• TABLE OF CONTENTS
HIDE
 Copyright
 Front Cover
 Title Page
 Table of Contents
 Preface
 I. Introduction
 II. Biology of pest mole crick...
 III. Biological control of mole...
 IV. Chemical control of mole...
 V. Further reading
 Reference
 Back Cover














Group Title: Bulletin - Agricultural Experiment Stations, University of Florida - 846
Title: Mole crickets in Florida
CITATION PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00027758/00001
 Material Information
Title: Mole crickets in Florida
Series Title: Bulletin Agricultural Experiment Stations, University of Florida
Physical Description: iv, 54 p. : ill (some col.) ; 23 cm.
Language: English
Creator: Sailer, R. I ( Reece Ivan ), 1915-
Walker, Thomas J ( Thomas Jefferson )
Publisher: Agricultural Experiment Stations, Institute of Food and Agricultural Sciences, University of Florida
Place of Publication: Gainesville Fla.
Publication Date: 1984
 Subjects
Subject: Mole crickets   ( lcsh )
Genre: government publication (state, provincial, terriorial, dependent)   ( marcgt )
bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Bibliography: p. 51-54.
Statement of Responsibility: R.I. Sailer ... et al., T.J. Walker, editor.
General Note: "December 1984."
Funding: Bulletin (University of Florida. Agricultural Experiment Station) ;
 Record Information
Bibliographic ID: UF00027758
Volume ID: VID00001
Source Institution: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: aleph - 000573439
oclc - 14158491
notis - ADA0633
issn - 0096-607X ;

Table of Contents
    Copyright
        Copyright
    Front Cover
        Page i
    Title Page
        Page ii
    Table of Contents
        Page iii
    Preface
        Page iv
    I. Introduction
        Page 1
        History
            Page 1
        Acknowledgments
            Page 2
    II. Biology of pest mole crickets
        Page 3
        Page 4
        Page 5
        Page 6
        Systematics and life cycles
            Page 3
        Species and their identifiication
            Page 3
            Geography
                Page 3
            Life cycles
                Page 7
            Dispersal
                Page 7
                Page 8
                Page 9
        Reproductive behavior
            Page 10
            Male calling
                Page 10
            Flight
                Page 11
            Phonotaxis and mate choice
                Page 12
                Page 13
            Mating behavior
                Page 14
            Egg laying and hatching
                Page 15
        Other behavior, damage, and sampling
            Page 16
            Feeding
                Page 16
            Burrowing
                Page 16
            Damage
                Page 17
            Population estimation
                Page 18
                Page 19
                Page 20
                Page 21
                Page 22
    III. Biological control of mole crickets
        Page 23
        Natural enemies
            Page 23
            Predators
                Page 23
            Parasites
                Page 23
                Page 24
                Page 25
                Page 26
                Page 27
            Past and present research on natural enemies of Florida's mole crickets
                Page 28
                Page 29
                Page 30
            Future research on natural enemies
                Page 31
            Diseases
                Page 32
                Pathogens and symtoms
                    Page 32
                Activity of metarhizium anisopliae against mole crickets
                    Page 33
                Future research
                    Page 33
                    Page 34
            Resistant varieties
                Page 35
                Page 36
                Bermudagrass selections
                    Page 37
                    Page 38
                Bahiagrass selections
                    Page 39
                Comparison of species and cultivars of turfgrass
                    Page 39
                Pasture grass selections
                    Page 40
                Host resistance potential for grasses
                    Page 40
    IV. Chemical control of mole crickets
        Page 41
        Beginnings
            Page 41
        Present day control
            Page 42
            Baits
                Page 42
                Page 43
                Page 44
                Page 45
            Sprays and granules
                Page 46
            Evaluation of chemical control
                Page 46
        Future prospects
            Page 47
            Addendum
                Page 48
    V. Further reading
        Page 49
        Page 50
    Reference
        Page 51
        Page 52
        Page 53
        Page 54
    Back Cover
        Page 55
Full Text





HISTORIC NOTE


The publications in this collection do
not reflect current scientific knowledge
or recommendations. These texts
represent the historic publishing
record of the Institute for Food and
Agricultural Sciences and should be
used only to trace the historic work of
the Institute and its staff. Current IFAS
research may be found on the
Electronic Data Information Source
(EDIS)

site maintained by the Florida
Cooperative Extension Service.






Copyright 2005, Board of Trustees, University
of Florida





















































Agricultural Experiment Stations
Institute of Food and Agricultural Sciences
University of Florida, Gainesville
F. A. Woods, Dean of Research


r' ^ '


-~ i~-~Sr"
S;Fia~tt~*~





December 1984


Mole Crickets in Florida
R. I. Sailer, J. A. Reinert, Drion Boucias, Philip Busey,
R. L. Kepner, T. G. Forrest, W. G. Hudson, T. J. Walker
(T. J. Walker, editor)


























Agricultural Experiment Stations
Institute of Food and Agricultural Sciences
University of Florida, Gainesville
F. A. Woods, Dean of Research


Bulletin 846












TABLE OF CONTENTS

Preface .......................... .. ............................ iv

I. INTRODUCTION ........................... .................. 1
History ................. ............. ............ 1
Acknowledgments ................ ........................ 2

II. BIOLOGY OF PEST MOLE CRICKETS ...................... 3
Systematics and Life Cycles ................................ 3
Species and their identification ........................... 3
Geography ......................................... 3
Life cycles ............................................ 7
Dispersal ................... .............................. 7
Reproductive Behavior .................................... 10
Male calling ......................................... 10
Flight ................................... ............. 11
Phonotaxis and mate choice ............................ 12
Mating behavior ....................................... 14
Egg laying and hatching ................................. 15
Other Behavior, Damage, and Sampling ..................... 16
Feeding ................................... ...... 16
Burrowing ............................. ............ 16
Damage .......... .............................. 17
Population estimation .............. ................. 18

III. BIOLOGICAL CONTROL OF MOLE CRICKETS .............. 23
Natural Enemies ................ ........................ 23
Predators ............................... ............. 23
Parasites .................. ............................. 23
Past and present research on natural enemies
of Florida's mole crickets .............................. 28
Future research on natural enemies ...................... 31
Diseases ................. ................................ 32
Pathogens and symptoms ............................... 32
Activity of Metarhizium anisopliae against mole crickets ... 33
Future research ....................................... 33
Resistant Varieties ....................................... 35
Bermudagrass selections ................................. 37
Bahiagrass selections .................................... 39
Comparison of species and cultivars of turfgrass ........... 39
Pasture grass selections ................ ............... 40
Host resistance potential for grasses .................... 40








IV. CHEMICAL CONTROL OF MOLE CRICKETS ................ 41
Beginnings .............. ......................... 41
Present Day Control ...................................... 42
Baits ................ ................................. 42
Sprays and granules ..................................... 46
Evaluation of chemical control ............................ 46
Future Prospects ...................................... 47
Addendum ............... ........... ................. 48

V. FURTHER READING ................................... 49

REFERENCES .............................. ............ 51







Preface
This bulletin is a report of the first five years ofIFAS's special mole
cricket project. It summarizes what has been learned of mole cricket
biology and describes what now seem the most promising avenues to
reducing or eliminating mole cricket damage in Florida.
T. J. Walker (Editor of this
bulletin and Co-ordinator
of the Mole Cricket Project
1979-date)
December 1983











I. INTRODUCTION

The most important insect pests of turf and pastures in Florida are
mole crickets. And their damage is not restricted to grasses. Their
feeding and tunneling also destroy seedlings of vegetables,
ornamentals, and tobacco. Insecticides commonly applied to control
mole crickets in lawns, golf courses, and seed beds are expensive and
not always effective. In pastures there is no control that is economi-
cally feasible; yet without control, stands of pasture grasses are
frequently so reduced as to require replanting. The annual cost of
mole crickets to Floridians is no less than $30 million.
In 1978, in response to increasing concerns of cattlemen, turf
managers, and home owners, IFAS intensified its research on mole
crickets. The resulting project was partially funded by a continuing
special appropriation from the State Legislature. Its primary goal
was to eliminate mole crickets as a problem in Florida (for example,
by establishing self-sustaining natural enemies). Its fall-back goal
was to make temporary control of mole crickets so economical that
even pastures could be protected (for example, by reducing costs of
insecticidal control by an order of magnitude). Because little was
known of mole cricket systematics, behavior, ecology, and population
dynamics, a third goal was given initial priority in order to improve
chances of achieving either of the other two-namely, ascertaining
fundamentals of mole cricket biology.
This bulletin summarizes the results of the first five years of
IFAS's mole cricket project. It describes the biology of pest mole
crickets, including new discoveries that helped delineate promising
pathways toward lessening the impact of mole crickets in Florida.
Under the headings of Biological Control and Chemical Control it
discusses progress and prospects relative to specific permanent and
temporary solutions. The three that now seem most promising are (1)
introduction of natural enemies from South America, (2) substitution
of resistant grasses for susceptible ones, and (3) development of a
cheap, safe, effective, insecticidal bait.

History
None of the three species of pest mole crickets in Florida is native.
All were accidentally introduced to the southeastern United States
about 80 years ago. By the 1930s they had spread to the vegetable
growing areas of central Florida. Their effects on truck crops became











I. INTRODUCTION

The most important insect pests of turf and pastures in Florida are
mole crickets. And their damage is not restricted to grasses. Their
feeding and tunneling also destroy seedlings of vegetables,
ornamentals, and tobacco. Insecticides commonly applied to control
mole crickets in lawns, golf courses, and seed beds are expensive and
not always effective. In pastures there is no control that is economi-
cally feasible; yet without control, stands of pasture grasses are
frequently so reduced as to require replanting. The annual cost of
mole crickets to Floridians is no less than $30 million.
In 1978, in response to increasing concerns of cattlemen, turf
managers, and home owners, IFAS intensified its research on mole
crickets. The resulting project was partially funded by a continuing
special appropriation from the State Legislature. Its primary goal
was to eliminate mole crickets as a problem in Florida (for example,
by establishing self-sustaining natural enemies). Its fall-back goal
was to make temporary control of mole crickets so economical that
even pastures could be protected (for example, by reducing costs of
insecticidal control by an order of magnitude). Because little was
known of mole cricket systematics, behavior, ecology, and population
dynamics, a third goal was given initial priority in order to improve
chances of achieving either of the other two-namely, ascertaining
fundamentals of mole cricket biology.
This bulletin summarizes the results of the first five years of
IFAS's mole cricket project. It describes the biology of pest mole
crickets, including new discoveries that helped delineate promising
pathways toward lessening the impact of mole crickets in Florida.
Under the headings of Biological Control and Chemical Control it
discusses progress and prospects relative to specific permanent and
temporary solutions. The three that now seem most promising are (1)
introduction of natural enemies from South America, (2) substitution
of resistant grasses for susceptible ones, and (3) development of a
cheap, safe, effective, insecticidal bait.

History
None of the three species of pest mole crickets in Florida is native.
All were accidentally introduced to the southeastern United States
about 80 years ago. By the 1930s they had spread to the vegetable
growing areas of central Florida. Their effects on truck crops became








so severe that the USDA undertook research at Sanford (1934-1939),
culminating in the distribution, in 1940, of 1,258 tons of arsenic bait
for mole cricket control in 12 Florida counties. Research and control
efforts were interrupted by World War II, but shortly afterwards,
mole crickets were effectively controlled by newly developed, cheap,
long-lasting insecticides-especially DDT and chlordane. Relief was
short-lived. Because of hazards to man and his environment, persist-
ent pesticides were banned or severely restricted, and the insecticides
permitted for mole cricket control became more expensive and less
lasting. At the same time, cattlemen in Florida were spending heav-
ily to change raw rangeland into improved bahiagrass pastures-a
favored food for mole crickets. By the mid '70s, mole crickets were
once again out of control. Pastures were being lost with no economic
recourse; turf managers were being forced to use increasingly costly
insecticides with increasingly uncertain results. This was the situa-
tion that prompted the research reported here.

Acknowledgments
Studies of pest mole crickets have been accelerated recently by a
cooperative agreement (7005-20240-022A) with USDA, a special
USDA grant (83-CRSR-2-2162), and a grant from the Florida Turf-
Grass Association.
During the past five years, 43 researchers have participated in the
IFAS mole cricket project. Eight of these are authors of sections of
this bulletin. The others are: C. S. Barfield, H. W. Beck, Guy Beug-
non, F. G. Bliz, D. G. Boucias, J. L. Castner, P. P. Cobb (Auburn
University), H. L. Cromroy, H. G. Fowler, G. N. Fritz, M. E. Green, P.
G. Koehler, K. O. Lawrence (Chemlawn Corp.), R. C. Littell, R. E.
Lynch (USDA), E. L. Matheny, Jr., C. M. McCoy, J. L. Nation, D. A.
Nickle (USDA), Ngo Dong, S. L. Poe, K. M. Portier, D. J. Schuster, K.
C. Shaw, L. N. Shaw, D. E. Short, A. Silveira-Guido, B. J. Smittle
(USDA), Bill Stackhouse, L. A. Stange (Fla. Div. Plant Ind.), Abate
Tsedeke, S. L. Walker, J. J. Williams, Sue Wineriter, and R. E.
Woodruff.










II. BIOLOGY OF PEST MOLE CRICKETS

Systematics and Life Cycles (T. J. Walker)

Species and their identification
Four species of mole crickets occur in Florida (Fig. 1). The northern
mole cricket, Neocurtilla hexadactyla, is a native species of no eco-
nomic importance. It is recognized by its four tibial dactyls (Fig. 3b).
The other three species are pests, have two tibial dactyls, and belong
to the genus Scapteriscus (Fig. 3c-e).
The tawny mole cricket, S. vicinus, has the tibial dactyls nearly
touching at their bases (Fig. 3c). The southern mole cricket, S. acle-
tus, and the short-winged mole cricket, S. abbreviatus, have a distinct
gap between the tibial dactyls, and in the latter species the dactyls
are slightly divergent (Fig. 3d, e). Other means of distinguishing the
species of Scapteriscus are the length of the trochantal blade (Fig.
4a-d), the dorsal pronotal pattern (Fig. 5a-d), and the mottling of the
hind legs (characteristic of the short-winged mole cricket). Only in
the short-winged mole cricket are the adult wings shorter than the
pronotum. (Large juveniles of other mole crickets also have short
wings, but the right and left ones do not overlap.)

Geography
Florida's three pest mole crickets were inadvertently introduced to
the southeastern United States from South America about 1900. It
seems likely that they traveled in ship's ballast that was dumped in
preparation for taking on heavy loads of timber or naval stores. The
short-winged mole cricket was first collected at Tampa and was
shortly thereafter taken in the vicinity of three other, widely sepa-
rated ports. It has spread slowly and is largely restricted to coastal
areas (Fig. 8).
The tawny mole cricket was first recorded at Brunswick, Georgia, a
major seaport at the time. It gradually spread, reaching south-
ernmost and westernmost Florida around 1960. The species is con-
tinuing to move westward along the Gulf (Fig. 7).
The southern mole cricket was introduced at no fewer than four
ports: Brunswick, in 1904; Charleston, in 1915; Mobile, in 1919; and
Port Arthur, Texas, in 1925 (Fig. 6b). The crickets introduced at
Brunswick and Mobile had mottled pronotums (Fig. 5b); those intro-
duced at the other two ports had dark pronotums with four pale spots













































le

Fig. 1. Florida mole crickets, a. Southern mole cricket. b. Tawny mole cricket.
c. Short-winged mole cricket. d. Northern mole cricket. e. Comparison of the four species.
(Photos by J. L. Castner)
^B^B is A^^^^
*LI J~fl' j^^r
?1~~~~~ "<-'B '^^^*'












(Photos by J. L. Castner)











Fig. 2-5. Identifying
features of Florida mole
crickets.


-pronotal pattern (Fig. 5a-d)


juvenile


3b

tympanum



rigt Northern
foreleg left foreleg
pulled downward 3d






Southern


Short-winged


Short-winged


4b




Tawny


Southern


5a 5b /










Southern Southern
(4-dot) (mottled)


Shortwinged
Short-winged


5d rt-wi










Short-winged


adult male










Key to Maps
Distribution by
E31910 E31930 E11980
Distribution of Southern
pronotal patterns
D4-dot E mottled


Southern


Fig. 6-9. Distribution of southeastern mole crickets and history of spread of pest species.


Tawny










II. BIOLOGY OF PEST MOLE CRICKETS

Systematics and Life Cycles (T. J. Walker)

Species and their identification
Four species of mole crickets occur in Florida (Fig. 1). The northern
mole cricket, Neocurtilla hexadactyla, is a native species of no eco-
nomic importance. It is recognized by its four tibial dactyls (Fig. 3b).
The other three species are pests, have two tibial dactyls, and belong
to the genus Scapteriscus (Fig. 3c-e).
The tawny mole cricket, S. vicinus, has the tibial dactyls nearly
touching at their bases (Fig. 3c). The southern mole cricket, S. acle-
tus, and the short-winged mole cricket, S. abbreviatus, have a distinct
gap between the tibial dactyls, and in the latter species the dactyls
are slightly divergent (Fig. 3d, e). Other means of distinguishing the
species of Scapteriscus are the length of the trochantal blade (Fig.
4a-d), the dorsal pronotal pattern (Fig. 5a-d), and the mottling of the
hind legs (characteristic of the short-winged mole cricket). Only in
the short-winged mole cricket are the adult wings shorter than the
pronotum. (Large juveniles of other mole crickets also have short
wings, but the right and left ones do not overlap.)

Geography
Florida's three pest mole crickets were inadvertently introduced to
the southeastern United States from South America about 1900. It
seems likely that they traveled in ship's ballast that was dumped in
preparation for taking on heavy loads of timber or naval stores. The
short-winged mole cricket was first collected at Tampa and was
shortly thereafter taken in the vicinity of three other, widely sepa-
rated ports. It has spread slowly and is largely restricted to coastal
areas (Fig. 8).
The tawny mole cricket was first recorded at Brunswick, Georgia, a
major seaport at the time. It gradually spread, reaching south-
ernmost and westernmost Florida around 1960. The species is con-
tinuing to move westward along the Gulf (Fig. 7).
The southern mole cricket was introduced at no fewer than four
ports: Brunswick, in 1904; Charleston, in 1915; Mobile, in 1919; and
Port Arthur, Texas, in 1925 (Fig. 6b). The crickets introduced at
Brunswick and Mobile had mottled pronotums (Fig. 5b); those intro-
duced at the other two ports had dark pronotums with four pale spots










II. BIOLOGY OF PEST MOLE CRICKETS

Systematics and Life Cycles (T. J. Walker)

Species and their identification
Four species of mole crickets occur in Florida (Fig. 1). The northern
mole cricket, Neocurtilla hexadactyla, is a native species of no eco-
nomic importance. It is recognized by its four tibial dactyls (Fig. 3b).
The other three species are pests, have two tibial dactyls, and belong
to the genus Scapteriscus (Fig. 3c-e).
The tawny mole cricket, S. vicinus, has the tibial dactyls nearly
touching at their bases (Fig. 3c). The southern mole cricket, S. acle-
tus, and the short-winged mole cricket, S. abbreviatus, have a distinct
gap between the tibial dactyls, and in the latter species the dactyls
are slightly divergent (Fig. 3d, e). Other means of distinguishing the
species of Scapteriscus are the length of the trochantal blade (Fig.
4a-d), the dorsal pronotal pattern (Fig. 5a-d), and the mottling of the
hind legs (characteristic of the short-winged mole cricket). Only in
the short-winged mole cricket are the adult wings shorter than the
pronotum. (Large juveniles of other mole crickets also have short
wings, but the right and left ones do not overlap.)

Geography
Florida's three pest mole crickets were inadvertently introduced to
the southeastern United States from South America about 1900. It
seems likely that they traveled in ship's ballast that was dumped in
preparation for taking on heavy loads of timber or naval stores. The
short-winged mole cricket was first collected at Tampa and was
shortly thereafter taken in the vicinity of three other, widely sepa-
rated ports. It has spread slowly and is largely restricted to coastal
areas (Fig. 8).
The tawny mole cricket was first recorded at Brunswick, Georgia, a
major seaport at the time. It gradually spread, reaching south-
ernmost and westernmost Florida around 1960. The species is con-
tinuing to move westward along the Gulf (Fig. 7).
The southern mole cricket was introduced at no fewer than four
ports: Brunswick, in 1904; Charleston, in 1915; Mobile, in 1919; and
Port Arthur, Texas, in 1925 (Fig. 6b). The crickets introduced at
Brunswick and Mobile had mottled pronotums (Fig. 5b); those intro-
duced at the other two ports had dark pronotums with four pale spots










II. BIOLOGY OF PEST MOLE CRICKETS

Systematics and Life Cycles (T. J. Walker)

Species and their identification
Four species of mole crickets occur in Florida (Fig. 1). The northern
mole cricket, Neocurtilla hexadactyla, is a native species of no eco-
nomic importance. It is recognized by its four tibial dactyls (Fig. 3b).
The other three species are pests, have two tibial dactyls, and belong
to the genus Scapteriscus (Fig. 3c-e).
The tawny mole cricket, S. vicinus, has the tibial dactyls nearly
touching at their bases (Fig. 3c). The southern mole cricket, S. acle-
tus, and the short-winged mole cricket, S. abbreviatus, have a distinct
gap between the tibial dactyls, and in the latter species the dactyls
are slightly divergent (Fig. 3d, e). Other means of distinguishing the
species of Scapteriscus are the length of the trochantal blade (Fig.
4a-d), the dorsal pronotal pattern (Fig. 5a-d), and the mottling of the
hind legs (characteristic of the short-winged mole cricket). Only in
the short-winged mole cricket are the adult wings shorter than the
pronotum. (Large juveniles of other mole crickets also have short
wings, but the right and left ones do not overlap.)

Geography
Florida's three pest mole crickets were inadvertently introduced to
the southeastern United States from South America about 1900. It
seems likely that they traveled in ship's ballast that was dumped in
preparation for taking on heavy loads of timber or naval stores. The
short-winged mole cricket was first collected at Tampa and was
shortly thereafter taken in the vicinity of three other, widely sepa-
rated ports. It has spread slowly and is largely restricted to coastal
areas (Fig. 8).
The tawny mole cricket was first recorded at Brunswick, Georgia, a
major seaport at the time. It gradually spread, reaching south-
ernmost and westernmost Florida around 1960. The species is con-
tinuing to move westward along the Gulf (Fig. 7).
The southern mole cricket was introduced at no fewer than four
ports: Brunswick, in 1904; Charleston, in 1915; Mobile, in 1919; and
Port Arthur, Texas, in 1925 (Fig. 6b). The crickets introduced at
Brunswick and Mobile had mottled pronotums (Fig. 5b); those intro-
duced at the other two ports had dark pronotums with four pale spots








arranged in a trapezoid (Figs. 5a and la). The present distribution of
mottled and four-dot forms is concordant with the original introduc-
tions except that peninsular Florida is occupied by the four-dot form
(either through spread southward along the coast from Charleston
or from an independent introduction at Jacksonville around 1924)
(Fig. 6a).
The exact origin of the Southeast's mole crickets is uncertain, but
the ports of Montevideo, Uruguay, and Buenos Aires, Argentina,
seem likely. Tawny and southern mole crickets are widespread in
northern Argentina, Uruguay, and southern Brazil. (Neither occurs
in the West Indies, although until recently the tawny mole cricket
was known as "Puerto Rican mole cricket" or changea.")

Life cycles
Mole crickets spend nearly all their lives underground. They begin
as eggs laid in clutches in underground chambers. The hatchlings
soon tunnel to the surface and feed in the upper soil and litter. During
the subsequent juvenile and adult stages, the crickets make and
occupy extensive tunnel systems. The number of juvenile stages
(separated by molts) has not been studied, but there are at least 6 or 7
stages. All juvenile stages resemble adults but lack wings (Fig. 2b)
until the last two juvenile stages, when noticeable wing buds appear.
Adult males differ from females in having their forewings specialized
for calling. The easiest-to-spot distinguishing feature of males is a
harp-shaped resonating cell (Fig. 2a).
From central Florida northward, tawny and southern mole crick-
ets have one generation each year. Most individuals of the former
species overwinter as adults, and most of the latter species overwin-
ter as large juveniles (Fig. 10). In southern Florida the tawny mole
cricket maintains its one-year life cycle (as in Fig. 10) but the south-
ern mole cricket has two generations annually. Geographical varia-
tion in the seasonal timing and number of annual generations is
evident from peaks of flight activity of adults (Fig. 11).
The life cycle of the short-winged mole crickets is not as well
understood. Apparently all stages occur at all seasons but with a
peak of egg laying in late spring or summer and a lesser peak in
winter.

Dispersal
Tawny and southern mole crickets sometimes fly in enormous
numbers. Their flights apparently serve two functions: (1) local








arranged in a trapezoid (Figs. 5a and la). The present distribution of
mottled and four-dot forms is concordant with the original introduc-
tions except that peninsular Florida is occupied by the four-dot form
(either through spread southward along the coast from Charleston
or from an independent introduction at Jacksonville around 1924)
(Fig. 6a).
The exact origin of the Southeast's mole crickets is uncertain, but
the ports of Montevideo, Uruguay, and Buenos Aires, Argentina,
seem likely. Tawny and southern mole crickets are widespread in
northern Argentina, Uruguay, and southern Brazil. (Neither occurs
in the West Indies, although until recently the tawny mole cricket
was known as "Puerto Rican mole cricket" or changea.")

Life cycles
Mole crickets spend nearly all their lives underground. They begin
as eggs laid in clutches in underground chambers. The hatchlings
soon tunnel to the surface and feed in the upper soil and litter. During
the subsequent juvenile and adult stages, the crickets make and
occupy extensive tunnel systems. The number of juvenile stages
(separated by molts) has not been studied, but there are at least 6 or 7
stages. All juvenile stages resemble adults but lack wings (Fig. 2b)
until the last two juvenile stages, when noticeable wing buds appear.
Adult males differ from females in having their forewings specialized
for calling. The easiest-to-spot distinguishing feature of males is a
harp-shaped resonating cell (Fig. 2a).
From central Florida northward, tawny and southern mole crick-
ets have one generation each year. Most individuals of the former
species overwinter as adults, and most of the latter species overwin-
ter as large juveniles (Fig. 10). In southern Florida the tawny mole
cricket maintains its one-year life cycle (as in Fig. 10) but the south-
ern mole cricket has two generations annually. Geographical varia-
tion in the seasonal timing and number of annual generations is
evident from peaks of flight activity of adults (Fig. 11).
The life cycle of the short-winged mole crickets is not as well
understood. Apparently all stages occur at all seasons but with a
peak of egg laying in late spring or summer and a lesser peak in
winter.

Dispersal
Tawny and southern mole crickets sometimes fly in enormous
numbers. Their flights apparently serve two functions: (1) local







Tawny mole cricket
T1111111111T1111111 IIT IIIII


Southern mole cricket







Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Fig. 10. Seasonal distribution of stages of tawny and southern mole crickets in northern and central Florida. Note the
difference between the two species in the proportion of overwintering crickets that are juveniles.









Tawny mole cricket


Southern mole cricket


* **' ASO N J AFM A 'M J J


ASON DJF MA'M ''JS 0N'DJ'F'M AM J'JA'





jN* D7^ JF A '


A'' ON DJ FM A M'JJ 'S'O'N'D'J'F'M'A'M '


Fig.11. Seasonality ofmole cricket flights in northern, central, and southern Florida. Small fall flights precede larger spring flights of
adults of a single generation (Fig. 10). In southern Florida, summer flights of southern mole crickets represent a second generation of
adults (hatched area).








searching for mates and new egg-laying or calling sites (see next
section) and (2) long range dispersal.
Evidence for dispersal flights can be found in the range expansion
of the two species since their introduction (Figs. 6, 7). More direct
proof of such flights is the large numbers of mole crickets landing in
areas far from where they developed-for example, in extensive
woods or on ships offshore. The fact that flying mole crickets are
attracted to the calling songs of males of the same species makes
possible the use of "sound traps" to sample mole cricket flights over
wooded areas as well as over their breeding areas (see p. 19). Calcula-
tions based on such samples yield estimates that 10 to 20% of south-
ern mole cricket flights and 40% or more of tawny mole cricket flights
are beyond the bounds of the breeding area of origin.
Flight range is uncertain, but marked mole crickets have been
recovered up to 3.8 km (2.4 miles) from the point of release. Max-
imum flight distances are surely farther. The economic impact of
dispersal flights is that areas freed of mole crickets are rapidly
reinfested and newly cleared land quickly acquires an inoculum of
mole crickets.


Reproductive Behavior (T. G. Forrest)

Male calling
Adult male mole crickets, like most other crickets, produce a call-
ing song that functions to bring the sexes together. Females some-
times respond to the call by walking or flying to the male (phonotaxis)
and mating with him. Songs of crickets inhabiting the same area at
the same time are usually species-typical-that is, each species has
its own calling song and a trained ear can easily tell the species of
cricket by hearing its song. Tawny and southern mole cricket calling
songs are continuous trills that differ in both the tone (carrier fre-
quency; 3.3 vs. 2.7 kHz respectively) and pulse rate (wingstroke rate;
130 vs. 50 pulses per sec) of the song. Songs of tawny mole crickets
are characteristically "buzzy" and have short intermittent silent
periods of less than 1 second. Southern mole cricket songs are more
musical and continuous. Both songs can be heard soon after sunset
coming from the ground where males call from specially excavated
chambers.
The calling chamber opens at the soil surface through a trumpet-
like (exponentially expanding) horn. This chamber is constructed
each evening 10 to 20 minutes before calling and is tuned by the
cricket to the frequency of his calling song. The tuning increases the








searching for mates and new egg-laying or calling sites (see next
section) and (2) long range dispersal.
Evidence for dispersal flights can be found in the range expansion
of the two species since their introduction (Figs. 6, 7). More direct
proof of such flights is the large numbers of mole crickets landing in
areas far from where they developed-for example, in extensive
woods or on ships offshore. The fact that flying mole crickets are
attracted to the calling songs of males of the same species makes
possible the use of "sound traps" to sample mole cricket flights over
wooded areas as well as over their breeding areas (see p. 19). Calcula-
tions based on such samples yield estimates that 10 to 20% of south-
ern mole cricket flights and 40% or more of tawny mole cricket flights
are beyond the bounds of the breeding area of origin.
Flight range is uncertain, but marked mole crickets have been
recovered up to 3.8 km (2.4 miles) from the point of release. Max-
imum flight distances are surely farther. The economic impact of
dispersal flights is that areas freed of mole crickets are rapidly
reinfested and newly cleared land quickly acquires an inoculum of
mole crickets.


Reproductive Behavior (T. G. Forrest)

Male calling
Adult male mole crickets, like most other crickets, produce a call-
ing song that functions to bring the sexes together. Females some-
times respond to the call by walking or flying to the male (phonotaxis)
and mating with him. Songs of crickets inhabiting the same area at
the same time are usually species-typical-that is, each species has
its own calling song and a trained ear can easily tell the species of
cricket by hearing its song. Tawny and southern mole cricket calling
songs are continuous trills that differ in both the tone (carrier fre-
quency; 3.3 vs. 2.7 kHz respectively) and pulse rate (wingstroke rate;
130 vs. 50 pulses per sec) of the song. Songs of tawny mole crickets
are characteristically "buzzy" and have short intermittent silent
periods of less than 1 second. Southern mole cricket songs are more
musical and continuous. Both songs can be heard soon after sunset
coming from the ground where males call from specially excavated
chambers.
The calling chamber opens at the soil surface through a trumpet-
like (exponentially expanding) horn. This chamber is constructed
each evening 10 to 20 minutes before calling and is tuned by the
cricket to the frequency of his calling song. The tuning increases the









efficiency of sound production by at least threefold. Intensities of
calling songs range from 50 to 90 dB (relative to 20pN/m2) at 15 cm
and are dependent upon male size and soil moisture.
The calling period is correlated with female activity and flight
periods. Male tawny mole crickets begin calling 10 to 20 minutes
after sunset, and southern mole cricket males start 15 minutes later.
Calling continues for about one hour (Fig. 12).


Flight

About the same time in the evening that males are building their
calling chambers, many females and some males are preparing for
flight. A small opening is made at the soil surface where the indi-
vidual returns periodically, seemingly to check weather conditions.


a
- calling



Tawny mole cricket
take-off

landing

10 20 30 40 50 60 70 80 0 100 110
Minutes after Sunset



b




Southern mole cricket
take-off
landing

10 20 30 40 50 60 70 80 90 100 110
Minutes after Sunset
Fig. 12. Timing of flight and calling for (a) tawny and (b) southern mole
crickets. Dark bars indicate when males of each species call. Unshaded
curves show the distribution of take-off times; shaded curves show the dis-
tribution of landing times. Both species fly and call for little more than an
hour beginning shortly after sunset, with tawny mole crickets preceding
southern mole crickets by 5 to 15 minutes.








Just prior to flight, while remaining inside its tunnel, the cricket
warms its flight muscles by trembling. When the muscles are
warmed and all is clear, the cricket runs from the burrow and im-
mediately takes to the air. Crickets usually remain in flight 5 to 20
minutes, but some may fly more than 40 minutes, perhaps traveling
as far as 8 km (5 miles). Flight usually ends in response to the male
calling song (Fig. 12); however, crickets are also attracted to and land
near lights.

Phonotaxis and mate choice
Mole crickets calling near one another must compete acoustically
for flying females, and females hear many calling males during a
single flight. If differences in calling songs are cues to differences
among males that are related to increases in female reproduction,
females should respond to and mate with some males more than
others. In general, females respond only to calling songs of their own
species, but within each species some males' songs are more attrac-
tive than others. A single male may attract as many as 27 females in
one calling period, while other males calling nearby attract few or
none. One aspect of the calling song that enables females to discrim-
inate is intensity. Flying mole crickets prefer to land at louder calls.
Louder males attract more females (and males, see below) than less
intense males, and the effect is greater than would be expected just
because a louder call travels farther.
Two factors that influence the loudness of a male's call are his size
and the moisture content of the soil used to construct his calling
chamber. Other things being equal, a large male produces a louder
calling song and attracts more females than smaller males. Thus by
going to louder calls, females choose to mate with larger males.
However, only about 30% of attracted females enter the burrow of the
calling male whose song triggered their landing. The others land
nearby and dig their own burrows, probably using the loudness of the
call to locate moist soil for oviposition. After the calling period is over,
males (those that have called and those that have landed near the
caller) probably search for, court, and mate with these females (Fig.
13).
Males also fly and respond to calling songs in much the same
manner as females; that is, they preferentially land at louder calling
songs. They thereby find an area where many females may have
landed. Such males sometimes enter the caller's burrow and fight
with him. Males may also use the louder calls to detect areas with
high soil moisture that will make their own calls louder and enhance
their ability to attract females.












too dry -
other 99
no food
(1-2 days)


9 molts to adult*


S--- 9 takes off


passed


if available
9 lands at max cf


t
9 moves directly
to f burrow


9 enters df
burrow

d continues calling

9 matures eggs* d stops calling 9 prevents c
10-14 days begins courting from calling

bad weather oY begins courting
good habitat
I 1 end of season mates with


9 lays eggs |
C forces 9 out
when she ceases
to accept spermatophores
(probably <24 hrs.)


9 leaves d
burrow


Fig. 13. Female reproductive behaviors. Behaviors indicated by heavier arrows are
better documented than those indicated by lighter arrows. (*Females may mate with
males that find them in their burrows.)








Mating behavior
Since most of the sexual behavior of mole crickets is subterranean,
few observations of their mating behavior have been made. Whether
or not males and females find each other by means other than phono-
taxis is not known, but courtship songs are heard at all times of the
day, indicating that phonotaxis does not always immediately precede
courtship and mating.
When males and females meet in a burrow, they often antennate
each other. Males may produce several sharp "aggressive" chirps
followed by softer courtship chirps. If the female is unreceptive, the
two will fight. During these bouts males and females produce aggres-
sive chirps, and bite and claw each other with forelegs until one is
driven from the burrow. If, on the other hand, the female is receptive,
the male will continue courting her acoustically, his forewings
slightly raised, producing rhythmic chirps often followed by short
trilling sequences. The courtship songs have similar tones and pulse
rates to the calling songs. The rate of the chirps varies in the courting
sequence and may depend on the female's behavior. Male tawny mole
crickets often hit their pronotum against the roof of the burrow
producing a rapid thumping sound (15 to 20 hits per second, 5 to 10
hits per burst, repeated at 1 to 2 second intervals). Pronotal thump-
ing may convey special information to the female or other nearby
crickets. It has not been observed in southern mole cricket males or in
females of either species.
If the male is facing the female in the burrow, he backs up while
continuing his courtship song. The female follows. The male, still
courting, turns to face away from the female. The male sometimes
pushes soil ahead of him in the burrow, enlarging the tunnel to make
room for the female to mount. As the female approaches, the male
lowers his wings and tries to back under her.
Coupling does not occur until about 40 seconds after mounting. At
this time a small spermatophore is extruded from the end of the
male's abdomen. The pair couple briefly, the spermatophore is passed
to the female, and she dismounts. The spermatophore is white and
ovate, about 1 mm in diameter, with a short curved tube on one side.
The tube is inserted into the female, and the sperm empties from the
spermatophore through the tube into her spermatheca, or sperm
storage organ. The small spermatophore is completely concealed
under the female's subgenital plate to protect it from being dislodged
by the tunnel walls.
Once the female dismounts, the male remains stationary for 8 to 10
minutes and then resumes courting. Sometimes he stops to enlarge
the tunnel. This stationary period is needed for the male to form a








new spermatophore. During the male's inactivity the female moves
about the tunnel, feeds, or grooms. When the male begins the next
courtship sequence, the female removes the spermatophore and may
eat it. The female may mount and copulate with the same male
several times, but once the female becomes unresponsive to the
male's courtship attempts, they fight until one of the pair leaves the
burrow system. During her adult life a female usually mates with
more than one male.

Egg laying and hatching

A male's parental duties are over after mating, but the female
must find suitable oviposition sites. Mole cricket dispersal flights are
adaptive in that already mated females can leave unsuitable areas
(for example, areas with poor food, high density of crickets, or low soil
moisture), locate, land, and oviposit in newly opened habitats. These
flights have contributed to the mole crickets' range expansion and
increasing pest status in the southeastern United States.
By landing near calling males, females can locate suitable mole
cricket habitats. Flying females are generally in an early stage of a
reproductive (clutch) cycle. Their primary oocytes are small, suggest-
ing that an egg clutch was laid just prior to flight. After landing, the
female spends 9 to 14 days maturing oocytes and then lays a clutch of
eggs (Fig. 13).
About 40 (25 to 60 depending on female size and age) gray or
brownish eggs are laid in a small ovoid (4 x 3 cm) chamber or egg cell.
Egg cells are constructed by females less than 24 hours before ovi-
position at depths of 9 to 30 cm, depending on soil type and moisture.
Once oviposition is complete, the female seals the entrance to the
chamber with soil and has no more contact with eggs or young.
Females sometimes lay more than one clutch before they fly again
and may mate with another male between egg clutches.
During the 20 days it takes the eggs to develop, they double in
volume from water absorption, and their color changes from gray to
milky white. All eggs of a clutch usually hatch within a 24-hour
period. Hatchlings are white or yellowish and turn bluish-black
within 24 to 48 hours. Juveniles often eat the egg shells (chorion) and
are known to cannibalize siblings. They escape from the egg cell
through a small tunnel dug straight up to the soil surface.









Other Behavior, Damage, And Sampling
(W. G. Hudson)

Feeding
For years it was assumed that all pest mole crickets were herbiv-
orous, but recent studies have shown this to be untrue. Examination
of gut contents indicates that diet of tawny and short-winged mole
crickets is mostly vegetarian. On the other hand, southern mole
crickets feed mostly on animal material (Fig. 14). Little is known
about the kinds of prey taken by southern mole crickets, but presum-
ably it includes a variety of soil inhabiting insects and other small
animals. Southern mole crickets eat some plant material, perhaps
when animal material is in short supply.
Mole crickets feed on a wide variety of plants and have caused
extensive damage to seedlings of tobacco, ornamentals, tomatoes and
other vegetables, and to sugarcane. The most serious damage in
recent years has been to pasture and turfgrass throughout the state.
All parts of the plant are eaten. At night, crickets often leave their
burrows to feed on above-ground parts, biting off stems and leaves,
which are dragged into the burrows to be eaten. Roots may be eaten
at any time.
Mole crickets have also been reported to feed on the underground
tubers of potato, carrot, and sweet potato, and on developing peanuts.

Burrowing
There is little quantitative information on burrowing activity of
mole crickets, although some "general conclusions about their sub-



Tawny Short-winged Southern


9% animal


88% plant% plant animal


Fig. 14. Feeding habits of pest mole crickets, indicated by examination of
gut contents. Note that tawny and short-winged mole crickets are mainly
herbivorous, whereas southern mole crickets are mainly carnivorous.









Other Behavior, Damage, And Sampling
(W. G. Hudson)

Feeding
For years it was assumed that all pest mole crickets were herbiv-
orous, but recent studies have shown this to be untrue. Examination
of gut contents indicates that diet of tawny and short-winged mole
crickets is mostly vegetarian. On the other hand, southern mole
crickets feed mostly on animal material (Fig. 14). Little is known
about the kinds of prey taken by southern mole crickets, but presum-
ably it includes a variety of soil inhabiting insects and other small
animals. Southern mole crickets eat some plant material, perhaps
when animal material is in short supply.
Mole crickets feed on a wide variety of plants and have caused
extensive damage to seedlings of tobacco, ornamentals, tomatoes and
other vegetables, and to sugarcane. The most serious damage in
recent years has been to pasture and turfgrass throughout the state.
All parts of the plant are eaten. At night, crickets often leave their
burrows to feed on above-ground parts, biting off stems and leaves,
which are dragged into the burrows to be eaten. Roots may be eaten
at any time.
Mole crickets have also been reported to feed on the underground
tubers of potato, carrot, and sweet potato, and on developing peanuts.

Burrowing
There is little quantitative information on burrowing activity of
mole crickets, although some "general conclusions about their sub-



Tawny Short-winged Southern


9% animal


88% plant% plant animal


Fig. 14. Feeding habits of pest mole crickets, indicated by examination of
gut contents. Note that tawny and short-winged mole crickets are mainly
herbivorous, whereas southern mole crickets are mainly carnivorous.









Other Behavior, Damage, And Sampling
(W. G. Hudson)

Feeding
For years it was assumed that all pest mole crickets were herbiv-
orous, but recent studies have shown this to be untrue. Examination
of gut contents indicates that diet of tawny and short-winged mole
crickets is mostly vegetarian. On the other hand, southern mole
crickets feed mostly on animal material (Fig. 14). Little is known
about the kinds of prey taken by southern mole crickets, but presum-
ably it includes a variety of soil inhabiting insects and other small
animals. Southern mole crickets eat some plant material, perhaps
when animal material is in short supply.
Mole crickets feed on a wide variety of plants and have caused
extensive damage to seedlings of tobacco, ornamentals, tomatoes and
other vegetables, and to sugarcane. The most serious damage in
recent years has been to pasture and turfgrass throughout the state.
All parts of the plant are eaten. At night, crickets often leave their
burrows to feed on above-ground parts, biting off stems and leaves,
which are dragged into the burrows to be eaten. Roots may be eaten
at any time.
Mole crickets have also been reported to feed on the underground
tubers of potato, carrot, and sweet potato, and on developing peanuts.

Burrowing
There is little quantitative information on burrowing activity of
mole crickets, although some "general conclusions about their sub-



Tawny Short-winged Southern


9% animal


88% plant% plant animal


Fig. 14. Feeding habits of pest mole crickets, indicated by examination of
gut contents. Note that tawny and short-winged mole crickets are mainly
herbivorous, whereas southern mole crickets are mainly carnivorous.








terranean behavior can be drawn. Under most conditions mole crick-
ets are found primarily in the top 20 to 25 cm (8 to 10 in.) of soil. This
is especially true ofjuveniles. In areas of high population density this
layer of soil is literally honeycombed with tunnels, through which the
mole crickets move forward and backward with equal agility. Tun-
nels as deep as 75 cm (30 in.) have been recorded, but these are
probably constructed only in times of extreme moisture or tempera-
ture stress. Surface burrows are usually the first indication of mole
cricket activity. These burrows look much like a miniature version of
the burrow of the common mole. Within turf or pasture, tawny mole
crickets tend to burrow in grassy areas, while southern mole crickets
prefer the bare, sandy areas. Burrowing may be so extensive that the
ground feels spongy to the step.
Although there may be movement at any time during the day or
night, the most active period is from late afternoon until near mid-
night. Mole crickets are least active during the middle of the
morning.


Damage
Mole crickets damage plants by feeding on them and by tunneling.
Feeding damage affects both underground and above-ground parts of
the plant. Heavily damaged turf or pasture grass has virtually no
root system and is easily pulled from the soil by grazing cattle or foot
traffic. The green shoots are also consumed, and even the tough,
fibrous stems are eaten in extreme cases. Damage to vegetable seed-
lings is often cutworm-like, although plants are generally severed at
or beneath rather than above the soil surface. Severed seedlings are
often pulled into tunnels where the foilage or entire plant may be
consumed. Damage is usually most severe in seedbeds or newly
transplanted fields.
Mechanical damage to plants is caused by the tunneling activity of
mole crickets and may be the principal detrimental effect of southern
mole crickets on grasses. There is evidence that all three mole crick-
ets cause damage to seedlings, probably because they damage the
developing root systems of the young plants by either feeding or
tunneling. Rhizoctonia root rot also increases in seedbeds infested
with mole crickets.
In most of Florida, damage to pasture and turfgrass is principally
due to feeding by tawny mole crickets. Southern mole crickets may
cause mechanical damage, especially in newly planted stands, but it
is doubtful that they cause significant damage to established stands.
The following evidence from several sources supports this conclusion.








1) Controlled experiments using known (and very high) densities of
mole crickets have repeatedly shown that southern mole crickets
have little effect on forage production. In no case have southern mole
crickets approached the destruction caused by tawny mole crickets in
the same experiments.
2) In field samples of pastures and turfgrass damaged by mole
crickets, southern mole crickets are either absent or greatly outnum-
bered by tawny mole crickets.
3) In areas of north Florida and south Alabama where southern
mole crickets have been present for years, mole crickets were not
considered pests until the tawny mole cricket extended its range into
those areas. Collection of mole crickets from damaged turf in Ala-
bama have included both species, but examinations of gut contents
have shown that southern mole crickets found in these incriminating
circumstances are still mostly carnivorous (two-thirds contain only
animal material) while tawny mole crickets are full of plant
material.
In areas where short-winged mole crickets occur (Fig. 8), they
contend with tawny mole crickets as the principal pest of grasses, and
they seem to be the only species attacking St. Augustinegrass.
Damage thresholds for mole crickets in turf and pastures have
been difficult to determine because of problems with sampling tech-
niques. Sampling in damaged and undamaged pastures using soil
flushing techniques (see below) has provided some idea of the popula-
tion density that can damage pasture grass. Severely damaged areas
of a pasture had an average sample density of 27 tawny mole crickets
per m2 (1 m2 = 11 sq.ft.). Undamaged areas of the same pasture had
an average of 12 tawnies per m2. Another bahiagrass pasture showed
some damage with a density of 9 tawnies per m2; the previous year
this pasture had 33 tawnies per m2 and had been severely damaged.
A nearby bermudagrass pasture that was beginning to suffer some
loss of stand had an average of 11 tawny mole crickets per m2.
Thresholds in turfgrass may be higher. Sampling with a tree spade
revealed 22 to 80 mole crickets per m2 in a heavily damaged golf
course.

Population estimation
The sketchy nature of our understanding of mole cricket ecology is
largely a result of problems encountered in sampling the insects in
the field. The following techniques have been (and continue to be)
employed, but none has yet overcome the main obstacle-how to
correlate sample numbers with true population density.








Sound traps These traps attract adults to the highly amplified
synthetic or recorded call of male mole crickets. A trap consists of a
caller situated over a collecting device that catches the crickets as
they land. The collecting device may be as simple as a child's wading
pool filled with water. The record catch for a "standard" sound trap
used by University of Florida researchers (with a 1.5 m diameter
wading pool as a collector) is 9000 mole crickets in one night. Sound
traps are effective only during the flight seasons in spring and fall
(Fig. 11), and only adults are taken. The numbers captured vary
widely from year to year (Table 1). Because so little is known about
how often, how far, and under what circumstances mole crickets fly,
it is difficult to interpret these variations. However, any lasting,
drastic reduction in mole cricket populations-such as hoped for
through the introduction of natural enemies (p. 23)-should be re-
vealed in a corresponding lasting, drastic reduction in trapping-
station catches.
Soil flushing The easiest and most economical method of field
sampling for mole crickets is flushing the soil with an aqueous solu-
tion of dishwashing liquid or insecticide. The two produce nearly



Table 1. Long-term variations in numbers of mole crickets captured at
standard sound-trapping stations at four sites.
Species and Gainesville- Gainesville- Ft.
Generation A C Bradenton Lauderdale
Southern mole
cricket
1978-79 2968 -
1979-80 5521 (26467)b -
1980-81 2292 8347 1376 479
1981-82 2043 5816 528 1065
1982-83 1783 12385 223 173
1983-84 4303 17358 2449 262
Tawny mole
cricket
1978-79 651 -
1979-80 1916 (1664)b -
1980-81 1898 1923 2313 265
1981-82 740 1037 3488 704
1982-83 2724 1800 4815 108
1983-84 10214 10265 1413 174
a August of the first year through July of the second. Except for southern mole crickets
in south Florida, this period includes a single generation (Fig. 22).
bPartial count, because station was not established until 1 Jan 1980.








identical results, making dishwashing liquid the better choice be-
cause it is cheaper and more readily available. The usual procedure is
to mix 15 mL of flushing agent with 4 liters of water, pour the solution
on the ground, and count the crickets as they surface. Four liters of
solution will cover an area about 0.5 m x 0.5 m, depending on soil
moisture, soil type, and ground cover. The sample density thus
obtained cannot at this point be called an estimate of true population
density (the relationship between sample numbers and true density
is unknown), but it is the best method available for comparing mole
cricket populations between locations or over time.
Pitfall traps Linear pitfall traps have shown promise in moni-
toring local population levels and represent one of the few sources of
live juvenile mole crickets for research use. Such traps are con-
structed by cutting a slot approximately 1 inch (2.5 cm) wide length-
wise in a section of 3-inch diameter PVC pipe. This pipe is then buried
in the ground so the slot is at or slightly below the soil surface, with
one end feeding into a 5-gallon bucket and the other end capped (Fig.
15). As with other sampling techniques, the relationship between
number taken and true mole cricket density is at present unknown.
Other Tawny and southern mole crickets can be captured at light
traps, sometimes in great numbers. However, lights do not attract as
many crickets as sound traps, and are also restricted to the flight
season. Digging after mole crickets, using tools ranging from shovels
to tree spades to specially constructed soil corers, has also been used
as a sampling technique but has not proved very effective. Shovels


Top View 1 / inch slot
end cap
.' -


5 gallon plastic pail

Side View
1 -/-/-F-c---

yj L soil level


-3 inch diameter PVC pipe
Fig. 15. Linear pitfall trap developed by K. O. Lawrence. Plastic jug
(dotted) can be placed in pail to facilitate removing the crickets (and other
animals).








are too slow to catch most of the mole crickets in an area, the tree
spade samples are of a volume that is difficult to work with, and the
core sampler provides results which are identical to soil flushing. All
are expensive in terms of time, manpower, and equipment and are
destructive to the pasture or turf being sampled. Estimates of tunnel-
ing activity have been used as population indicators, but so many
factors affect this activity (such as species of cricket, sex, size, soil
moisture, and temperature) that these estimates are of limited value.
Development of an effective sampling technique (or determining
the relationship between numbers sampled with existing techniques
and true density in a given area) remains a priority goal of mole
cricket research. Without such a technique, study of mole cricket
population ecology and scientific evaluation of potential control
methods is difficult.











III. BIOLOGICAL CONTROL
OF MOLE CRICKETS

Natural Enemies (R. I. Sailer)

Most organisms, whether plant or animal, have a variety of natu-
ral enemies. These include predators that consume their hosts and
parasites that develop in or on the body of individual hosts and cause
their death. There are also pathogens, such as certain fungi, bacteria,
and viruses, that infect and kill their hosts (see next section).
No predators are known to depend entirely on mole crickets for
their food supply, but in one genus of wasp the larvae of all species
must develop as parasites of mole crickets. Mole crickets are also
attacked by species of parasitic flies and by certain nematodes.

Predators
A surprisingly large number of vertebrate and invertebrate animals
have been observed to prey on mole crickets. Crop contents of birds
were studied in 1916 when mole crickets were very abundant in
Puerto Rico. Results indicated that a variety of birds ate significant
numbers of mole crickets and that over half the food of the Cuban
green heron and more than a quarter of the food of the Puerto Rican
sparrow hawk consisted of mole crickets. Toads, skunks, armadillos,
raccoons, and foxes have also been observed to eat mole crickets.
A number of predacious insects also capture and destroy many
mole crickets. These include carabid beetles of the genus Calosoma,
tiger beetles of the genus Tetracha, and an elaterid beetle belonging
to the genus Pyrophorus. Assassin bugs of the species Sirthenea
carinata have been reported to be abundant in areas where mole
crickets were numerous and have recently been observed to attack
and feed on these insects in the laboratory. Even the imported fire ant
acts as a predator on mole crickets.

Parasites
Before discussing parasites in greater detail, attention should be
called to some confusion relating to use of the term parasite. As used
by parasitologists the term is applied to organisms that live in or on
hosts without causing their death. The parasite is normally very
much smaller than its host and belongs to a very different taxonomic











III. BIOLOGICAL CONTROL
OF MOLE CRICKETS

Natural Enemies (R. I. Sailer)

Most organisms, whether plant or animal, have a variety of natu-
ral enemies. These include predators that consume their hosts and
parasites that develop in or on the body of individual hosts and cause
their death. There are also pathogens, such as certain fungi, bacteria,
and viruses, that infect and kill their hosts (see next section).
No predators are known to depend entirely on mole crickets for
their food supply, but in one genus of wasp the larvae of all species
must develop as parasites of mole crickets. Mole crickets are also
attacked by species of parasitic flies and by certain nematodes.

Predators
A surprisingly large number of vertebrate and invertebrate animals
have been observed to prey on mole crickets. Crop contents of birds
were studied in 1916 when mole crickets were very abundant in
Puerto Rico. Results indicated that a variety of birds ate significant
numbers of mole crickets and that over half the food of the Cuban
green heron and more than a quarter of the food of the Puerto Rican
sparrow hawk consisted of mole crickets. Toads, skunks, armadillos,
raccoons, and foxes have also been observed to eat mole crickets.
A number of predacious insects also capture and destroy many
mole crickets. These include carabid beetles of the genus Calosoma,
tiger beetles of the genus Tetracha, and an elaterid beetle belonging
to the genus Pyrophorus. Assassin bugs of the species Sirthenea
carinata have been reported to be abundant in areas where mole
crickets were numerous and have recently been observed to attack
and feed on these insects in the laboratory. Even the imported fire ant
acts as a predator on mole crickets.

Parasites
Before discussing parasites in greater detail, attention should be
called to some confusion relating to use of the term parasite. As used
by parasitologists the term is applied to organisms that live in or on
hosts without causing their death. The parasite is normally very
much smaller than its host and belongs to a very different taxonomic











III. BIOLOGICAL CONTROL
OF MOLE CRICKETS

Natural Enemies (R. I. Sailer)

Most organisms, whether plant or animal, have a variety of natu-
ral enemies. These include predators that consume their hosts and
parasites that develop in or on the body of individual hosts and cause
their death. There are also pathogens, such as certain fungi, bacteria,
and viruses, that infect and kill their hosts (see next section).
No predators are known to depend entirely on mole crickets for
their food supply, but in one genus of wasp the larvae of all species
must develop as parasites of mole crickets. Mole crickets are also
attacked by species of parasitic flies and by certain nematodes.

Predators
A surprisingly large number of vertebrate and invertebrate animals
have been observed to prey on mole crickets. Crop contents of birds
were studied in 1916 when mole crickets were very abundant in
Puerto Rico. Results indicated that a variety of birds ate significant
numbers of mole crickets and that over half the food of the Cuban
green heron and more than a quarter of the food of the Puerto Rican
sparrow hawk consisted of mole crickets. Toads, skunks, armadillos,
raccoons, and foxes have also been observed to eat mole crickets.
A number of predacious insects also capture and destroy many
mole crickets. These include carabid beetles of the genus Calosoma,
tiger beetles of the genus Tetracha, and an elaterid beetle belonging
to the genus Pyrophorus. Assassin bugs of the species Sirthenea
carinata have been reported to be abundant in areas where mole
crickets were numerous and have recently been observed to attack
and feed on these insects in the laboratory. Even the imported fire ant
acts as a predator on mole crickets.

Parasites
Before discussing parasites in greater detail, attention should be
called to some confusion relating to use of the term parasite. As used
by parasitologists the term is applied to organisms that live in or on
hosts without causing their death. The parasite is normally very
much smaller than its host and belongs to a very different taxonomic











III. BIOLOGICAL CONTROL
OF MOLE CRICKETS

Natural Enemies (R. I. Sailer)

Most organisms, whether plant or animal, have a variety of natu-
ral enemies. These include predators that consume their hosts and
parasites that develop in or on the body of individual hosts and cause
their death. There are also pathogens, such as certain fungi, bacteria,
and viruses, that infect and kill their hosts (see next section).
No predators are known to depend entirely on mole crickets for
their food supply, but in one genus of wasp the larvae of all species
must develop as parasites of mole crickets. Mole crickets are also
attacked by species of parasitic flies and by certain nematodes.

Predators
A surprisingly large number of vertebrate and invertebrate animals
have been observed to prey on mole crickets. Crop contents of birds
were studied in 1916 when mole crickets were very abundant in
Puerto Rico. Results indicated that a variety of birds ate significant
numbers of mole crickets and that over half the food of the Cuban
green heron and more than a quarter of the food of the Puerto Rican
sparrow hawk consisted of mole crickets. Toads, skunks, armadillos,
raccoons, and foxes have also been observed to eat mole crickets.
A number of predacious insects also capture and destroy many
mole crickets. These include carabid beetles of the genus Calosoma,
tiger beetles of the genus Tetracha, and an elaterid beetle belonging
to the genus Pyrophorus. Assassin bugs of the species Sirthenea
carinata have been reported to be abundant in areas where mole
crickets were numerous and have recently been observed to attack
and feed on these insects in the laboratory. Even the imported fire ant
acts as a predator on mole crickets.

Parasites
Before discussing parasites in greater detail, attention should be
called to some confusion relating to use of the term parasite. As used
by parasitologists the term is applied to organisms that live in or on
hosts without causing their death. The parasite is normally very
much smaller than its host and belongs to a very different taxonomic























































Fig. 16a-d. Larra bicolor, a digger wasp that parasitizes mole crickets. Fig. 16a.
Female laying egg on temporarily paralyzed mole cricket. Fig. 16b. Young larva feeding
ectoparasitically on active mole cricket host. Fig. 16c. Mature larva consuming remains
of host. Fig. 16d. Female feeding on flowers. (Photos by J. L. Castner)


24

























































25








group. Both adult and immature stages live in the host; for example,
round worms and tape worms that parasitize man and other verte-
brates. When entomologists use the term parasite, they generally are
referring to an organism that kills its host and is parasitic only in the
immature stages. The adults are free living, and most are insects that
attack other insects. The following discussion focuses on parasites of
this type.
Known parasites of mole crickets belong to three groups: digger
wasps of the genus Larra, parasitic flies of the genus Euphasiopteryx,
and entomophagous nematodes.
Digger wasps Wasps of the genus Larra (family Larridae) are
the most specialized natural enemies of mole crickets now known.
Unlike other digger wasps, which are generally predators, all species
are parasitic. Worldwide, 65 species of Larra are known. Most are
inhabitants of the tropics. Only one species, Larra analis, a parasite
of the northern mole cricket, is native to the United States. Sixteen
species have been recorded from South America, where the variety of
species of mole crickets belonging to the genus Scapteriscus is
greatest.
Much of what is known of the behavior and biology of larrid wasps
is the result of mole cricket invasions of Hawaii and Puerto Rico. In
Hawaii in the early part of this century, African mole crickets were
causing serious damage to newly planted sugarcane. When an en-
tomologist was sent to Australia, southeastern Asia, Africa, and
South America to find natural enemies suitable for introduction, he
discovered 12 species of Larra that develop as parasites of mole
crickets-2 from Australia, 3 from the Philippines, and 7 from South
America. One species, L. luzonensis, was successfully introduced into
Hawaii. Unfortunately, little is known of the consequences, other
than what might be inferred by diminished interest in and present
absence of concern about the economic importance of mole crickets in
Hawaii.
In Puerto Rico, mole crickets thought to have been introduced in
guano shipped from South America were the most injurious agricul-
tural pest from 1876 to 1902. They remained a serious problem at
least until the 1930s, when Puerto Rican entomologists introduced
Larra bicolor from Brazil. By 1938 this wasp was successfully estab-
lished, and by 1942 it occurred throughout the island. Again, there is
no documented assessment of results.
Apart from preferences for different species of mole crickets, the
biology of the various species of Larra is remarkably similar.
Females do not construct a nest, and prey paralysis is temporary with
the host reviving soon after egg deposition (Fig. 16a). The Larra larva








lives ectoparasitically until almost full grown (Fig. 16b). It then kills
and consumes the rest of its host (Fig. 16c). Larra females lay about
30 eggs in their life span. Full-grown larvae form underground co-
coons of sand grains cemented together with a secretion from the
labial glands. In the case of L. bicolor the adult wasps emerge from
their cells 60 to 80 days after eggs are laid on the hosts. At Fort
Lauderdale, where 88 females from Puerto Rico were released in
early June 1981, three generations were produced by December. In
1982, the first adults were seen in March. Thus, at least four genera-
tions are probably produced annually at that location.
The behavior of adults of several Larra species has been studied,
and all species exhibit the same general pattern. Females hunt for
mole crickets during early morning and mid-afternoon hours. During
the middle of the day they visit flowers where they feed on nectar and
possibly pollen (Fig. 16d). The different species not only have indi-
vidual preferences as to kind of mole cricket they parasitize, but they
also tend to frequent the flowers of different plant species. For exam-
ple, adults ofL. bicolor are most commonly found on the flowers of
two quite unrelated plants, Spermacoce verticillata (Rubeaceae) and
Hyptis atrorubens (Lamiaceae). The close association of L. bicolor
with these plants in Brazil and Puerto Rico suggests that it may not
inhabit areas where these plants are not available. At the site in Fort
Lauderdale where L. bicolor is now established, several small plots of
S. verticillata were planted and were blooming when the wasps were
first released. These plots have since been used to monitor the pres-
ence and increase of the wasp population.
Fortunately, S. verticillata is native to south Florida; however, in
order for L. bicolor to become an effective enemy of mole crickets in
central and northern Florida, it may be necessary to establish S.
verticillata or other suitable flowering plants in areas where mole
cricket populations are high.
Tachinid flies All species of true flies of the family Tachinidae
develop as parasites of other insects. Some groups of species are
restricted to closely related kinds of host insects; for example, the
group Ormiini consists of species that parasitize katydids and other
night-singing Orthoptera. The best known species of this group,
Euphasiopteryx ochracea, has been attracted to the tape-recorded
song of the southern mole cricket, though its normal hosts are field
crickets. Although larvae dissected from female E. ochracea and
placed on adult southern mole crickets have developed successfully,
none have been reared from field-collected mole crickets in the
United States. However, small numbers of a related species, E.
depleta, have been reared from mole crickets collected at Belem,








Brazil, and there is a high probability that other South American
species of the genus Euphasiopteryx are specialized as enemies of
mole crickets and can profitably be introduced into the United States.
Nematodes Among the several subgroups of nematodes that are
known to parasitize insects, the families Mermithidae and Steinerne-
matidae are especially likely to include natural enemies of mole
crickets.
The mermithids are widely distributed in aquatic and terrestrial
habitats and attack a wide range of insects and other arthropods. The
fact that certain genera are commonly restricted to specific groups of
insects suggests that a careful search of the homeland areas of Scap-
teriscus will reveal one or more species of mermithids that should be
introduced into Florida. Agameris decaudata, a parasite of grasshop-
pers, is an example of this group. After emerging from a dying host
they normally go into the soil, molt to the adult stage, and mate. The
female then lays eggs on vegetation to be eaten by feeding grasshop-
pers.
The insectivorous nematodes of the family Steinernematidae be-
have differently from the mermithids. Instead of developing singly in
hosts, the infective juvenile stage introduces a symbiotic bacterium
into the host. The bacterium causes death of the host in about 24
hours. The nematodes complete their development in the host ca-
daver and mate; the females lay very large numbers of eggs that
quickly hatch to produce second generation adults. The offspring of
these adults convert the host cadaver into a mass of juvenile nema-
todes that seek out new hosts. Within this group, species of the genus
Neoaplectana are probably of greatest interest for purposes of mole
cricket control.
In addition to recent discovery in South America of a mole cricket
infected by a species of Neoaplectana, there is another reason for
interest in these nematodes-they can be cultured on artificial
media, and one species, N. carpocapsae, is currently produced com-
mercially for control of soil insects and wood tunneling insect larvae.
Research is now in progress to determine whether strains of N.
carpocapsae or other species of Neoaplectana may prove useful when
applied as a biocide for immediate control of mole crickets.

Past and present research on natural enemies
of Florida's mole crickets
During the 1930s mole crickets became a serious economic problem
in central Florida, and research on their biology and control was
undertaken shortly before the outbreak of World War II. At this time








some attention was given to natural enemies. Most were incidental
predators that had no demonstrable effect on mole cricket popula-
tions. However, two kinds of pathogenic fungi were found, one of
which (Syngliocladium urella) was thought to materially reduce
populations of the tawny mole crickets.
There appeared to be no effective native natural enemies in
Florida; however, L. bicolor had recently been introduced into Puerto
Rico, and an attempt was made to introduce that wasp into Florida.
Work was started in 1941 and disrupted by the war. Following the
war, an effort was made to obtain stock of L. bicolor from Belem,
Brazil, and in 1947 one shipment of 138 adults and 40 parasitized
mole crickets was received. All adults and all but one of the parasi-
tized crickets were dead on arrival.
At this same time, as a result of the discovery of DDT, entomolo-
gists were shifting their attention from biological to chemical control.
Other highly effective insecticides, including chlordane, were soon
discovered. In 1949, the annual report of the Florida Agricultural
Experiment Station noted that research on biological control had
been terminated because tests had shown DDT and chlordane to be
highly effective against mole crickets and these were recommended
for control.
Research on biological control was not resumed until 1978, when
the present mole cricket project was initiated. Since both the tawny
and the southern mole crickets were thought to be of South American
origin, the best approach to biological control would have been work
in South America to discover areas of origin and associated natural
enemies of the species that invaded Florida and other areas of south-
eastern United States. When found, enemy species could then be
studied and evaluated as candidates for introduction before being
sent to Florida for further study and field colonization. With limited
resources this was not initially possible; however, L. bicolor could be
obtained from Puerto Rico where it had been introduced in 1938.
Although no new information regarding the wasp had been published
since 1942, correspondence with Puerto Rican entomologists indi-
cated that it was commonly seen at several locations on the island.
Successful introduction ofL. bicolor into Florida from Puerto Rico
appeared unlikely because the colonizing stock used to establish the
species in Puerto Rico came from Belem, Brazil, a city located almost
on the equator. In the history of biological control no beneficial
species native to the equatorial zone has been successfully introduced
into regions of temperate climate. But, since it was not yet possible to
undertake work in South America, and gaining experience with
Larra was judged important, study of L. bicolor was begun.








Beginning in 1978 annual trips were made to Puerto Rico to learn
where to find and how to collect and ship adult Larra, as well as to
obtain information about behavior of the wasps and about their host
mole crickets. Plantings ofS. verticillata were established at Gaines-
ville, Fort Lauderdale, and elsewhere in order to create conditions
favorable for field colonization of Larra.
Experience gained during 1978-1980 made possible the successful
establishment of L. bicolor on a golf course in Fort Lauderdale in
1981. Introductions at Gainsville, Tampa, and Bradenton failed. The
colony at Fort Lauderdale increased during the summer and fall of
1981, and weekly counts of adults on 12 small plots of flowering
Spermacoce indicated that the population doubled in 1982. The in-
creasing trend continued during the spring and summer of 1983.
Unfortunately, in late summer, except for a narrow strip bearing the
Spermacoce plantings, the golf course was treated with ethylene
dibromide (EDB) and the Larra population dropped abruptly to a
very low level. By winter it had partially recovered.
Apart from continuing observation of the L. bicolor population at
the Fort Lauderdale site, domestic research has concerned the biol-
ogy and behavior of the wasp. In June 1983,83 female wasps collected
in Puerto Rico were released at prepared sites on Ram's Horn Ranch
in Hillsborough County. A second release, consisting of 47 females,
was made in December. Ram's Horn Ranch has been selected as the
principal field location for future release of promising enemy species
because it has a large acreage ofbahiagrass pasture heavily infested
with mole crickets and is centrally located with respect to the mole
cricket problem. Also, a cooperative management together with the
high cost of chemical control insures against disruptive actions of the
kind that jeopardized continuance of L. bicolor at Fort Lauderdale.
For reasons already indicated, little work was done in South Amer-
ica during the period 1978-1982. However, information derived from
taxonomic study of museum specimens pointed to the Rio de la Plata
area of South America as the source of the mole crickets responsible
for Florida's pest problem. This was supported by survey of Uruguay
and Buenos Aires Province, Argentina, in late October and early
November 1980. Additional evidence was obtained indicating that
both tawny and southern mole crickets occurred in Uruguay and
northern Argentina.
The foreign research effort was increased during 1981 when an
agreement was negotiated with Professor A. Silveira-Guido of Mon-
tevideo, Uruguay, who then began studies on mole cricket biology
and on natural enemies. During 1982, through operation of sound
traps simulating the calling songs of Florida tawny and southern








mole crickets, he obtained further proof of the conspecificity of the
populations in Florida and Uruguay. He also collected two species of
Larra wasps, subsequently identified as burmeisteri and gastrica.
With funds from USDA, it was possible in 1982 to augment the
foreign research effort through employment of a postdoctoral re-
-search entomologist with extensive South American experience.
From mid-October to the end of December 1982, he visited Brazil,
Uruguay, Argentina, and Paraguay to select a location from which to
conduct a two-year study of natural enemies associated with mole
crickets in South America. After taking into account such factors as
access to field populations of mole crickets, availability of support
facilities, and proximity to an international airport, he concluded
that the Instituto de Biociencias, Universidad Estadual de Sao Paulo,
at Rio Claro, S.P., Brazil was best suited as a location from which to
conduct his research. In July 1983, he moved to Rio Claro and started
his research program.

Future research on natural enemies
During the period 1978-83 much was learned regarding the biol-
ogy and behavior of mole crickets. The identity, distribution, and
area of origin was established for the three species of Scapteriscus
now found in Florida. Apart from information acquired from litera-
ture, much less progress was made in learning of natural enemies of
mole crickets and in determining which species are suitable candi-
dates for introduction into Florida. As a result of USDA funds
awarded to the project in 1983, it should now be possible to obtain this
information as well as to collect or rear stock of promising enemy
species. This is the primary objective of the research centered at Rio
Claro, Brazil.
While efforts will be made to find and assess the potential value of
all organisms that attack species of Scapteriscus in southern Brazil,
Uruguay, and adjacent areas, present knowledge indicates that the
most useful candidates for introduction are likely to be found among
those that are parasitic or pathogenic. In the case of the parasitic
wasps of the genus Larra, there is the possibility of a high degree of
host specificity, with each species specialized to attack a single spe-
cies of Scaperiscus. It may be that more than one species will have to
be imported and colonized. If found and successfully introduced, these
wasps should bring about a very substantial reduction in mole cricket
populations. However, this cannot be accomplished quickly. With
currently available resources and expertise, additional enemy spe-
cies should be found, evaluated, and introduced by 1987. Additional








time will be needed to insure that they are distributed throughout
Florida and to allow their effects to be evaluated.
Future studies of L. bicolor will involve efforts to colonize it at
other locations in south and central Florida. Effort will also be made
to obtain information on host range, dispersal, and other factors that
influence its population numbers and effectiveness as an enemy of
mole crickets in Florida. These same studies will be undertaken on
other species of Larra as well as on other kinds of mole cricket
enemies as these are found in South America and shipped to Florida.
For immediate biological control, particularly on golf courses and
other high-value land areas, nematodes of the genus Neoaplectana
and pathogens show the greatest promise. Here, more time is likely
to be required in the research phase, but if an effective agent is found,
it can be quickly exploited.


Diseases (Drion Boucias)

Pathogens and symptoms
The entomopathogenic fungi are the only reported microorganisms
that have been determined to be causal agents of disease in mole
cricket populations. The green muscardine, Metarhizium anisopliae,
is the predominant fungal pathogen associated with mole crickets.
This fungus is characterized by the production of compact colums of
green pigmented spores conidiaa) on the external surface of diseased
insects. The white muscardine, Beauveria bassiana, a fungus which
produces singly-borne white conidia on infected hosts, has been de-
tected on several mole cricket cadavers. Two other entomopathogenic
fungi, an saria species that produces unbranched, thick, white stalk-
like structures and a Sorosporella species characterized by its brick
red mycelium-spore complex, have been identified from nymphal and
adult tawny mole crickets, respectively.
To date no protozoan, viral, or bacterial agent has been reported to
be a mole cricket pathogen. The reasons for this apparent lack of
detection of these causal agents may be the lack of an extensive
survey of various geographical populations of mole crickets for the
presence of disease, and the fact that mole crickets are soil-inhabiting
insects, making it extremely difficult to collect diseased larvae prior
to purification and/or contamination of the cadavers by soil sap-
rophytes. Fungal infections, resulting in the production of a mum-
mified cadaver, are somewhat protected from degradation by the soil
microflora.
The relative susceptibility of mole crickets to viral, bacterial, and








time will be needed to insure that they are distributed throughout
Florida and to allow their effects to be evaluated.
Future studies of L. bicolor will involve efforts to colonize it at
other locations in south and central Florida. Effort will also be made
to obtain information on host range, dispersal, and other factors that
influence its population numbers and effectiveness as an enemy of
mole crickets in Florida. These same studies will be undertaken on
other species of Larra as well as on other kinds of mole cricket
enemies as these are found in South America and shipped to Florida.
For immediate biological control, particularly on golf courses and
other high-value land areas, nematodes of the genus Neoaplectana
and pathogens show the greatest promise. Here, more time is likely
to be required in the research phase, but if an effective agent is found,
it can be quickly exploited.


Diseases (Drion Boucias)

Pathogens and symptoms
The entomopathogenic fungi are the only reported microorganisms
that have been determined to be causal agents of disease in mole
cricket populations. The green muscardine, Metarhizium anisopliae,
is the predominant fungal pathogen associated with mole crickets.
This fungus is characterized by the production of compact colums of
green pigmented spores conidiaa) on the external surface of diseased
insects. The white muscardine, Beauveria bassiana, a fungus which
produces singly-borne white conidia on infected hosts, has been de-
tected on several mole cricket cadavers. Two other entomopathogenic
fungi, an saria species that produces unbranched, thick, white stalk-
like structures and a Sorosporella species characterized by its brick
red mycelium-spore complex, have been identified from nymphal and
adult tawny mole crickets, respectively.
To date no protozoan, viral, or bacterial agent has been reported to
be a mole cricket pathogen. The reasons for this apparent lack of
detection of these causal agents may be the lack of an extensive
survey of various geographical populations of mole crickets for the
presence of disease, and the fact that mole crickets are soil-inhabiting
insects, making it extremely difficult to collect diseased larvae prior
to purification and/or contamination of the cadavers by soil sap-
rophytes. Fungal infections, resulting in the production of a mum-
mified cadaver, are somewhat protected from degradation by the soil
microflora.
The relative susceptibility of mole crickets to viral, bacterial, and








protozoan agents isolated from other orthopteran insects has not yet
been fully assessed. However, preliminary bioassays with three
microsporidian isolates, Nosema locusta, N. acridophagus, and N.
cuneatum, incorporated in food substrate (106 spores/gram) demons-
trated that neither southern nor tawny mole crickets supported
pathogen development.

Activity of Metarhizium anisopliae
against mole crickets
The majority of recent research on mole cricket pathogens has
involved laboratory studies on Metarhizium anisopliae. Conidia of
this fungus will germinate on the host insect, penetrate through the
cuticle via germ tube formation, and multiply in the hemocoel and
other tissues, causing eventual mummification of the host. Under
proper environmental conditions (that is, high humidity), filaments
(conidiophores) will emerge through the cuticle, producing large
numbers of progeny conidia (Fig. 17) arranged in a typical palisade
pattern (Fig. 18). These conidia, being produced in aggregate bundles
and having hygroscopic properties, are well adapted for survival in
the soil ecosystem. In fact, many M. anisopliae strains have been
isolated from soil inhabiting insects.
The relative activity ofM. anisopliae against mole crickets appears
to depend upon both the specific strain of M. anisopliae assayed and
on the particular mole cricket species tested. Various M. anisopliae
isolates (Table 2) were bioassayed in the laboratory against first
instar nymphs of southern and tawny mole crickets (Table 3). The
MATR-20, MADA-24, and MAPG-77 strains originally isolated from
scarab beetle hosts (Table 2) were highly virulent to both species and
produced large numbers of progeny conidia on 100% of the mole
cricket cadavers. Other M. anisopliae isolates appeared either to be
less virulent or were not capable of producing progeny conidia on
cadavers. The MC-3057 and MC-3059 strains, originally isolated
from tawny mole cricket nymphs, although capable of killing mole
crickets under laboratory conditions, did not produce conidia on the
resulting cadavers. The production of progeny conidia is considered
to be of paramount importance for the establishment ofM. anisopliae
as a long-term microbial agent.

Future research
To date an extensive survey of entomopathogens of mole crickets
has not been undertaken. One would expect that this insect, like
other arthropods, is susceptible to a wide spectrum of disease agents.








protozoan agents isolated from other orthopteran insects has not yet
been fully assessed. However, preliminary bioassays with three
microsporidian isolates, Nosema locusta, N. acridophagus, and N.
cuneatum, incorporated in food substrate (106 spores/gram) demons-
trated that neither southern nor tawny mole crickets supported
pathogen development.

Activity of Metarhizium anisopliae
against mole crickets
The majority of recent research on mole cricket pathogens has
involved laboratory studies on Metarhizium anisopliae. Conidia of
this fungus will germinate on the host insect, penetrate through the
cuticle via germ tube formation, and multiply in the hemocoel and
other tissues, causing eventual mummification of the host. Under
proper environmental conditions (that is, high humidity), filaments
(conidiophores) will emerge through the cuticle, producing large
numbers of progeny conidia (Fig. 17) arranged in a typical palisade
pattern (Fig. 18). These conidia, being produced in aggregate bundles
and having hygroscopic properties, are well adapted for survival in
the soil ecosystem. In fact, many M. anisopliae strains have been
isolated from soil inhabiting insects.
The relative activity ofM. anisopliae against mole crickets appears
to depend upon both the specific strain of M. anisopliae assayed and
on the particular mole cricket species tested. Various M. anisopliae
isolates (Table 2) were bioassayed in the laboratory against first
instar nymphs of southern and tawny mole crickets (Table 3). The
MATR-20, MADA-24, and MAPG-77 strains originally isolated from
scarab beetle hosts (Table 2) were highly virulent to both species and
produced large numbers of progeny conidia on 100% of the mole
cricket cadavers. Other M. anisopliae isolates appeared either to be
less virulent or were not capable of producing progeny conidia on
cadavers. The MC-3057 and MC-3059 strains, originally isolated
from tawny mole cricket nymphs, although capable of killing mole
crickets under laboratory conditions, did not produce conidia on the
resulting cadavers. The production of progeny conidia is considered
to be of paramount importance for the establishment ofM. anisopliae
as a long-term microbial agent.

Future research
To date an extensive survey of entomopathogens of mole crickets
has not been undertaken. One would expect that this insect, like
other arthropods, is susceptible to a wide spectrum of disease agents.































Fig. 17. Scanning electron micrograph of first nymphal instar of cadaver of tawny
mole cricket infected with M. anisopliae (MADA-31).


Fig. 18. Scanning electron micrograph of an individual M. anisopliae "palisade"
produced on cadaver of tawny mole cricket. Note tremendous numbers of conidia within
each palisade.








Table 2. Host distribution of the Metarhizium anisopliae isolates.
Isolate Insect host Location
MATRAW 20 Fuller rose weevil Bradley Jet., Florida
MADA 24 citrus root weevil Apopka, Florida
MADA 31 citrus root weevil Puerto Rico
MAPG 77 Fuller rose weevil Duetle, Florida
MARB 297 "Rhinoceros beetle" Apia, West Samoa
MARB298 "Rhinoceros beetle" Apia, West Samoa
MABP 456 "Brown planthopper" Irri, Philippines
MADA 472 Dasygnathus scarab beetle Australia
MC-3057 tawny mole cricket Uruguay (1982)
MC-3059 tawny mole cricket Uruguay (1982)



For example, M. anisopliae is a pathogen that is isolated frequently
from mole crickets. This fungus, infectious to a wide spectrum of
soil-inhabiting insect pests and amenable to mass production on
artificial substrates, is currently being studied for its potential as a
microbial control agent. The protocols currently being developed for
the production, formulation, and application ofM. anisopliae against
other soil inhabiting and pasture land insect pests are directly appli-
cable to managing mole cricket populations with M. anisopliae.
IFAS research has demonstrated that certain strains of this fungus
are highly virulent to and are capable of sporulating on southern and
tawny mole cricket nymphs. Ideally, one would like to be able to
inoculate M. anisopliae into mole cricket populations and establish it
as long term biological control agent. Current research is underway
to further define the relative activity of M. anisopliae isolates against
mole crickets. Future research will involve small plot evaluations of
the "high" virulent isolates against natural populations of mole
crickets. This research, aimed at quantifying the virulence, persist-
ence, and spread of this disease under field conditions, is vitally
important for developing M. anisopliae as an effective microbial
control against mole cricket populations.

Resistant Varieties
(James A. Reinert and Philip Busey)

The greatest damage by mole crickets in Florida is to turf and
pasture grasses. Most significant is destruction of bahiagrass and
bermudagrass by the tawny mole cricket and of bermudagrass and
St. Augustinegrass by short-winged mole crickets. Resistant culti-
vars are needed in these instances as a safe, economical control











Table 3. Response of first instar nymphs of tawny and southern mole crickets
contaminated with Metarhizium anisopliae conidia.
Tawny mole cricket Southern mole cricket
Percent Mean time to M. anisopliae Percent Mean time to M. anisopliae
Fungal strain* mortality death (days) sporulation** mortality death (days) sporulation
MATRW-20 100 10 100 100 6 100
MADA-24 100 4 100 100 5 100
MADA-31 0 0 0 100 5 100
MAPG-77 100 6 100 100 6 100
MARB-297 0 0 0 29 6 100
MARB-298 100 6 100 24 14 0
MARB-456 100 9 100 42 5 100
MADS-472 100 10 60 53 14 20
MC-3057 100 4 0 -
MC-3059 40 13 0 -

*Spores applied to soil at a rate of 0.5 mg/g soil.
**Percent dead larvae producing spores.









strategy. The potential role of turf and pasture as alternate hosts for
mole crickets invading vegetables and other corps must also be con-
sidered. We have, therefore, evaluated turfgrass cultivars and spe-
cies for resistance to pest mole crickets. We have included an evalua-
tion of promising germplasm and prospective cultivar releases for
mole cricket resistance, in some cases supporting cultivar releases
with timely information on genetic vulnerability to mole crickets.

Bermudagrass selections

Most of the bermudagrass cultivars grown as turfgrass were evalu-
ated for resistance to adult tawny mole crickets (Table 4) in two
experiments in screened field cages (Fig. 19). This mole cricket dam-
ages bermudagrass by feeding on the roots and leaves, and by loosen-
ing and uprooting the plants as it tunnels. Feeding damage in these
experiments was characterized by extensive severing of upright leafy
shoots from the spaced, container-grown transplants. Damage esti-
mates were based upon visual ratings and relative reduction in
harvested plants.














i -.'






Fig. 19. Plugs of bermudagrass cultivars and selections in test for resist-
ance to mole crickets (Table 4). Plugs were planted in replicated, paired,
screen cages (10 x 10 x 7 ft, L x W x H). Adult mole crickets were introduced
into one of each cage pair and allowed to feed. Damage estimates were made
by comparing shoot harvests of plants grown with mole crickets to those
grown without. (Note high preference for 'Texturf 10,' lower center, com-
pared to several of the other selections.)









Table 4. Tawny mole cricket damage and growth reduction to
bermudagrasses in field cage experiments.
Dry wt. Visual
Bermudagrass reduction of damage Resistance
genotype clippings* rating** score


PI-290659
FL-2400
PI-291586
Ormond
Santa Ana
T-72-54
Tifway
Tifway II
Tifgreen II
FB-109
FB-119
U-3
Tifdwarf
Tifgreen
Sunturf


Experiment #1
-3 a
6 ab
7 ab
22 ab
25 ab
24 ab
22 ab
19 ab
37 ab
29 ab
31 ab
38 ab
36 ab
38 ab
62 b
Experiment #2


6a
6a
13 ab
13 ab
12 ab
14 ab
16 ab
30 ab
14 ab
25 ab
25 ab
37 abc
52 bc
52 bc
74 c


Tifgreen 3 a 21 ab 88
Ormond 23 ab 8 a 85
Sunturf 18 a 24 abc 79
Midiron 28 abc 14 a 79
Hardy 29 abc 21 ab 75
FB-119 45 abcd 13 a 71
Tiffine 38 abc 28 abc 67
Tufcote 44 abcd 32 abc 62
Northrup King 78098 43 abcd 33 abc 62
Tifway 44 abed 30 abc 63
Pee Dee 54 abcd 32 abc 57
Everglades 71 bcd 29 abc 50
Tuffy 71 bcd 30 abc 50
Texturf-1F 79 cd 53 bc 34
Texturf-10 92 d 55 c 27
SOURCE: Taken in part from Reinert and Busey (in preparation).
NOTE: Data reflect average of four replications. Letters after figures in middle col-
umns show statistically significant differences. Only those figures in the same column
and experiment that have no letter in common are significantly different.
*All foliage was removed at or just above ground level (check and mole cricket-
inoculated plants treated alike). Reduction due to mole crickets was figured on a
percentage of control.
**Visually estimated percentage of dead or dying foliage.
tResistance scre = 100 clipping dry weight + visual rating
2








All bermudagrass selections tested were damaged, but 'Ormond',
FL2400, PI-290659, and PI-291586 showed the least damage in these
studies. Conflicting results were produced with 'Tifgreen' and
'Sunturf' in the two experiments, whereas 'Tifway' and FB-119, also
common to both studies, each received similar and intermediate
damage. PI-291586 is currently under consideration for cultivar re-
lease for use as low maintenance turf.
Several of the same bermudagrass cultivars were tested for resist-
ance to the short-winged mole cricket. This species also fed exten-
sively on the plants. FB-119, Ormond, and 'Common' were least
damaged by this species while Tifgreen and Tifway were both se-
verely injured. Tifgreen and Tifway are the principal bermuda-
grasses grown on Florida golf courses, and both are severely damaged
by mole crickets under field conditions.

Bahiagrass selections
Bahiagrass selections were evaluated for resistance to both the
southern and tawny mole cricket. Twenty lines ofbahiagrass, includ-
ing 'Argentine' and 'Paraguayan', were tested against high popula-
tions of southern mole crickets in tank cages. This species does not
feed extensively on grass plants but does most of its damage by
uprooting the grass or loosening the soil around the roots, causing
them to desiccate. Five experimental lines (FL-1950, FL-1962,
FL-1990, FBA-20, and FBA-21) and Argentine sustained little or no
damage in the study. All of these except FBA-21 produced more shoot
growth when southern mole crickets were present than when they
were excluded. Paraguayan and FBA-14 were the most damaged in
the study, and the latter also showed severe damage in a field plot.
Eleven diploid, Pensacola-type bahiagrass selections were exposed
to field levels of tawny mole crickets in screened field cages (Fig. 19)
and sustained 59% to 88% damage within 11 weeks after inoculation
with 4.8 mole crickets per square meter.

Comparison of species and
cultivars of turfgrass
Field observations have suggested that mole crickets prefer certain
species of warm season turfgrasses. When selections of St. Augus-
tinegrass, bahiagrass, bermudagrass, centipedegrass, and zoysia-
grass were exposed to populations of southern mole crickets, greatest
damage was sustained by 'Bitterblue' St. Augustinegrass, Tifway
bermudagrass, and 'Emerald' zoysiagrass, in descending order.








All bermudagrass selections tested were damaged, but 'Ormond',
FL2400, PI-290659, and PI-291586 showed the least damage in these
studies. Conflicting results were produced with 'Tifgreen' and
'Sunturf' in the two experiments, whereas 'Tifway' and FB-119, also
common to both studies, each received similar and intermediate
damage. PI-291586 is currently under consideration for cultivar re-
lease for use as low maintenance turf.
Several of the same bermudagrass cultivars were tested for resist-
ance to the short-winged mole cricket. This species also fed exten-
sively on the plants. FB-119, Ormond, and 'Common' were least
damaged by this species while Tifgreen and Tifway were both se-
verely injured. Tifgreen and Tifway are the principal bermuda-
grasses grown on Florida golf courses, and both are severely damaged
by mole crickets under field conditions.

Bahiagrass selections
Bahiagrass selections were evaluated for resistance to both the
southern and tawny mole cricket. Twenty lines ofbahiagrass, includ-
ing 'Argentine' and 'Paraguayan', were tested against high popula-
tions of southern mole crickets in tank cages. This species does not
feed extensively on grass plants but does most of its damage by
uprooting the grass or loosening the soil around the roots, causing
them to desiccate. Five experimental lines (FL-1950, FL-1962,
FL-1990, FBA-20, and FBA-21) and Argentine sustained little or no
damage in the study. All of these except FBA-21 produced more shoot
growth when southern mole crickets were present than when they
were excluded. Paraguayan and FBA-14 were the most damaged in
the study, and the latter also showed severe damage in a field plot.
Eleven diploid, Pensacola-type bahiagrass selections were exposed
to field levels of tawny mole crickets in screened field cages (Fig. 19)
and sustained 59% to 88% damage within 11 weeks after inoculation
with 4.8 mole crickets per square meter.

Comparison of species and
cultivars of turfgrass
Field observations have suggested that mole crickets prefer certain
species of warm season turfgrasses. When selections of St. Augus-
tinegrass, bahiagrass, bermudagrass, centipedegrass, and zoysia-
grass were exposed to populations of southern mole crickets, greatest
damage was sustained by 'Bitterblue' St. Augustinegrass, Tifway
bermudagrass, and 'Emerald' zoysiagrass, in descending order.








Other selections of St. Augustinegrass, bahiagrass, and bermuda-
grass were either not damaged or moderately damaged. A selection of
centipedegrass was also moderately damaged. Among the selections
of St. Augustinegrass, bahiagrass, or bermudagrass, those with the
finest texture were most damaged, while the coarser selections were
stimulated to produce more shoot growth in the presence of southern
mole crickets.

Pasture grass selections
Pensacola and Argentine bahiagrass were also compared to four
cultivars of Hemarthria grass in forced-feeding studies. Cultivars
were 80 to 90% damaged by tawny mole crickets; 'Floralta' Hemar-
thria was the least damaged. However, southern mole crickets caused
only 8 to 38% damage to the same grasses under the same conditions.
Argentine bahiagrass showed little damage from southern mole
crickets, possibly due to the mole cricket's greater preference for
Hemarthria. Also, earlier tests with southern mole crickets, have
shown that Argentine may be the least preferred of the bahiagrass
types. In previous studies, 'Tifton 44' bermudagrass also exhibited
tolerance to the southern mole cricket. Follow up studies have shown
that in situations where Floralta is the only pasture grass available,
mole cricket damage is as severe as might be expected in bahiagrass.
Field testing of Tifton 44 has yet to be concluded. The possible
resistance in Argentine bahiagrass also needs to be closely examined.

Host resistance potential for grasses
Field observations suggest that mole crickets prefer bahiagrass
and bermudagrass. Mole crickets also show a preference for the finer
textured selections within each grass species. Levels of resistance or
tolerance to mole cricket feeding and tunneling damage have been
shown within species of bermudagrass, bahiagrass, and St. Augus-
tinegrass.
The relative contribution of nonpreference and host plant toler-
ance needs to be more carefully defined. Obviously, in a large mono-
culture such as a golf course or a pasture, nonpreference in the
absence of other resistance mechanisms would be of limited economic
value. Because the plants were widely spaced in these studies, inher-
ent differences in density of growth among species were largely
masked. However, potential for resistance has been documented, and
further studies promise to identify cultivars that can be planted with
substantially reduced risk of mole cricket damage.








Other selections of St. Augustinegrass, bahiagrass, and bermuda-
grass were either not damaged or moderately damaged. A selection of
centipedegrass was also moderately damaged. Among the selections
of St. Augustinegrass, bahiagrass, or bermudagrass, those with the
finest texture were most damaged, while the coarser selections were
stimulated to produce more shoot growth in the presence of southern
mole crickets.

Pasture grass selections
Pensacola and Argentine bahiagrass were also compared to four
cultivars of Hemarthria grass in forced-feeding studies. Cultivars
were 80 to 90% damaged by tawny mole crickets; 'Floralta' Hemar-
thria was the least damaged. However, southern mole crickets caused
only 8 to 38% damage to the same grasses under the same conditions.
Argentine bahiagrass showed little damage from southern mole
crickets, possibly due to the mole cricket's greater preference for
Hemarthria. Also, earlier tests with southern mole crickets, have
shown that Argentine may be the least preferred of the bahiagrass
types. In previous studies, 'Tifton 44' bermudagrass also exhibited
tolerance to the southern mole cricket. Follow up studies have shown
that in situations where Floralta is the only pasture grass available,
mole cricket damage is as severe as might be expected in bahiagrass.
Field testing of Tifton 44 has yet to be concluded. The possible
resistance in Argentine bahiagrass also needs to be closely examined.

Host resistance potential for grasses
Field observations suggest that mole crickets prefer bahiagrass
and bermudagrass. Mole crickets also show a preference for the finer
textured selections within each grass species. Levels of resistance or
tolerance to mole cricket feeding and tunneling damage have been
shown within species of bermudagrass, bahiagrass, and St. Augus-
tinegrass.
The relative contribution of nonpreference and host plant toler-
ance needs to be more carefully defined. Obviously, in a large mono-
culture such as a golf course or a pasture, nonpreference in the
absence of other resistance mechanisms would be of limited economic
value. Because the plants were widely spaced in these studies, inher-
ent differences in density of growth among species were largely
masked. However, potential for resistance has been documented, and
further studies promise to identify cultivars that can be planted with
substantially reduced risk of mole cricket damage.










IV. CHEMICAL CONTROL OF
MOLE CRICKETS (R. L. Kepner)

Beginnings

The introduction, in about 1900, and subsequent spread of pest
mole crickets quickly aroused the attention of farmers along much of
coastal Georgia. By 1909 mole cricket damage had become so severe
that immediate remedial action was warranted. In 1910 the state of
Georgia initiated a three year study to determine an economically
effective means for controlling this serious threat. Various control
techniques such as metal barriers, light traps, and repellants were
attempted but had limited success. It was concluded that poison baits
concocted of various mashes of bran, corn, or cottonseed meal, mixed
with paris green or calcium arsenate, offered the best control.
Poison baits, broadcast over the soil surface or buried in trenches
around individual plants, provided the only economically effective
control of mole crickets for several decades. As mole crickets con-
tinued their spread through the southeastern United States, recom-
mended bait formulations were changed in response to claims of
improved control by incorporating low grade flour or egg mash. In
addition to poison baits, soil treatments with lead arsenate and soil
fumigation with carbon bisulfide or calcium cyanide were recom-
mended for control by 1930.
Mole crickets had spread well into Florida by the 1930s and popula-
tions soon reached epidemic levels. In 1940, central Florida's vege-
table growing areas experienced the worst infestation of mole crick-
ets ever reported in the United States. Hundreds of growers made
appeals for assistance, and in response the USDA Bureau of Entomol-
ogy and Plant Quarantine set up an emergency mole cricket control
program. Growers were supplied with 120 pounds of 71/2% calcium
arsenate, bran bait per acre to be applied in three applications of 40
pounds each throughout the growing season. Studies were initiated
to develop a more economical bait but were soon suspended because of
World War II.
Until the mid-1940s, poison baits were the most efficient means of
suppressing mole cricket populations, but with the advent of syn-
thetic insecticides in 1944, a new era of control began. Persistent,
highly toxic contact poisons such as DDT and chlordane were found to
be very effective against mole crickets. Baits were no longer consid-
ered the best method of control, and efforts to improve bait formula-










IV. CHEMICAL CONTROL OF
MOLE CRICKETS (R. L. Kepner)

Beginnings

The introduction, in about 1900, and subsequent spread of pest
mole crickets quickly aroused the attention of farmers along much of
coastal Georgia. By 1909 mole cricket damage had become so severe
that immediate remedial action was warranted. In 1910 the state of
Georgia initiated a three year study to determine an economically
effective means for controlling this serious threat. Various control
techniques such as metal barriers, light traps, and repellants were
attempted but had limited success. It was concluded that poison baits
concocted of various mashes of bran, corn, or cottonseed meal, mixed
with paris green or calcium arsenate, offered the best control.
Poison baits, broadcast over the soil surface or buried in trenches
around individual plants, provided the only economically effective
control of mole crickets for several decades. As mole crickets con-
tinued their spread through the southeastern United States, recom-
mended bait formulations were changed in response to claims of
improved control by incorporating low grade flour or egg mash. In
addition to poison baits, soil treatments with lead arsenate and soil
fumigation with carbon bisulfide or calcium cyanide were recom-
mended for control by 1930.
Mole crickets had spread well into Florida by the 1930s and popula-
tions soon reached epidemic levels. In 1940, central Florida's vege-
table growing areas experienced the worst infestation of mole crick-
ets ever reported in the United States. Hundreds of growers made
appeals for assistance, and in response the USDA Bureau of Entomol-
ogy and Plant Quarantine set up an emergency mole cricket control
program. Growers were supplied with 120 pounds of 71/2% calcium
arsenate, bran bait per acre to be applied in three applications of 40
pounds each throughout the growing season. Studies were initiated
to develop a more economical bait but were soon suspended because of
World War II.
Until the mid-1940s, poison baits were the most efficient means of
suppressing mole cricket populations, but with the advent of syn-
thetic insecticides in 1944, a new era of control began. Persistent,
highly toxic contact poisons such as DDT and chlordane were found to
be very effective against mole crickets. Baits were no longer consid-
ered the best method of control, and efforts to improve bait formula-








tions diminished. Though not as effective as contact poisons, baits
were still used, since they were economical and growers were accus-
tomed to them. The only change was that chlordane became the
toxicant.
Chlorinated hydrocarbon insecticides, especially chlordane in the
form of baits, sprays, and dusts, became the standard control agents
for mole crickets because they offered economical, long-term control.
In the early 1970s, problems with residues on food and forage crops
caused most of these insecticides to be removed from use in many
areas where mole crickets were a problem. In addition to restrictions
on its use, chlordane had become less toxic to mole crickets by the
mid-1970s, probably because the crickets had developed resistance.
With the loss of efficient, long-term control agents, IFAS Extension
personnel began a continuing series of screening tests in search of
new insecticides effective against mole crickets. Some were found,
and these are the basis of present chemical control. However, these
insecticides proved less persistent and more costly, and, as a result,
mole crickets became increasingly difficult and expensive to control.

Present Day Control

Chemical control of mole crickets is presently accomplished by soil
treatments with baits, sprays, and granules.

Baits
In many instances, such as in pastures, chemical control is limited
to the use of baits, since irrigation and/or soil incorporation is not
feasible. Though baits are the most economical of all control mea-
sures, costs are still high (Table 5), and control is not always
adequate.
Controlling mole crickets with poison baits can be very effective if
applications are properly timed. The use of baits takes advantage of
mole cricket feeding behavior. Under favorable conditions, mole
crickets will come to the surface and forage for food. While burrowing
through the soil, they periodically leave their tunnels and search the
soil surface. It is this behavior that makes the use of baits so attrac-
tive. If a poison bait is present when a large percentage of mole
crickets are feeding on the surface, control will be maximized. It is
best to treat with baits during the summer months after eggs have
hatched and weather conditions are most favorable (see Fig. 10). If
baits are applied early in the summer when the nymphs are small,
control can be obtained before significant damage has occurred.








tions diminished. Though not as effective as contact poisons, baits
were still used, since they were economical and growers were accus-
tomed to them. The only change was that chlordane became the
toxicant.
Chlorinated hydrocarbon insecticides, especially chlordane in the
form of baits, sprays, and dusts, became the standard control agents
for mole crickets because they offered economical, long-term control.
In the early 1970s, problems with residues on food and forage crops
caused most of these insecticides to be removed from use in many
areas where mole crickets were a problem. In addition to restrictions
on its use, chlordane had become less toxic to mole crickets by the
mid-1970s, probably because the crickets had developed resistance.
With the loss of efficient, long-term control agents, IFAS Extension
personnel began a continuing series of screening tests in search of
new insecticides effective against mole crickets. Some were found,
and these are the basis of present chemical control. However, these
insecticides proved less persistent and more costly, and, as a result,
mole crickets became increasingly difficult and expensive to control.

Present Day Control

Chemical control of mole crickets is presently accomplished by soil
treatments with baits, sprays, and granules.

Baits
In many instances, such as in pastures, chemical control is limited
to the use of baits, since irrigation and/or soil incorporation is not
feasible. Though baits are the most economical of all control mea-
sures, costs are still high (Table 5), and control is not always
adequate.
Controlling mole crickets with poison baits can be very effective if
applications are properly timed. The use of baits takes advantage of
mole cricket feeding behavior. Under favorable conditions, mole
crickets will come to the surface and forage for food. While burrowing
through the soil, they periodically leave their tunnels and search the
soil surface. It is this behavior that makes the use of baits so attrac-
tive. If a poison bait is present when a large percentage of mole
crickets are feeding on the surface, control will be maximized. It is
best to treat with baits during the summer months after eggs have
hatched and weather conditions are most favorable (see Fig. 10). If
baits are applied early in the summer when the nymphs are small,
control can be obtained before significant damage has occurred.












Table 5. Insecticide formulations labeled in Florida for control of mole crickets on various crops.
Rate Approximate cost of
Treatment Formulation (lb A.I./acre) Registered use materials per acre
BAITS: Malathion 2% 1-2 turf, pasture $24-48
Trichlorfon 5% 1-1.5 pasture, $15-22


SPRAYS:


GRANULES:


Carbaryl 20%
SPropoxur 2%
Chlorpyrifos 0.5%
Diazinon 2 EC
Diazinon 4 EC
Propoxur 70WP
Ethoprop 10G
Isofenphos 5G
Isofenphos 1.5G
Diazinon 14G


field crops
turf, pasture
turf
turf
turf
vegetables
turf
turf
turf
turf
vegetables,
field crops


$25-50
$130-260
$55-82
$32
$6
$60-120
$88
$64
$135
$90


IMPORTANT NOTE: These insecticides are currently (1983) labeled for use in Florida, but registrations are subject to change at any time. Consult
your county extension agent for up-to-date control recommendations.








Treatments during the spring months, when mostly adults are pres-
ent, are not recommended, because adults do not accept baits as
readily and the chances of reinfestation from subsequent flights and
unhatched eggs are high.
Baits should be applied in the late afternoon or early evening,
preferably soon after a rain. Mole crickets are most active on the
surface at such times, when the soil is moist and temperatures are
warm. If the bait is applied when the soil is dry or temperatures are
cool, few crickets will be active, and little of the bait will be con-
sumed. Sunlight and high daytime temperatures will then quickly
degrade the bait and control efforts will be wasted. Also, rain soon
after a bait application will leach out the insecticide and render the
bait ineffective.
The proper timing of a bait application is a crucial step toward
improving chances of control, but it does not insure success. Control
ultimately depends on the proportion of mole crickets feeding and the
amount of bait consumed. In an attempt to increase the effectiveness
of chemical control and to make baits more economical, intensive
investigations into bait formulation were undertaken.
These investigations have dealt with each of the four basic compo-
nents of baits.
1) Attractants To improve the chances of mole crickets finding a
bait, it should be made attractive from a distance. Such a bait could
draw the cricket to the poison instead of relying on the cricket
accidentally stumbling upon it as it randomly forages for food. Not
only would attractive baits make control more efficient, but less bait
would be required.
More than 45 materials were tested as possible food attractants for
both tawny and southern mole crickets. Tawny mole crickets were
not attracted to any of the materials tested, but southern mole crick-
ets showed attraction to rancid hamburger and fish meal. These
materials are not suitable bait additives, since they are highly un-
stable and costly, and are only attractive to the pest mole cricket of
least economic importance.
2) Feeding stimulants Mole crickets, in their random search
for food, will sample food before ingesting it. If the material is found
palatable, its chances of being consumed are very high. Therefore
adding a feeding stimulant to a bait should greatly enhance the
probability that it will be accepted and consumed.
Over the years amyl acetate and molasses have been advocated as
additives to enhance bait consumption. However, recent IFAS tests
have shown that amyl acetate does not encourage feeding by either
mole cricket species, and at high concentrations is a deterrent. There-








fore, it should not be used. Molasses proved moderately attractive to
tawny mole crickets but not to southern mole crickets. Addition of
molasses for pasture and turf is justified since tawny mole crickets do
most of the damage in these areas. Several products are excellent
feeding stimulants for both types of mole crickets. These are COAX
(commercial feeding stimulant-$2.00/lb), brewers concentrate
(brewery byproduct, $0.02/lb), malt extract ($0.46/lb), and crude cot-
tonseed or soybean oil in combination with sucrose ($0.25/lb). Brew-
ers concentrate appears to be a good additive for mole cricket baits,
since it is slightly cheaper than molasses ($0.03/lb) and is signifi-
cantly more attractive.
3) Carriers Carriers constitute the bulk of the bait and can
account for greater than 50% of the material costs. Mole cricket baits
have been formulated on a variety of materials, varying considerably
in cost, including vermiculite ($0.45/lb), cottonseed meal ($0.16/lb),
wheat bran ($0.15/lb), laying mash ($0.14/lb), cracked corn ($0.12/lb),
corncob grits ($0.10/lb), and peanut hulls ($0.04/lb). Wheat bran,
cracked corn, and laying mash are highly acceptable to mole crickets.
Addition of feeding stimulants appears to enhance their acceptance
only moderately. This is probably because these materials have some
nutritional value and are therefore naturally palatable. Inert com-
pounds such as peanut hulls, vermiculite, and corncob grits are not
very palatable alone but become highly acceptable when feeding
stimulants are added. Baits formulated with either corncob grits or
sawdust, incorporated with malt extract, have proved as effective as
laying mash-molasses bait in field plot tests. These results suggest
that material costs can be reduced by at least 50% if baits are formu-
lated on a cheap, readily available carrier such as peanut hulls.
4) Toxicants Mole crickets are susceptable to most insecticides.
Of those most likely to be used in baits, chlorpyrifos (Dursban) and
trichlorfon (Dylox, Proxol) are the most toxic and are effective as
0.5% and 1% baits applied at 2 pounds A.I./acre. These insecticides
are expensive at $8 to $17/lb A.I. Malathion and carbaryl (Sevin) are
not as toxic and therefore require more concentrated baits for good
control but are significantly less expensive at $3 and $5/lb A.I.,
respectively. Baits formulated as 20% carbaryl or 2% malathion have
both shown excellent control when applied at 2 lb A.I./acre. There-
fore, using more concentrated baits formulated from less toxic insec-
ticides could reduce toxicant costs as much as 50%.
5) Other components An ideal bait should remain both highly
acceptable and toxic for an extended period of time under field condi-
tions. The addition of waterproofing agents, antioxidants, or UV light








sunscreens to bait formulations may prolong the field life and there-
fore enhance control. No compounds of these types have been tested.


Sprays and granules
Soil treatments with sprays and granules are generally more effec-
tive than baits in controlling mole crickets since they do not rely on
feeding, and timing is not as critical. But they generally require more
insecticide per acre, are more expensive, and require some sort of
irrigation or soil incorporation. Irrigation serves two purposes: 1) it
carries the insecticide into the root zone; and 2) it encourages mole
crickets to be active in the upper layer of soil where they can come
into contact with the poison. Control with any of these treatments is
best during the summer months for reasons previously explained.
Ethoprop (Mocap) and isofenphos (Oftanol) are the best chemical
treatments for mole crickets in turf. Isofenphos offers up to several
months of control with a single application and is most efficient when
applied early in the summer. Propoxur (Bayon) and diazinon
(Sarolex) sprays are not as effective in turf as other registered for-
mulations.


Evaluation of chemical control
In the past, the effectiveness of chemical controls has been evalu-
ated by counting dead or moribund crickets on the surface. This can
be misleading, since many crickets die below the surface. A chemical
causing high mortality below ground could be rated ineffective. Addi-
tionally, the initial population is unknown, and movement into and
out of the treated area is not accounted for. A technique developed to
evaluate true chemical efficacy involves placing a known number of
mole crickets in soil-filled cages or containers buried at soil level
prior to treatment. Control can be accurately determined by counting
the mortality both above and below the surface. This technique,
though labor-intensive, is the most reliable means of comparing
toxicity of chemical treatments.
Flushing live crickets from the soil with mild soap and pyrethrin
solutions can also be used to evaluate chemical control. Because of
the inefficiency of flushing, only a relative measure of control can be
obtained. Other techniques involve measuring the extent of tunnel-
ing/damage or the number of plants destroyed. Like the flush, these
techniques give only a relative measure of control but are useful for
evaluating the long-term effects of chemical treatments.


46








sunscreens to bait formulations may prolong the field life and there-
fore enhance control. No compounds of these types have been tested.


Sprays and granules
Soil treatments with sprays and granules are generally more effec-
tive than baits in controlling mole crickets since they do not rely on
feeding, and timing is not as critical. But they generally require more
insecticide per acre, are more expensive, and require some sort of
irrigation or soil incorporation. Irrigation serves two purposes: 1) it
carries the insecticide into the root zone; and 2) it encourages mole
crickets to be active in the upper layer of soil where they can come
into contact with the poison. Control with any of these treatments is
best during the summer months for reasons previously explained.
Ethoprop (Mocap) and isofenphos (Oftanol) are the best chemical
treatments for mole crickets in turf. Isofenphos offers up to several
months of control with a single application and is most efficient when
applied early in the summer. Propoxur (Bayon) and diazinon
(Sarolex) sprays are not as effective in turf as other registered for-
mulations.


Evaluation of chemical control
In the past, the effectiveness of chemical controls has been evalu-
ated by counting dead or moribund crickets on the surface. This can
be misleading, since many crickets die below the surface. A chemical
causing high mortality below ground could be rated ineffective. Addi-
tionally, the initial population is unknown, and movement into and
out of the treated area is not accounted for. A technique developed to
evaluate true chemical efficacy involves placing a known number of
mole crickets in soil-filled cages or containers buried at soil level
prior to treatment. Control can be accurately determined by counting
the mortality both above and below the surface. This technique,
though labor-intensive, is the most reliable means of comparing
toxicity of chemical treatments.
Flushing live crickets from the soil with mild soap and pyrethrin
solutions can also be used to evaluate chemical control. Because of
the inefficiency of flushing, only a relative measure of control can be
obtained. Other techniques involve measuring the extent of tunnel-
ing/damage or the number of plants destroyed. Like the flush, these
techniques give only a relative measure of control but are useful for
evaluating the long-term effects of chemical treatments.


46









Future Prospects

Unless resistant grass varieties or introduced biological control
agents become established and permanently reduce the economic
impact of mole crickets, chemical insecticides will remain indispen-
sable for controlling mole cricket populations. At present, chemicals
provide the only known means for quickly suppressing large num-
bers of crickets with predictable results.
Sprays and granules are the most effective insecticidal treatments
for mole crickets, but their use will remain expensive and will be
limited to non-forage crops. However, the recent registration of
isofenphos, a long-term residual insecticide, will help reduce the cost
of control in turf by lessening the need for additional treatments later
in the season.
Because of recent concerns over indiscriminate and excessive use of
pesticides, chemical control with poison baits has received renewed
emphasis and is most promising for several reasons. Not only are
baits relatively cost effective, but they offer an efficient and ecologi-
cally selective use of broad spectrum insecticides. Though they are
the safest and most economical method of control, poison baits are,
nonetheless, too expensive for practical application in pastures. Cat-
tlemen and hay producers are left with no acceptable means of con-
trolling mole crickets. There are some approaches promising more
affordable chemical control.
1) As described in an earlier section, costs can be dramatically
reduced by using less expensive materials in formulation, namely
carriers and toxicants. A bait formulated from peanut hulls, brewers
concentrate, and 2% malathion, and applied at recommended rates
(1-2 lb A.I./acre), could cost as little as $5.00/acre.
2) The use of applicator-formulated baits offers a significant sav-
ings in material costs. A user-formulated 2% malathion bait (Table 6)



Table 6. Formulation and approximate cost of materials for a
applicator-formulated 2% malathion bait.
Material Amount* Approximate cost**
Laying mash 100 lbs $13.50
Crude molasses 2 qts $ 0.50
Water 1-5 qts -
Malathion 2 lbs $ 6.00
TOTAL $20.00
*Formula developed by P. G. Koehler and D. E. Short.
**Based on 1983 prices of quantities locally available to growers.









will save the grower more than 50% in the cost of materials when
compared to a similar preformulated bait (Table 5) applied at the
same rate.
3) Recent research suggests that the amount of material necessary
for effective control can be substantially reduced. Chlorpyrifos 0.5%
bait was 85% effective when applied at less than one-third the recom-
mended rate (0.2 lb A.I./acre) to an extremely high population den-
sity (100 crickets/sq m). Laboratory tests have shown that malathion
baits are effective at rates as low as 0.5 lb A.I./acre. Additionally the
use of an applicator formulated 2% malathion bait (Table 6) has
given excellent control in pastures when applied at 0.4 lb A.I./acre,
less than half the recommended rate.
Though chemical control can be made more economical, it can only
offer temporary relief. More permanent solutions are necessary to
eliminate the severe impact of mole crickets. With pesticides becom-
ing increasingly expensive and their environmental side effects of
greater concern, selective and intelligent use of insecticides will be
essential to mole cricket management in the future.

Addendum
Tests completed in 1984 proved that a single application of
grower-formulated 4% malathion bait could give season-long control
of mole crickets for as little as $3.50/acre in material costs.
The most effective bait tested was similar to the one described in
Table 6, but was formulated with twice as much malathion (4%) and,
instead of molasses, incorporated 5% crude cottonseed oil and 10%
sucrose. Lab trials established that cottonseed oil and sucrose in-
duced more feeding than molasses, and small-scale field trials de-
monstrated that 4% malathion was more efficacious than 2% (when
applied at the same A.I. / acre rate). Persistence studies found that the
4% bait retains half its toxicity for as long as 30 days under field
conditions. In full-scale field trials in heavily infested bahiagrass
pastures, one application of grower-formulated bait at 0.5 lb A.I./
acre gave greater than 95% control for at least two months.












V. FURTHER READING
Much of the material in this bulletin has been published in greater
detail elsewhere. References of special importance to the section on
"Systematics and Life Cycles" are Hayslip 1943; Ngo and Beck 1982;
Nickle and Castner 1984; Ulagaraj 1975; Walker and Fritz 1983;
Walker and Nickle 1981; Walker, Reinert, and Schuster 1983. Refer-
ences of similar importance to "Reproductive Behavior" are Forrest
1980, 1983a; Ulagaraj 1976; Ulagaraj and Walker 1973, 1975; and
references to "Other Behavior, Damage, and Sampling" are Law-
rence 1982; Matheny 1981; Matheny, Kepner, and Portier 1983;
Reinert 1983a; Short and Koehler 1979; Walker 1982; Walker and
Dong 1982; Williams and Shaw 1982.
References pertinent to "Biological Control of Mole Crickets" in-
clude Adamson 1942; Castner 1984; Mangold 1978; Smith 1935;
Williams 1928; Wolcott 1938, 1939, 1940, 1941; and references to
"Chemical Control of Mole Crickets" include Koehler and Short
1976a, 1976b; Short and Driggers 1973; Short and Koehler 1980b;
Ulagaraj 1974.
Finally, four earlier bulletins have dealt extensively with the
biology of introduced pest mole crickets. Two of these concern U.S.
pest species (Worsham and Reed 1912, Thomas 1928) and two con-
cern similar Puerto Rican species (Barrett 1902, Van Zwaluwenberg
1918). A few aspects of mole cricket biology and control reported in
these bulletins have not been reported elsewhere. The same is true of
the current bulletin, which includes the results of previously unpub-
lished research at the University of Florida.












REFERENCES

References marked with an asterisk (*) are project-related publications of
research results; those marked with a dagger (t) are master's theses written
by students working on the project. Project-related, general-information
publications are not listed here because they are superceded by this bulletin.

Adamson, A. M. 1942. Mole cricket parasites of the genus Larra in Trinidad.
Rev. Appl. Entomol., Ser. A. 30:515.
Barrett, O. W. 1902. The change, or mole cricket (Scapteriscus didactylus
Latr.) in Porto Rico. Porto Rico Agri. Exp. Stn. Bull. 2:1-19.
*Beugnon, Guy. 1981. Orientation of southern mole cricket, Scapteriscus
acletus, landing at a sound source. Fla. Entomol. 64(4):463-468.
tCastner, J. L. 1983. Biology and ecology of the mole cricket parasitoid Larra
bicolor. 80 pp.
*Castner, J. L. 1984. Suitability of Scapteriscus spp. mole crickets (Orthop-
tera: Gryllotalpidae) as hosts of Larra bicolor (Hymenoptera: Sphecidae).
Entomophaga 29 (3):323-329.
*Castner, J. L., and H. G. Fowler. 1984. Gut content analyses of Puerto Rican
mole crickets (Orthoptera: Gryllotalpidae: Scapteriscus). Fla. Entomol. 67
(3):479-481.
*Castner, J. L., and H. G. Fowler. 1984. Distribution of mole crickets
(Orthoptera: Gryllotalpidae: Scapteriscus) and the mole cricket parasitoid
Larra bicolor (Hymenoptera: Sphecidae) in Puerto Rico. Fla. Entomol. 67
(3):481-484.
*Castner, J. L., and J. L. Nation. 1984. Cuticular lipids for species recogni-
tion of mole crickets (Orthoptera: Gryllotalpidae) I. Scapteriscus didacty-
lus, Scapteriscus imitatus, and Scapteriscus vicinus. Fla. Entomol. 67:
155-160.
*Forrest, T. G. 1980. Phonotaxis in mole crickets: its reproductive signi-
ficance. Fla. Entomol. 63(1):45-53.
tForrest, T. G. 1981. Acoustic behavior, phonotaxis, and mate choice in two
species of mole crickets (Gryllotalpidae: Scapteriscus). 72 pp.
*Forrest, T. G. 1983. Calling songs and mate choice in mole crickets. Pages
185-204 in D. T. Gwynne and G. K. Morris, eds. Orthopteran mating
systems: sexual competition in a diverse group of insects. Westview,
Boulder, Colo.
*Forrest, T. G. 1983. Phonotaxis and calling in Puerto Rican mole crickets
(Orthoptera: Gryllotalpidae). Ann. Entomol. Soc. Am. 76(4): 797-799.
*Fowler, H. G., and J. N. Kochalka. 1985. New Record of Euphasiopteryx
depleta (Diptera: Tachinidae) from Paraguay: Attraction to broadcast calls
of Scapteriscus acletus (Orthoptera: Gryllotalpidae). Fla. Entomol. 68
(1):225-226.
*Fritz, G. N. A technique for separating mole crickets from soil. Fla.
Entomol. 66: 360-362.
tGreen, M. E. 1982. Laboratory and field methods for evaluating toxic baits
for control of the mole crickets, Scapteriscus acletus and Scapteriscus
vicinus. 39 pp.








Hayslip, N. C. 1943. Notes on biological studies of mole crickets at Plant City,
Florida. Fla. Entomol. 26(3):33-46.
tKepner, R. L. 1981. Evaluation of bait attractants for mole crickets Scapter-
iscus vicinus and S. acletus (Orthoptera: Gryllotalpidae). 38 pp.
*Kleyla, P. C., and G. Dodson. 1978. Calling behavior and spatial distribu-
tion of two species of mole crickets in the field. Ann. Entomol. Soc. Am.
71(4):602-604.
Koehler, P. G., and D. E. Short. 1976. Control of mole crickets in pas-
turegrass. J. Econ. Entomol. 69(2):229-232.
Koehler, P. G., and D. E. Short. 1976. Pasture mole cricket control with an
applicator-formulated bait. Fla. Entomol. 59(2):180-182.
Lawrence, K. L. 1982. A linear pitfall trap for mole crickets and other soil
arthropods. Fla. Entomol. 65(3):376-377.
*Mangold, J. R. 1978. Attraction ofEuphasiopteryx ochracea, Corethrella sp.
and gryllids to broadcast songs of the southern mole cricket. Fla. Entomol.
61(2):57-61.
*Matheny, E. L., Jr. 1981. Contrasting feeding habits of pest mole cricket
species. J. Econ. Entomol. 74:444-445.
*Matheny, E. L., Jr., and R. L. Kepner. 1980. Maxillae of the mole crickets,
Scapteriscus acletus Rehn and Hebard and S. vicinus Scudder (Orthoptera:
Gryllotalpidae): a new means of identification. Fla. Entomol. 63(4):512-
514.
*Matheny, E. L., Jr., R. L. Kepner, and K. M. Portier. 1983. Landing distribu-
tion and density of two sound-attracted mole crickets (Orthoptera: Gryllo-
talpidae: Scapteriscus). Ann. Entomol. Soc. Am. 76(1):278-281.
*Matheny, E. L., Jr., A. Tsedeke, and B. J. Smittle. 1981. Feeding response of
mole cricket nymphs (Orthoptera: Gryllotalpidae: Scapteriscus) to radio-
labeled grasses with, or without, alternative food available. J. Ga.
Entomol. Soc. 16:492-495.
*Nation, J. L. 1983. Specialization in the alimentary canal of mole crickets.
Int. J. Insect Morph. Embryol. 12(4):201-210.
*Ngo, Dong and H. W. Beck. 1982. Mark-release of sound-attracted mole
crickets: flight behavior and implications for control. Fla. Entomol.
65(4):531-538.
*Nickerson, J. C., D. E. Snyder, and C. C. Oliver. 1979. Acoustical burrows
constructed by mole crickets. Ann. Entomol. Soc. Am. 72(3):438-440.
*Nickle, D. A., and J. L. Castner. 1984. Introduced species of mole crickets in
the United States, Puerto Rico, and the Virgin Islands (Orthoptera: Gryllo-
talpidae). Ann. Entomol. Soc. Am. 77:(4):450-465.
*Reinert, J. A. 1983. Foraging sites of the southern mole cricket, Scapteriscus
acletus (Orthoptera: Gryllotalpidae). Proc. Fla. State Hort. Soc. 96:
149-151.
*Reinert, J. A. 1983. Mole crickets are damaging to turfgrass and
ornamentals. Proc. South. Nurserymans Assoc. Res. Conf. 28:150-151.
*Schuster, D. J. 1981. Mole cricket control in tomato, 1980. Insecticide
Acaricide Tests 6:98-99.
*Schuster, D. J. 1982. Mole cricket control in tomato, 1981. Insecticide
Acaricide Tests 7:122.
*Schuster, D. J. 1983. Mole cricket control in tomato, 1982. Insecticide
Acaricide Tests 8:147.
*Schuster, D. J. 1984. Mole cricket control in tomato, 1983. Insecticide
Acaricide Tests 9:199-200.








Short, D. E., and D. P. Driggers. 1973. Field evaluation of insecticides for
controlling mole crickets in turf. Fla. Entomol. 56(1):19-23.
*Short, D. E., and P. G. Koehler. 1979. A sampling technique for mole
crickets and other pests in turfgrass and pasture. Fla. Entomol. 62(3):282-
283.
*Short, D. E., and P. G. Koehler. 1980. Evaluation of pesticides against mole
crickets. Insecticide Acaricide Tests 5:196.
*Short, D. E., and P. G. Koehler. 1980. Location as a variable for evaluating
pesticides against mole crickets. J. Ga. Entomol. Soc. 15:241-245.
*Short, D. E., and J. A. Reinert. 1982. Biology and control of mole crickets in
Florida. Pages 119-124 in H. D. Neimczyk and B. G. Joyner, eds. Advances
in turfgrass entomology. Hammer Graphics, Piqua, Ohio.
Smith, C. E. 1935. Larra analis Fabricius, a parasite of the mole cricket
Gryllotalpa hexadactyla Perty. Proc. Entomol. Soc. Wash. 37(4):65-87.
*Taylor, T. R. 1979. Crop contents of two species of mole crickets, Scapteris-
cus acletus and S. vicinus (Orthoptera: Gryllotalpidae). Fla. Entomol.
62(3):278-279.
Thomas W. A. 1928. The Porto Rican mole cricket. USDA Farmers' Bull.
1561:1-9.
tTsedeke, Abata. 1979. Plant material consumption and subterranean move-
ments of mole crickets (Orthoptera: Gryllotalpidae: Scapteriscus) as deter-
mined by radioisotope techniques with notes on materials for laboratory
feeding. 72 pp.
Ulagaraj, S. M. 1974. Effects of insecticides on mole crickets (Orthoptera:
Gryllotalpidae: Scapteriscus). Fla. Entomol. 57(4):414.
Ulagaraj, S. M. 1975. Mole crickets: ecology, behavior, and dispersal flight
(Orthoptera: Gryllotalpidae: Scapteriscus). Environ. Entomol. 4(2):265-
273.
Ulagaraj, S. M. 1976. Sound production in mole crickets (Orthoptera: Gryllo-
talpidae: Scapteriscus). Ann. Entomol. Soc. Am. 69(2):299-306.
Ulagaraj, S. M., and T. J. Walker. 1973. Phonotaxis of crickets in flight:
Attraction of male and female crickets to male calling songs. Science
182:1278-1279.
Ulagaraj, S. M., and T. J. Walker. 1975. Response of flying mole crickets to
three parameters of synthetic songs broadcast outdoors. Nature
253(5492):530-532.
Van Zwaluwenberg, R. H. 1918. The change or West Indian mole cricket.
Porto Rico Agric. Exp. Stn. Bull. 23:1-28.
tWalker, S. L. 1979. Population estimation, damage evaluation and be-
havioral studies on mole crickets Scapteriscus vicinus and S. acletus
(Orthoptera: Gryllotalpidae). 83 pp.
*Walker, T. J. 1982. Sound traps for sampling mole cricket flights (Orthop-
tera: Gryllotalpidae: Scapteriscus). Fla. Entomol. 65(1):105-110.
*Walker, T. J., and G. N. Fritz. 1983. Migratory and local flights in mole
crickets, Scapteriscus spp. (Gryllotalpidae). Environ. Entomol. 12(3):953-
958.
*Walker, T. J., R. C. Littel, and Ngo Dong. 1982. Which mole crickets damage
bahiagrass pastures? Fla. Entomol. 65(1):110-116.
*Walker, T. J., and J. L. Nation. 1982. Sperm storage in mole crickets: fall
matings fertilize spring eggs in Scapteriscus acletus. Fla. Entomol.
65(2):283-285.
*Walker, T. J., and Ngo Dong. 1982. Mole crickets and pasture grasses:








damage by Scapteriscus vicinus but not by S. acletus (Orthoptera: Gryllo-
talpidae). Fla. Entomol. 65(3):300-306.
*Walker, T. J., and D. A. Nickle. 1981. Introduction and spread of pest mole
crickets: Scapteriscus vicinus and S. acletus reexamined. Ann. Entomol.
Soc. Am. 74:158-163.
*Walker, T. J., J. A. Reinert, and D. J. Schuster. 1983. Geographical varia-
tion in flights of mole crickets, Scapteriscus spp. (Orthoptera: Gryllotalpi-
dae). Ann. Entomol. Soc. Am. 76(3): 507-517.
Williams, F. X. 1928. Studies in tropical wasps-their hosts and associates
with descriptions of new species. Entomol. Ser., Hawaii Sugar Planters'
Assoc. Exp. Stn. Bull. 19:1-179.
*Williams, J. J., and L. N. Shaw. 1982. A soil corer for sampling mole
crickets. Fla. Entomol. 65(1):192-194.
Wolcott, G. N. 1938. The introduction into Puerto Rico of Larra americana
Saussure, a specific parasite of the changea," or Puerto Rican mole-cricket,
Scapteriscus vicinus Scudder. J. Agri. Univ. Puerto Rico 22(2):193-218.
Wolcott, G. N. 1939. A parasite of the Puerto Rican mole-cricket. Science 89
(2318):508-509.
Wolcott, G. N. 1940. A Tachinid parasite of Puerto Rican change. J. Econ.
Entomol. 33(1):202.
Wolcott, G. N. 1941. The establishment in Puerto Rico of Larra americana
Saussure. J. Econ. Entomol. 34:53-56.
Worsham, E. L., and W. V. Reed. 1912. The mole cricket (Scapteriscus
didactylus Latr.). Ga. Agri. Exp. Stn. Bull. 101:251-263.
*Yu, S. J. 1982. Microsomal oxidases in the mole crickets, Scapteriscus
acletus Rehn and Hebard and Scapteriscus vicinus Scudder. Pest. Biochem.
Physiol. 17:170-176.










































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