stock investor and has the capability 

to fund the two professorships that will require about 2 

million dollars each. He may well have additional funding 

to provide support for several graduate students that would 

study under each of the professors (Moser 2000).

Moser also plans to will the contents of his office in the 

Alexandria Research Center to the Southern Research 

Station with the provision that the contents remain attached 

to the Forest Insect Research Work Unit at Pineville and 

not be dispersed to other research units or universities.5

This would preserve the major collection of mite slides that 

will be key to the identification of new mite species.

Although these plans are in place, Moser has no intention 

of fading away. He “has files stuffed with unpublished 

material” and has a lot of things that he still wants to 

do. For instance, he has cooperative agreements with a 

number of scientists dealing with leaf-cutting ants and 

bark beetle mites. Several of these collaborations will 

result in descriptions of new mites associated with bark 

5

Barnett, J.P.; Streett, D.A.; Blomquist, S.R. 2012. Oral history interview 

of March 7, 2012, with John C. Moser. Cassette tape. Pineville, LA: U.S. 

Department of Agriculture Forest Service, Southern Research Station, 

Insects, Diseases, and Invasive Plants Research.

beetles in North and Central America, with keys to these 

species (Moser 2000). Others will focus on increasing the 

understanding of how mites interact with their bark beetle 

hosts and with each other.

Opportunities exist as well to document an interesting 

case of biological control in Chile, where the tip moth, 

Rhycionia bouliana, is being attacked by a parasitoid 

pyemotid mite. Another instance relates to describing 

a new species of Bakerdania, a phoretic pyemotid mite 

found in association with the SPB mycangial fungus, 

Ceratocystiopsis, found in SPB galleries in Guatemala 

(Moser 2000).

The role of mites in the biological control of insect pests 

and transmittal of tree diseases are areas of research that 

must be pursued and may offer opportunities to manage 

both major insect pests and plant diseases in a more 

innovative and productive way. 

Charles Triplehorn (left); long-time friend and college roommate, 

Susan Fisher; Chair of Entomology Department, Moser; and 

John Wenzel, curator of Ohio Museum of Biodiversity, at the 

event of Moser’s bequest to Ohio State University for an endowed 

chair for insect systematics.

IMPLICATIONS FOR FUTURE RESEARCH
26

John Moser has worked for the Southern Research Station 

as a dedicated and productive research scientist and a 

volunteer for over 50 years, and he continues to work in 

his office daily and on most weekends. However, when 

we examine John’s research in greater detail, we find 

a cutting-edge research program with publications in 

several of the world’s top scientific journals, including 

four Science articles and two articles in Nature. Early in 

his career, John’s work with Murray Blum was some of 

the earliest behavioral studies showing that ant foraging 

trails were marked by secretions (Blum and others 1964). 

Later, John was the first to report that the roach inquiline, 

A. fungicola, responded to the pheromone trail emitted by 

a different species, the town ant. John’s work with R.M. 

(Milt) Silverstein established the volatile nature of the 

town ant alarm pheromone and the presence of at least 

two compounds of the town ant trail-marking pheromone 

consisting of volatile and nonvolatile components. John’s 

later collaboration with Silverstein led to the isolation and 

chemical identification of the first ant trail pheromone 

by Tumlinson and others (1971, 1972) that was the major 

volatile component of the trail-marking substance produced 

by the town ant. John and Silverstein later investigated 

the biological responses of enantiomers of the principal 

alarm pheromone of Atta texana. Enantiomers are 

nonsuperimposable mirror images of the same molecule 

which have identical physical and chemical properties. This 

was the first report of insects being able to differentiate 

between enantiomers of the alarm pheromone. John was 

also one of the first to report on the complete excavation 

and mappings of a leaf-cutting ant nest (Moser 2006). 

When John’s research focused more on SPB, he (and 

Bridges) were one of the first to demonstrate that the 

ophiostomatoid fungus, O. minus (formerly Ceratocystis 

minor) was transmitted phoretically by two phoretic mites 

on southern pine beetles (Bridges and Moser 1983, 1984). 

Throughout John’s career he has untaken research problems 

that have interested him, and this initiative led to his 

significant scientific career as a Forest Service scientist.

John Moser’s efforts have brought international recognition 

to the Forest Service, Southern Research Station for 

its excellence in scientific research. His expertise in 

the biology of leaf-cutting ants continues to be sought 

world-wide where those ants are present. The effort 

on mites brings new understanding of the biology of 

major forest insect pests and their insect associates. 

Increased knowledge of these relationships now provides 

an opportunity to understand the role of mites in the 

transmission of some of our major plant diseases and 

potential biological control of insect pests.

Moser’s efforts, especially as a volunteer over the last 20 

years, have brought recognition and respect to the Insects, 

Diseases, and Invasive Plants Research Work Unit to which 

he is attached. Seldom has a volunteer research scientist 

contributed so much to a research program and gained 

international respect in the process. In recognition of his 

contributions to the taxonomy of forest insects, 20 species 

of the associates of town ants, SPB, and other forest beetles 

have been named in his honor (see appendix).

It is important to recognize that during most of the period 

that Moser has volunteered his time and expertise, the 

Insects, Diseases, and Invasive Plants Research Work Unit 

in Pineville, LA, has provided technical support for his 

efforts. Since 2001, Stacy R. Blomquist, who has a master’s 

degree in entomology, has assisted Moser on a part-time 

basis. Her talents have done much to increase the scientific 

productivity and recognition of the program.

The effort of Moser to focus on research that has been 

generally unappreciated has shown what such dedication 

can accomplish. Not only has his research been 

internationally accepted, but it now offers prospects for 

a greater understanding of disease movement in forest 

trees. This commitment continues thanks to his funding 

of major university professorships that will support study 

and training in insect systematics. These chairs will assure 

that scientists with appropriate taxonomic skills will be 

available to continue his program on understanding the role 

of mites and associated species in forest entomology. 

John Moser in his office with shelves holding about half of his 

mite collection. Each box on the shelves contains 100 slides, and 

each slide is numbered—the current number is 53,097. That is a 

lot of mite slides!

SIGNIFICANCE OF MOSER’S CONTRIBUTIONS
27

Aksit, T.; Cakmak, I.; Moser, J. 2007. Attack by Pyemotes johnmoseri

(Acari: Pyemotidae) on Hypoborus ficus (Coleoptera: Scolytidae) in fig 

trees in Turkey. Experimental and Applied Acarology. 41(4): 251-254.

Al-Atawi, F.; Klompen, H.; Moser, J.C. 2002. Redescription of 

Schizosthetus lyriformis (McGraw and Farrier, 1969) (Parasitiformes: 

Parasitidae), with revision of the genus. International Journal of 

Acarology. 28(4): 341-360.

Andryszak, N.A.; Payne, T.L.; Dickens, J.C. [and others]. 1990. Antennal 

olfactory responsiveness of the Texas leaf cutting ant (Hymenoptera: 

Formicidae) to trail pheromone and its two alarm substances. Journal 

of Entomological Science. 25(4): 593-600.

Barnett, J.P.; Pearson, H.A.; Haywood, J.D. 2011. Louisiana’s Palustris 

Experimental Forest: 75 years of research that transformed the 

South. Gen. Tech. Rep. SRS-148. Asheville, NC: U.S. Department of 

Agriculture Forest Service, Southern Research Station. 64 p.

Barras, S.J.; Perry, T. 1972. Fungal symbionts in the prothoracic 

mycangium of Dendroctonus frontalis (Coleopt.: Scolytidae). 

Zeitschrift fűr Angewandte Entomologie. 71: 95-104.

Bennett, W.H. 1958. The Texas leaf-cutting ant. Forest Pest Leaflet 23. 

Washington, DC: U.S. Department of Agriculture, Forest Service. 3 p.

Blackwell, M.; Bridges, J.R.; Moser, J.C.; Perry, T.J. 1986a. 

Hyperphoretic dispersal of a Pyxidiophora anamorph. Science. 

232: 993-995.

Blackwell, M.; Perry, T.J.; Bridges, J.R.; Moser, J.C. 1986b. A new 

species of Pyxidiophora and its Thaxteriola anamorph. Mycologia. 

78(4): 605-612.

Blackwell, M.; Moser, J.C.; Wiśniewski, J. 1989. Ascospores of 

Pyxidiophora on mites associated with beetles in trees and wood. 

Mycological Research. 92(4): 397-403.

Blum, M.S.; Moser, J.C.; Cordero, A.D. 1964. Chemical releasers of 

social behavior. II. Source and specificity of the odor trail substances 

in four attine genera (Hymneoptera: Formicidae). Psyche. 71(1): 1-7.

Bridges, J.R. 1983. Mycangial fungi of Dendroctonus frontalis

(Coleoptera: Scolytidae) and their relationship to beetle population 

trends. Environmental Entomology. 12: 858-861.

Bridges, J.R.; Moser, J.C. 1983. Role of two phoretic mites in 

transmission of bluestain fungus, Ceratocystis minor. Ecological 

Entomology. 8: 9-12.

Bridges, J.R.; Moser, J.C. 1984. A continuous mass-rearing technique 

for the southern pine beetle (Coleoptera: Scolytidae). Journal of the 

Georgia Entomological Society. 19(4): 480-482.

Bridges, J.R.; Moser, J.C. 1986. Relationship of phoretic mites (Acari: 

Tarsonemidae) to the bluestaining fungus, Ceratocystis minor, in 

trees infested by southern pine beetle (Coleoptera: Scolytidae). 

Environmental Entomology. 15(4): 951-953.

Broce, A.B.; Zurek, L.; Kalisch, J.A. [and others]. 2006. Pyemotes herfsi

(Acari: Pyemotidae), a mite new to North America as the cause of bite 

outbreaks. Journal of Medical Entomology. 43(3): 610-613.

Bruce, W.A. 1983. Mites as biological control agents on stored 

product pests. In: Hoy, M.A.; Cunningham, G.L.; Knutson, L., eds. 

Proceedings of the work conference, Biological control of pests by 

mites. Berkeley, CA. University of California Press: 74-78.

Burjanadze, M.; Moser, J.C.; Zimmermann, G.; Kleespies, R. 2008. 

Antagonists of the spruce bark beetle Ips typographus L. (Coleoptera: 

Scolytidae) of German and Georgian populations. In: Ehlers, R.U.; 

Enkerli, J.; Glazer, I. [and others], eds. Insect pathogens and insect 

parasitic nematodes. IOBC/wprs Bulletin. 31: 245-250.

Cardoza, Y.J.; Moser, J.C.; Klepzig, K.D.; Raffa, K.F. 2008. Multipartite 

symbioses among fungi, mites, nematodes, and the spruce beetle, 

Dendroctonus rufipennis. Environmental Entomology. 37(4): 956-963. 

Chillcott, J.G. 1965. New species and stages of Nearctic Fannia R.D. 

(Diptera: Muscidae) associated with nests of Hymenoptera. The 

Canadian Entomologist. 97(6): 640-647.

Cross, E.A.; Moser, J.C. 1971. Taxonomy and biology of some 

pyemotidae (Acarina: Tarsonemoidea) inhabiting bark beetle galleries 

in North American conifers. Acarologia. 8(1): 47-64.

Cross, E.A.; Moser, J.C. 1975. A new, dimorphic species of Pyemotes and 

a key to previously-described forms (Acarina: Tarsonemoidea). Annals 

of the Entomological Society of America. 68(4): 723-732.

Cross, E.A.; Moser, J.C.; Rack, G. 1981. Some new forms of Pyemotes

(Acarina: Pyemotidae) from forest insects, with remarks on 

polymorphism. International Journal of Acarology. 7: 179-196.

Ebermann, E; Moser, J.C. 2008. Mites (Acari: Scutacaridae) 

associated with the red imported fire ant, Solenopsis invicta Buren 

(Hymenoptera: Formicidae), from Louisiana and Tennessee, U.S.A. 

International Journal of Acarology. 34(1): 55-69. 

Echols, H.W. 1966. Texas leaf-cutting ants controlled with pelleted Mirex 

bait. Journal of Economic Entomology. 19: 628-631.

Gagné, R.J.; Moser, J.C. 1997. Celticecis, a genus of gall midges (Diptera: 

Cecidomyiidae), newly reported for the Western Palearctic Region. 

Proceedings of the Entomological Society of Washington. 99(4): 756.

Hermann, H.R.; Moser, J.C.; Hunt, A.N. 1970. The Hymenopterous 

poison apparatus. X. Morphological and behavioral changes in Atta 

texana (Hymenoptera: Formicidae). Annals of the Entomological 

Society of America. 63(6): 1552-1558.

Hofstetter, R.W.; Cronin, J.T.; Klepzig, K.D. [and others]. 2006. 

Antagonisms, mutualisms and commensalisms affect outbreak 

dynamics of the southern pine beetle. Oecologia. 147: 679-691.

Hofstetter, R.W.; Moser, J.C.; McGuire, R. 2009. Observations on the 

mite Schizosthetus lyriformis (Acari: Parasitidae) preying on bark 

beetle eggs and larvae. Entomological News. 120(4): 397-400.

LITERATURE CITED

Acknowledgments

We wish to express our gratitude to Brian Sullivan, U.S. Department of Agriculture Forest Service, Pineville, LA; 

Bobbe Fitzgibbon, U.S. Department of Agriculture Forest Service, Flagstaff, AZ; and Kier Klepzig, U.S. Department of 

Agriculture Forest Service, Asheville, NC, for their critical reviews of the earlier draft of this manuscript.
28

Hunter, P.E.; Moser, J.C. 1968. Pseudoparasitus thatcheri n. sp. (Acarina: 

Dermanyssidae, Laelapinae) associated with southern pine beetles. 

Florida Entomologist. 51(2): 119-123.

Hunter, P.E.; Rosario, R.M.T.; Moser, J.C. 1989. Two new species of 

Ereynetes (Acari: Prostigmata: Eyeynetidae) associated with bark 

beetles. Journal of Entomological Science. 24(1): 16-20.

Johnson, H.R. 1944. Control of the Texas leaf-cutting ant with methyl 

bromide. Journal of Forestry. 42: 130-132.

Kalúz, S.; Mašán, P.; Moser, J.C. 2003. Morphology and ecology of 

Schizosthetus simulatrix (Acari, Mesostigmata) associated with 

galleries of bark beetles (Scolytidae). Biologia, Bratislava. 

58(2): 165-172.

Khaustov, A.A. 1998. Mites of the genus Pyemotes (Acariformes, 

Pyemotidae) associated with bark beetles (Coleoptera: Scolytidae) 

in the Crimea. Zoologicheskii Zhurnal. 77: 1419-1424 (Russian, with 

English summary).

Khaustov, A.A.; Moser, J.C. 2008. Two new species of mites of the 

genera Petalomium Cross and Caesarodispus Mahunka (Acari: 

Heterostigmata: Neopygmephoridae, Microdispidae) associated with 

Solenopsis invicta Buren (Hymenoptera: Formicidae) from the U.S.A. 

International Journal of Acarology. 34(2): 115-121.

Kiełczewski, B.; Moser, J.C.; Wiśniewski, J. 1983. Surveying the 

Acarofauna associated with Polish Scolytidae. Bulletin de la Societe 

des Amis. 22: 151-159.

Kingsolver, J.M.; Stephan, K.; Moser, J.C. 2006. A new species of 

Lasconotus (Coleoptera: Colydiidae) from Arizona and South Dakota, 

U.S.A. Entomological News. 117(1): 53-56.

Kinn, D.N. 1976. Key to mites commonly associated with the southern 

pine beetle. Res. Note SO-214. New Orleans, LA: U.S. Department of 

Agriculture Forest Service, Southern Forest Experiment Station. 11 p.

Klepzig, K.D.; Moser, J.C.; Lombardero, M.J. [and others]. 2001a. 

Mutualism and antagonism: ecological interactions among bark 

beetles, mites and fungi. In: Jeger, M.J.; Spence, N.J., eds. Biotic 

interactions in plant-pathogen associations. Wallingford, UK: CAB 

International, CABI Publishing: 237-267.

Klepzig, K.D.; Moser, J.C.; Lombardero, M.J. [and others]. 2001b. 

Symbiosis and competition: complex interactions among beetles, fungi 

and mites. Symbiosis. 30: 83-96.

Krantz, G.W.; Moser, J.C. 2012. A new genus and species of 

Macrochelidae (Acari: Mesostigmata) associated with the 

Texas leafcutting ant, Atta texana (Buckley) in Louisiana, USA. 

International Journal of Acarology. 38(7): 576-582.

Lanier, G.N.; Peacock, J.W. 1981. Vectors of the pathogen. In: Stipes, 

R.J.; Campana, R.J., eds. Compendium of Elm diseases. The 

American Phytopathological Society, St. Paul, MN:14-16.

Leach, J.G.; Orr, L.W.; Christensen, C. 1934. The interrelationships of 

bark beetles and blue-staining fungi in felled Norway pine timber. 

Journal of Agricultural Research. 49: 315–341.

Levieux, J.; Lieutier, F.; Moser, J.C.; Perry, T.J. 1989. Transportation of 

phytopathogenic fungi by the bark beetle Ips sexdentatus Boerner and 

associated mites. Journal of Applied Entomology. 108:1-11.

Lombardero, M.J.; Ayres, M.P.; Hofstetter, R.W. [and others]. 2003. 

Strong indirect interactions of Tarsonemus mites (Acarina: 

Tarsonemidae) and Dendroctonus frontalis (Coleoptera: Scolytidae). 

Oikos. 102: 243-252.

Lombardero, M.J.; Klepzig, K.D.; Moser, J.C.; Ayres, M.P. 2000. 

Biology, demography and community interactions of Tarsonemus 

(Acarina: Tarsonemidae) mites phoretic on Dendroctonus frontalis

(Coleoptera: Scolytidae). Agricultural and Forest Entomology. 

2: 193-202.

Magowski, W.L.; Lindquist, E.E.; Moser, J.C. 2005. Giselia arizonica, 

a new genus and species of mite (Acari: Tarsonemidae) associated 

with bark beetles of the genus Pseudopityophthorus (Coleoptera: 

Scolytidae) in North America. The Canadian Entomologist. 

137: 648-656.

Magowski, W.L.; Moser, J.C. 1993. Three new species of the genus 

Acanthomastix Mahunka, 1972 from United States and Poland (Acari: 

Dolichocybidae). Entomologische Mitteilungen aus dem Zoologischen 

Museum Hamburg. 11(147): 1-20.

Magowski, W.L.; Moser, J.C. 2003. Redescription of Tarsonemus 

minimax and definition of its species-group in genus Tarsonemus

(Acari: Tarsonemidae) with descriptions of two new species. Annals of 

the Entomological Society of America. 96(4): 345-368.

Majewski, T.; Wiśniewski, J. 1978. New species of parasitic fungi 

occurring on mites (Acarina). Acta Mycology. 14: 3-12.

Mahunka, S.; Moser, J.C. 1980. Scutacarus scolyti sp. n., a new 

Scutacarid species (Acari: Tarsonemina) from Germany. 

Parasitologica Hungarica. 13: 99-103.

Mahunka, S.; Moser, J.C. 1981-1982. New data to the knowledge of the 

tarsonemids (Acari) living on bark beetles. Parasitologica Hungarica. 

14: 87-89.

Miller, M.C.; Moser, J.C.; McGregor, M. [and others]. 1987. Potential for 

biological control of native North American Dendroctonus beetles 

(Coleoptera: Scolytidae). Annals of the Entomological Society of 

America. 80(3): 417-428.

Moser, J.C. 1956. A new species of Torymus (Hymenoptera: Torymidae) 

parasitic on Pachypsylla celtidis-vesicula Riley (Chermidae) with 

notes on its biology and other parasitoids attacking the same host at 

Columbus, Ohio. Journal of the Kansas Entomological Society. 

29(2): 57-62.

Moser, J.C. 1960. The case of the innocent ants. Forests & People. 

10(4): 1-4.

Moser, J.C. 1962. Probing the secrets of the town ant. Forests & People. 

12(4): 12-13, 40-41.

Moser, J.C. 1963. Contents and structure of Atta texana nest in summer. 

Annals of the Entomological Society of America. 56(3): 286-291.

Moser, J.C. 1964. Inquiline roach responds to trail-marking substance of 

leaf-cutting ants. Science. 143(3610): 1048-1049.

Moser, J.C. 1965. The interrelationships of three gall makers and their 

natural enemies, on hackberry (Celtis occidentalis L.). Bull. 402. 

Albany, NY: New York State Museum and Science Service. 95 p.

Moser, J.C. 1967a. Trails of the leafcutters. Natural History. 

LXXVI(1): 32-35.

Moser, J.C. 1967b. Mating activities of Atta texana (Hymenoptera, 

Formicidae). Insectes Sociaux. XIV(3): 295-312.

Moser, J.C. 1970. Pheromones of social insects. In: Wood, D.L.; 

Silverstein, R.M.; Nakajima, M., eds. Control of insect behavior by 

natural products. New York, NY: Academic Press: 162-172.

Moser, J.C. 1975a. Biosystematics of the straw itch mite with special 

reference to nomenclature and dermatology. Transactions of the Royal 

Entomological Society of London. 127(2): 185-191.
29

Moser, J.C. 1975b. Mite predators of the southern pine beetle. Annals of 

the Entomological Society of America. 68(6): 1113-1116.

Moser, J.C. 1976a. Phoretic carrying capacity of flying southern pine 

beetles (Coleoptera: Scolytidae). The Canadian Entomologist. 

108: 807-808.

Moser, J.C. 1976b. Surveying mites (Acarina) phoretic on the southern 

pine beetle (Coleoptera: Scolytidae) with sticky traps. The Canadian 

Entomologist. 108: 809-813.

Moser, J.C. 1979. Parasitengona mites (Acarina: Prositigmata) associated 

with flying adults of the southern pine beetle. International Journal of 

Acarology. 5(1): 24-28.

Moser, J.C. 1981. Transfer of a Pyemotes egg parasite phoretic on western 

pine bark beetles to the southern pine beetle. International Journal of 

Acarology. 7: 197-202.

Moser, J.C. 1983. Town ant. In: Payne, T.L.; Billings, R.F.; Coulson, R.N.; 

Kulhavy, D.L., eds. History, status, and future needs for entomology 

research in southern forests. Proceedings of the 10th anniversary of the 

East Texas Forest Entomology Seminar, October 6-7, 1983. Misc. Publ. 

MP-1553. Kurth Lake, TX: Texas Agricultural Experiment Station: 

47-52.

Moser, J.C. 1985. Use of sporothecae by phoretic Tarsonemus mites 

to transport ascospores of coniferous bluestain fungi. Transactions 

British Mycological Society. 84: 750-753.

Moser, J.C. 1986. Estimating timber losses from a town ant colony with 

aerial photographs. Southern Journal of Applied Forestry. 10(1): 45-47.

Moser, J.C. 1989. Inoculative release of an exotic predator for the 

biological control of the black turpentine beetle. In: Kulhavy, D.L.; 

Miller, M.C. eds. Potential for biological control of Dendroctonus

and Ips bark beetles. Nacogdoches, TX: Stephen F. Austin State 

University, School of Forestry, Center for Applied Studies: 189-200.

Moser, J.C. 2000. Events leading to one person’s career in forest 

entomology. A.D. Hopkins address. In: Proceedings, 44th Southern 

forest insect work conference, July 31-Aug. 3, 2000. Memphis, TN. 

URL: www.fsl.orst.edu/wfiwc/awards/speeches/moser-address.htm. 

[Date accessed: March 20, 2012].

Moser, J.C. 2006. Complete excavation and mapping of a Texas leafcutting ant nest. Annals of the Entomological Society of America. 

99(5): 891-897.

Moser, J.C.; Blomquist, S.R. 2011. Phoretic arthropods of the red 

imported fire ant in central Louisiana. Annals of the Entomological 

Society of America. 104(5): 886-894.

Moser, J.C.; Blum, M.S. 1963. Trail marking substance of the Texas leafcutting ant: source and potency. Science. 140(3572): 1228.

Moser, J.C.; Bogenschűtz, H. 1984. A key to the mites associated with 

flying Ips typographus in South Germany. Zeitschrift fűr angewandte 

Entomologie. 5: 437-450.

Moser, J.C.; Branham, S.J. 1988. Bugs that eat bugs. Forest Farmer. 

47(3): 17-20.

Moser, J.C.; Bridges, J.R. 1983. Technique for rearing mite-free southern 

pine beetle, Dendroctonus frontalis Zimmermann (Coleoptera: 

Scolytidae), adults. Annals of the Entomological Society of America. 

76(6): 942-945.

Moser, J.C.; Bridges, J.R. 1986. Tarsonemus (Acarina: Tarsonemidae) 

mites phoretic on the southern pine beetle (Coleoptera: Scolytidae): 

attachment sites and numbers of bluestain (Ascomycetes: 

Ophiostomataceae) ascospores carried. Proceedings of the 

Entomological Society of Washington. 88(2): 297-299.

Moser, J.C.; Browne, L.E. 1978. A nondestructive trap for Dendroctonus 

frontalis Zimmerman (Coleoptera: Scolytidae). Journal of Chemical 

Ecology. 4(1): 1-7.

Moser, J.C.; Brownlee, R.C.; Silverstein, R.M. 1968. Alarm pheromones 

of the ant, Atta texana. Journal of Insect Physiology. 14: 529-535.

Moser, J.C.; Cross, E.A. 1975. Phoretomorph: a new phoretic phase 

unique to the Pyemotidae (Acarina: Tarsonemoidea). Annals of the 

Entomological Society of America. 68(5): 820-822.

Moser, J.C.; Cross, E.A.; Roton, L.M. 1971. Biology of Pyemotes 

parviscolyti (Acarina: Pyemotidae). Entomophaga. 16(1): 367-379.

Moser, J.C.; Dell, T.R. 1979. Predictors of southern pine beetle flight 

activity. Forest Science. 25(2): 217-222.

Moser, J.C.; Dell, T.R. 1980. Weather factors predicting flying populations 

of a clerid predator and its prey, the southern pine beetle. In: Berryman, 

A.A.; Safranyik, L., eds. Dispersal of forest insects: evaluation, 

theory, and management implications. Proceedings of the 2nd IUFRO 

conference, Sandpoint, ID. Pullman, WA: Washington State University, 

Cooperative Extension Service, Conference Office: 266-278.

Moser, J.C.; Della Lucia, T.M.C.; Bento, J.M.; Cameron, R.S. 1998. 

Morphological and behavioral differences between day and nightflying leafcutting ant alates of the genus Atta (Hymenoptera: 

Formicidae). In: McManus, M.L.; Liebhold, A.M., eds. Proceedings: 

Population dynamics, impacts, and integrated management of forest 

defoliating insects. Gen. Tech. Rep. NE-247. Radnor, PA: U.S. 

Department of Agriculture Forest Service, Northeastern Forest 

Experiment Station: 353.

Moser, J.C.; Eidmann, H.H.; Regnander, J.R. 1989. The mites associated 

with Ips typographus in Sweden. Annales Entomologici Fennici. 

55(1): 23-27.

Moser, J.C.; Fitzgibbon, B.A.; Klepzig, K.D. 2005a. The Mexican pine 

beetle, Dendroctonus mexicanus: first record in the United States and 

co-occurrence with the southern pine beetle—Dendroctonus frontalis

(Coleoptera: Scolytidae or Curculionidae: Scolytinae). Entomological 

News. 116(4): 235-243.

Moser, J.C.; Gagne, R.J. [In press]. The North American gall midges 

(Diptera: Cecidomyiidae) of hackberries (Cannabaceae: Celtis spp.).

Moser, J.C.; Kiełczewski, B.; Wiśniewski, J.; Bałazy, S. 1978. Evaluating 

Pyemotes dryas (Vitzthum1923) (Acari: Pyemotidae) as a parasite of 

the southern pine beetle. International Journal of Acarology. 

4(2): 67-70.

Moser, J.C.; Konrad, H.; Blomquist, S.R.; Kirisits, T. 2010. Do mites 

phoretic on elm bark beetles contribute to the transmission of Dutch 

elm disease. Naturwissenschaften. 97: 219-227.

Moser, J.C.; Konrad, H.; Kirisits, T.; Carta, L.K. 2005b. Phoretic mites 

and nematode associates of Scolytus multistriatus and Scolytus 

pygmaeus (Coleoptera: Scolytidae) in Austria. Agricultural and Forest 

Entomology. 7: 169-177.

Moser, J.C.; Lewis, J.R. 1981. Multiple nest queens of Atta texana

(Buckley 1860): (Hymenoptera: Formicidae). Turrialba 31(3): 256-257.

Moser, J.C.; Macías-Sámano, J.E. 2000. Tarsonemid mite associates of 

Dendroctonus frontalis (Coleoptera: Scolytidae): implications for the 

historical biogeography of D. frontalis. The Canadian Entomologist. 

132: 765-771.

Moser, J.C.; Neff, S.E. 1971. Pholeomyia comans (Diptera: Milichiidae) 

an associate of Atta texana: larval anatomy and notes on biology. 

Zeitschrift für angewandte Entomologie. 69(4): 343-348.
30

Moser, J.C.; Perry, T.J.; Bridges, J.R.; Yin, H. 1995. Ascospore dispersal 

of Ceratocystiopsis ranaculosus, a mycangial fungus of the southern 

pine beetle. Mycologia. 87(1): 84-86.

Moser, J.C.; Perry, T.J.; Furuta, K. 1997. Phoretic mites and their 

hyperphoretic fungi associated with flying Ips typographus japonicus

Niijima (Col., Scolytidae) in Japan. Journal of Applied Entomology. 

121: 425-428.

Moser, J.C.; Perry, T.J.; Solheim, H. 1989. Ascospores hyperphoretic on 

mites associated with Ips typographus. Mycological Research. 

93(4): 513-517.

Moser, J.C.; Reeve, J.D.; Bento, J.M.S. [and others]. 2004. Eye size and 

behaviour of day- and night-flying leafcutting ant alates. Journal of 

Zoology, London. 264: 69-75.

Moser, J.C.; Roton, L.M. 1970. Tagging mites with aerosol paint. Annals 

of the Entomological Society of America. 63(6): 1784.

Moser, J.C.; Roton, LM. 1971. Mites associated with southern pine bark 

beetles in Allen Parish, Louisiana. The Canadian Entomologist.

103: 1775-1798.

Moser, J.C.; Roton, L.M. 1972. Reproductive compatibility between 

two widely separated populations of Pyemotes scolyti. (Acarina: 

Pyemotidae). The Pan-Pacific Entomologist. 48(2): 97-99. 

Moser, J.C.; Silverstein, R.M. 1967. Volatility of trail marking substance 

of the town ant. Nature. 215(5097): 206-207.

Moser, J.C.; Smiley, R.L.; Otvos, I.S. 1987. A new Pyemotes (Acari: 

Pyemotidae) reared from the Douglas-fir cone moth. International 

Journal of Acarology. 13(2): 141-147.

Moser, J.C.; Smiley, R.L.; Otvos, I.S.; Lorio, P.L., Jr. 1986. Pyemotes n. 

sp. (Acari: Pyemotidae), a potential biological control agent for coneand-seed insects. In: Roques, A., ed. Proceedings of the 2nd conference 

of the cone and seed insects working party S2. 07-01. Briancon, 

France: International Union of Forestry Research Organisations and 

Institut National de La Recherche 

Agronomique: 307-308.

Moser, J.C.; Thatcher, R.C.; Pickard, L.S. 1971. Relative abundance 

of southern pine beetle associates in East Texas. Annals of the 

Entomological Society of America. 64(1): 72-77.

Moser, J.C.; Thompson, W.A. 1986. Temperature thresholds related to 

flight of Dentroctonus frontalis Zimm. (Col.: Scolytidae). Agronomie. 

6(10): 905-910.

Moser, J.C.; Vercammen-Grandjean, P.H. 1979. Megophthrombium 

gracile n. sp. and Diathrombium diaphane n.g., n.sp. (Acarina: 

Trombidiidae), two larval parasites of adult southern pine beetles. 

International Journal of Acarology. 5(1): 18-23.

Moser, J.C.; Wilkinson, R.C.; Clark, E.R. 1974. Mites associated with 

Dendroctonus frontalis Zimmerman (Scolytidae: Coleoptera) in 

Central America and Mexico. Turrialba. 24(4): 379-381.

Neelands, R.W. 1959. Exposing the town ant. Forests & People. 

9(4): 16, 19, 50.

Nelson, R.M. 1934. Effect of bluestain fungi on southern pines attacked 

by bark beetles. Phytopathologische Zeitschrift. 7: 327-353.

OConnor, B.M.; Moser, J.C. 1985. Phylogenetic relationships of the 

Algophagidae (Acari: Astigmata), with descriptions of a new 

subfamily, genus, and species. Annals of the Entomological Society 

of America. 78(6): 783-789.

Pernek, M.; Hrasovec, B.; Matosevic, D. [and others]. 2008. Phoretic 

mites of three bark beetles (Pityokteines spp.) on Silver fir. Journal of 

Pest Science. 81: 35-42.

Pernek, M.; Wirth, S.; Blomquist, S.R. [and others]. 2012. New 

associations of phoretic mites on Pityokteines curvidens (Coleoptera, 

Curculionidae, Scolytinae). Central Europe Journal of Biology. 

7(1): 63-68. 

Riley, R.G.; Silverstein, R.M.; Moser, J.C. 1974a. Biological responses 

of Atta texana to its alarm pheromone and the enantiomer of the 

pheromone. Science. 183(4126): 760-762.

Riley, R.G.; Silverstein, R.M.; Moser, J.C. 1974b. Isolation, identification, 

synthesis and biological activity of volatile compounds from the heads 

of Atta ants. Journal of Insect Physiology. 20: 1629-1637.

Robinson, S.W.; Moser, J.C.; Blum, M.S.; Amante, E. 1974. Laboratory 

investigations of the trail-following responses of four species of leafcutting ants with notes on the specificity of a trail pheromone of Atta 

texana (Buckley). Insectes Sociaux. 21: 87-94.

Roton, L.M. 1978. Mites phoretic on the southern pine beetle: when and 

where they attach. The Canadian Entomologist. 110: 557-558.

Sacktor, B.; Thomas, G.M.; Moser, J.C.; Bloch, D.I. 1953. 

Dephosphorylation of adenosine triphosphate by tissues of the 

American cockroach, Periplaneta americana (L.). Biological Bulletin. 

105(1): 166-173.

Sonnet, P.E.; Moser, J.C. 1972. Synthetic analogs of the trail pheromone 

of the leaf-cutting ant, Atta texana (Buckley). Journal of Agricultural 

and Food Chemistry. 20(6): 1191-1194.

Sonnet, P.E.; Moser, J.C. 1973. Trail pheromones: responses of the Texas 

leafcutting ant, Atta texana to selected halo- and cyanopyrrole-2-

aldehydes, ketones, and esters. Environmental Entomology. 

2(5): 851-854.

Spangler, P.J. 1962. A new species of the Genus Oosternum and a key 

to the U.S. species (Coleoptera: Hydrophilidae). Proceedings of the 

Biological Society of Washington. 75: 97-100.

Smiley, R.L.; Moser, J.C. 1968. New species of mites from pine (Acarina: 

Tarsochelidae, Eupalopsellidae, Caligonellidae, Cryptognathidae, 

Raphignathidae, and Neophyllobiidae). Proceedings of the 

Entomological Society of Washington. 70(4): 307-317.

Smiley, R.L.; Moser, J.C. 1970. Three cheyletids found with pine bark 

beetles (Acarina: Cheyletidae). Proceedings of the Entomological 

Society of Washington. 72(2): 229-236.

Smiley, R.L.; Moser, J.C. 1974. New Tarsonemids associated with bark 

beetles (Acarina: Tarsonemidae). Annals of the Entomological Society 

of America. 67(4): 639-665.

Smiley, R.L.; Moser, J.C. 1976. Two new phoretomorphic Siteroptes from 

galleries of the southern pine beetle. Beiträge zur Entomologie Berlin. 

26(1): 307-322.

Smiley, R.L.; Moser, J.C. 1984a. Notes and a key to separate normal and 

heteromorphic males of Pyemotes giganticus Cross, Moser, and Rack 

and P. dimorphus Cross and Moser (Acari: Pyemotidae). International 

Journal of Acarology. 10(1): 11-14.

Smiley, R.L.; Moser, J.C. 1984b. A new Microdispodides (Acari: 

Pygmephoridae) associated with a western bark beetle. International 

Journal of Acarology. 10(1): 19-22.

Smiley, R.L.; Moser, J.C. 1985. A new species, key to females, and 

distribution records for Heterotarsonemus (Acari: Tarsonemidae). 

International Journal of Acarology. 11(4): 247-253.

Stephen, F.M.; Berisford, C.W.; Dahlsten, D.L. [and others]. 1993. 

Invertebrate and microbial associates. In: Schowalter, T.D.; Filip, 

G.M., eds. Beetle-pathogen interactions in conifer forests. New York, 

NY: Academic Press, Harcourt, Brace & Company: 128-153.
31

Takov, D.; Pilarska, D.; Moser, J. 2009. Phoretic mites associated 

withspruce bark beetle Ips typographus L. (Curculionidae: Scolytinae) 

from Bulgaria. Acta Zoologica Bulgarica. 61(3): 293-296.

Taylor, A.D.; Hayes, J.L.; Roton, L.; Moser, J.C. 1992. A phloem 

sandwich allowing attack and colonization by bark beetles 

(Coleoptera: Scolytidae) and associates. Journal of Entomological 

Science. 27(4): 311-316.

Triplehorn, C.A.; Moser, J.C. 1970. Two new species of Corticeus from 

Mexico and Honduras (Coleoptera: Tenebrionidae). The Coleopterists’ 

Bulletin. 24: 47-50.

Tumlinson, J.H.; Moser, J.C.; Silverstein, R.M. [and others]. 1972. A 

volatile trail pheromone of the leaf-cutting ant, Atta texana. Journal 

of Insect Physiology. 18: 809-814.

Tumlinson, J.H.; Silverstein, R.M.; Moser, J.C. [and others]. 1971. 

Identification of the trail pheromone of a leaf-cutting ant, Atta texana. 

Nature. 234: 348-349.

Turchin, P.; Taylor, A.D.; Reeve, J.D. 1999. Dynamical role of predators 

in population cycles of a forest insect: an experimental test. Science. 

285: 1068-1071.

Waller, D.A.; Moser, J.C. 1990. Invertebrate enemies and nest associates 

of the leaf-cutting ant, Atta texana (Buckley) (Formicidae, Attini). In: 

Vander Meer, R.K.; Jaffe, K.; Cedeno, A., eds. Applied myrmecology, 

a world perspective. Boulder, CO: Westview Press: 256-273.

Walter, D.E.; Moser, J.C. 2010. Gaeolaelaps invictianus, a new and 

unusual species of hypoaspidine mite (Acari: mesostigmata: 

Laelapidae) phoretic on the red imported fire ant Solenopsis invicta

Buren (Hymenoptera: Formicidae) in Louisiana, USA. International 

Journal of Acarology. 36(5): 399-407.

Walter, E.V.; Seaton, L.; Mathewson, A.A. 1938. The Texas leaf-cutting 

ant and its control. Circular 494. Washington, DC: U.S. Department of 

Agriculture, Bureau of Entomology and Plant Quarantine. 19 p.

Warter, S.L.; Moser, J.C.; Blum, M.S. 1962. Some correlations between 

the foraging behavior of common nighthawks, Chordeiles minor

(Forster), and the swarming behavior of two species of ants, Atta 

texana (Buckley) and Iridomyrmex pruinosis (Roger). Proceedings of 

the Louisiana Academy of Sciences. 25: 42-46.

Weatherby, J.C.; Moser, J.C.; Gagné, R.J.; Wallace, H.N. 1989. Biology 

of a pine needle sheath midge, Contarina acuta Gagne (Diptera: 

Cecidomyiidae), on loblolly pine. Proceedings of the Entomological 

Society of Washington. 91(3): 346-349.

Wirth, S.; Moser, J.C. 2008. Interactions of Histiostomatid mites 

(Astigmata) and leafcutting ants. In: Bertrand, M.; Kreiter, S.; McCoy, 

K.D.; Migeon, A.; Navajas, M.; Tixier, M.-S.; Vial, L., eds. Integrative 

Acarology, Proceedings of the 6th European Congress, European 

Association of Acarologists: 378-384.

Wirth, S.: Moser, J.C. 2010. Histiostoma blomquisti n. sp. (Acari: 

Astigmata: Histiostomatidae), a phoretic mite of the red imported fire 

ant, Solenopsis invicta Buren (Hymenoptera: Formicidae). Acarologia. 

50(3): 357-371.

Woodring, J.P.; Moser, J.C. 1970. Six new species of anoetid mites 

associated with North American Scolytidae. The Canadian 

Entomologist. 102(10): 1237-1257.

Woodring, J.P.; Moser, J.C. 1975. Description of Histiostoma 

conjuncta (new comb.) (Acari: Anoetidae), an associate of Central 

American bark beetles. Proceedings of the Entomological Society of 

Washington. 77(1): 83-86.

Woodruff, R.E.; Cartwright, O.L. 1967. A review of the Genus Euparixia

with description of a new species from nests of leaf-cutting ants in 

Louisiana (Coleoptera: Scarabaeidae). Proceedings of the United 

States National Museum. 123(3616): 1-21.
32

Species of arthropods and a fungus named in recognition of John C. Moser’s contributions to the taxonomy of forest 

insects. 

Species name and describer

Year 

designated Description of species

Fannia moseri Chillcott 1965 Fly (Diptera) found in nests of leaf-cutting ants feeding on fresh 

detritus.

Histiogaster moseri Woodring 1966 Mite found in boring dust of the southern pine beetle (D. 

frontalis) from Pinus oocarpa in Teguciqualpa, Honduras.

Dendrolaelaps moseri (Hurlbutt) 1967 A predatory mite found associated in galleries of elm bark beetles 

(Scolytus multistriatus) in Delaware, OH.

Euparixia moseri Woodruff & Cartwright 1967 Beetle found associated with leaf-cutting ant (Atta texana) nests 

near Flatwoods, LA.

Tarsonemus moseri Smiley 1967

A fungivore mite collected from the inner galleries of loblolly 

pine with southern pine beetle (Dendroctonus frontalis) in 

Elizabeth, LA.

Laelaspis moseri (Hunter & Glover) 1968 Mite found associated with ants; role in the ant colony is 

unknown.

Neivamyrmex moseri Watkins 1968 Ant collected from excavated surface subsoil of leaf-cutting ants 

(Atta texana) seven miles west of Alexandria, LA.

Nenteria moseri Hirschmann 1972

Mite found in inner bark of Endlicher pine (Pinus rudis) and 

in southern pine beetle (Dendroctonus frontalis) galleries of 

Guatemala.

Oplitis moseri Hirschmann 1972 Mite found phoretic on fire ants (Solenopsis invicta).

Trichouropoda moseri Hirschmann 1972 Mite found associated with bark beetle (Scolytidae) galleries in 

Canada.

Uroobovella moseri Hirschmann 1972 Mite collected from bark beetle (Scolytidae) galleries in 

Honduras.

Metagynella moseri Hirschmann 1975 Mite found associated with the Bess beetle (Odontotaenius 

disjunctus).

Thaxteriola moseri Majewsky & Wiśniewski 1978 Fungus found associated with mites from Poland.

Proctolaelaps moseri Wiśniewski 1980 Mite found living in the galleries of bark beetles in Poland.

Metagynella moserisimilis Hiramatsu 1981 Mite found associated with a stag beetle (Lucanidae) in Taiwan.

Dendrolaelaps moserisimilis Shcherbak 1984 A predatory mite found under the bark of a poplar tree in Poltava, 

Ukraine

Julolaelaps moseri Hunter & Rosario 1986 Mite found on large millipede (family Spirostreptidae) in the 

Insect Zoo, Smithsonian Institute, Washington, DC

Neobonzia moseri Smiley 1992 A predatory mite of small arthropods that was found on the 

outside bark of a loblolly pine in Elizabeth, LA.

Pyemotes moseri Yu & Liang 1996 A parasitic mite found on bark beetles (Cryphalus sp.) infesting 

Chinese hickory trees from Hebei, China.

Pyemotes johnmoseri (Khaustov) 2004 A mite which preys on fig tree bark beetles (Hypoborus ficus) in 

Turkey.

APPENDIX
Barnett, J.P.; Streett, D.A.; Blomquist, S.R. 2013. Town ants: 

the beginning of John Moser’s remarkable search for knowledge. 

Gen. Tech. Rep. SRS-182. Asheville, NC: U.S. Department of 

Agriculture Forest Service, Southern Research Station. 32 p. 

John C. Moser’s career spans over 50 years, and his research has 

focused on understanding the biology of town ants (Atta texana) 

and phoretic mites and other associates of ants and pine bark 

beetles. His approach to developing methods for the control of these 

pests has been to understand more completely the biology of these 

organisms. This research approach has established Moser as an 

internationally recognized expert in leaf-cutting ant biology and as a 

premier authority on phoretic mites and other associates of pine bark 

beetles. John’s efforts have led to the largest collection (over 30,000 

specimens) of mites associated with forest insects in the world. This 

knowledge is leading entomologists, ecologists, and pathologists to 

consider the possible role of mites in insect-fungal symbioses and 

disease transmission in trees.

Keywords: Atta texana, Dendroctonus frontalis, Dutch elm disease, 

fungal ascospores, mites and other pine bark beetle associates, 

southern pine beetles, southern pines, Texas leaf-cutting ants.

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