PARASITES IN LEAP HOPPERS FRO!', KANSAS GRASSLANDS (HOMOPfERA:CICAD£LLIDAE)
by
ROBERT SHOOT BALDRIDGE
B. S., Baylor University, 1366
A MASTER'S THESIS
submitted in partial fulfillment of the
requirements for the deqrze.
MASTER OF SCIENCE
Department of Entomology
KANSAS STATE UNIVERSITY Ken hat tan, Kansas
1569 :j?d by: , )
•
CS ^"—oO TABLE 0F CONTENTS
INTRODUCTION • 1
LITERATURE REVIEW 2
Dryinidae (Hymenoptera) 2
Hal ictophagidae (Streps iptera) , 2,
Dorilaidae (Diptera) c
MATERIALS AND METHODS . 3
Grass'and Collections 3
Procedure for Handling the Dead Parasitized Leafhoppers ; jq
Rearing Cages ,,
Procedure for Handling Live Leafhopper Material \k
Preparation of Adult Dryinids for Identification 18 RESULTS AND DISCUSSION 19
Kansas Collection Material 19
Location of Dryinid Larval Sacs in Leafhopper Hosts 23
Position of Strepsiptera Parasites In their Hosts •• 25
Summary of Identified Strepsiptera from Kansas Hosts 25
Notes on a Male Dryinid (Tribe: Gona topod i n I in its Late Larval to Adult Develooment .. •••• 27 Rearing of Dryinids from Exitia nus ex_itjosus~ (Uhler) from Bermuda ..,.". grass , ...... ,1
Dryinids from Col lect ions and Rearing Studies 3]
Parasitic Worms Noted in Kansas Leafhoppers 3/, 1 I I
SUMMARY AND CONCLUS IONS 35
ACKNOWLEDGMENTS 39
LITERATURE CITED i4
APPEND IX Z»3 INTRODUCTION
The leafhopper family Cicadellidae is one of the largest in North America
and the world (DeLong, 19^3). They use varied groups of plants as food and
oviposit ion hosts. Certain species are of major economic importance. Most
work involving economic effects has dealt with species infesting high in-
come crops such as sugar cane, rice, sugar beets, apples and grapes.
Oman (IS^) listed four major types of leafhopper injury to plants:
(1) damage caused by the transmission of virus diseases; (2) introducing
foreign matter during feeding which mechanically interrupted the normal
translocation in plants; (3) direct removal of liquid plant foods by feed-
ing; and, C<) such injury as was caused by oviposition and defecation.
The injuries inflicted by leafhoppers are usually not as obvious as
those of other herbivorous insects (Sanders and DeLong, 1920). Annual
damage to grazing and hay production by high populations in grassland is
probably of economic proportion (Osborn, 1915; Fattig, 1955; Young, 1959).
Leafhoppers can be partially controlled by man and insect agents.
Agricultural practices can be applied to suppress large populations of
leafhoppers (Osborn, 1915). These insects are also controlled to vary-
ing degrees by insect predators and parasites, Limited work has been done to determine predators and parasites operating in grassland biomes, particular!/ in Kansas.
This study was an attempt to determine parasites found infecting leafhoppers in native prairie grasslands in Kansas, using both field collecting and rearing data. The discussion was limited primarily to .
three orders of leaf hopper parasites: Hymenoptera, Streps iptera and
Diptera. These forms are those usually listed as parasitizing nymphal
and adult leafhoppers (Lawson, 1320), and were the most frequently
collected kinds of parasites.
LITERATURE REVIEW
The study of parasites of leafhcppers from the grasslands of Kansas
has been neglected and only occasional references are noted in taxo-
nomlc treatments of leafhoppers occurring in the grassland biome. These
usually concerned the three main groups of internal parasites (Lawson,
1320) or the presence of immature forms of one particular group (Blocker,
1367).
Dryinidae (Hymenoptera)
Members of the family Dryinidae are primarily internal parasites of
auchenorhynchus Homoptera (Richards, 1333). The life histories of this
group have been studied more than the other two parasitic groups to be
mentioned
Perkins (1305, 1306, 1912) and Fenton (1918) have done extensive work in many areas of dry in id biology and systemai ics . Perkins dealt with Hawaiian dryinids and species sent to him bv his co-workers in the
United States; Fenton primarily studied North American Dryinidae.
Richards (1333) did a taxonomic study of the British Dryinidae.
Douglas (1830), Ajnslie (1320), Centner (1326), Esaki and Hashimoto
Stoiner (1335), 0336), Know! Ion (1337), Cook (19*1) ""d Barrett et ,
al . (1$65) have studied the biology of various dryinid parasites of leaf-
hoppers, mostly of the genera Aphelopus Palm., Gonatopus Ljungh and
Pachygonatopus Fenton
According to Fenton (1918) adult drylnids were found in grass, forbs
and woody vegetation. Wingless females sought their prey on foot while
winged forms flew actively in search of prey. The female upon nearing
the prey stopped, made a sudden jump at the prey, and seized it with
the chelate front tarsi. She then stung the host, causing a paralyzed
condition. The female dryinid might: (1) consume the host; (2) ovi-
posit in the host through either the cuticle or intersegmental membrane;
or (3) release the host unharmed., The reason for the latter was unclear.
Richards (1939) indicated that five larval stages were probably
involved in most of the life cycles. Barrett et al . (I965) recorded
five larval instars In Pachygonatopus minimus Fenton. The first four
molts, which were not typical larval molts, took place in the body of
the host. Each preceding larval exuvium was split longitudinally down
the back, yielding an exuvium which adhered to the body of the next larval
instar. The succeeding larval stage darkened, giving the appearance of a tab or sac on the body of the host. The last larval stage liquified and consumed the body contents of the host, dropped from the host and pupated in a cocoon of silken material (Fenton, 1913; Barrett et al., 13&5)
The last larval molt was the only typical one (Barrett et al., 1 965) -
The larvae moved by means o[ peristal ic body movements in search of suitable pupation habitats (Fenton, 1918). The cocoon might be formed on vegetation, at soil level, or in the soil. It miaht be formed r )
y>
only of silken material or have soil particles or incorporated parts of
the larval sac.
Only the female is known to actively prey on leaf hoppers. Both
sexes, however, feed on sugar solutions (Fenton, 1918). In nature,
certain d'ryinids feed on honeydew secreted by their hosts (Perkins,
1905).
The females are longer-lived than the males. Barrett et at. ( 1 965)
recorded females of P_. minimus Fenton as living from 5 to k6 days. The
males lived from less than a day to four days, probably surviving only
long enough to copulate (Perkins, 1905). Very little is known of male
life history (Richards, 1953),
Pathogenetic reproduction occurred to a large extent in this
group (Fenton, 1918); this was based on laboratory rearing data and the
low number of males recorded from field collections. Polyembryony also
has been recorded from one species.
The effect of this parasitic family as a controlling factor of leaf-
hopper populations is uncertain. Field collections may indicate errone-
ously low or high populations of these parasites on leafhopper hosts (Law-
son, 1920). The controlling effects of P_. minimus Fenton on Macrosteles
fasc if ron s (Stal), with 37% parasitism recorded, were considered un-
important (Barrett et al., 1965). "
Hal ictophagidae s t ( Scrcp i p e a
Leafhopper parasites of the order Strepsiptera have been reported by Pierce (1909, 1918) and Bohart (1941, 19*0). The latter presented the most comprehensive treatment of North American Strepsiptera and includes
a summary of leafhopper hosts, systematics and life history information.
The family Hal ictophag idae is the primary stylopid parasite (Bohart,
12^3) and exhibits a high degree of sexual dimorphism. This informa-
tion, as wel i as morphological differences between stages in tiie develop-
ment of larval and pupal forms, was reviewed by Pierce (1903) and Bohart
(I94t , 1343).
The female of Hal ictophagidae lacks antennae, eyes, legs, wings, palpi
and distinct body segments (Bohart, 19'4l). It is permanently sealed in
its puparium within the host and appears externally as a flattened, semi-
circular to oblong tab (the cepha lothorax) on the body of the host. The
abdomen, within the host, is devoid of obvious characters except for one
to three brood canals.
The males are free-living, and seldom live more than two days
(Bohart, 19^1). They have wel 1 -developed fan-shaped metathoracic wings.
The mesothoracic wings are reduced to pseudohal teres.
A typical life, cycle was described by Bohart ( 1 94 1 ) . The female was actively sought by the male. Copulation took place through an aperture in the cephalothoi ax of the female. The sperm fertilized a mass of eggs developing simultaneously within the abdomen of the female.
Ultimately one to five thousand larvae developed and escaped through the cephalic aperture. These triungulin larvae got onto vegetation and awaited a host. They used r.heir sharply-edged heads to penetrate the body of the host, where a series of hypermetamorphic molts took place prior to adult emergence. T hose that became males formed an exarate pupa inside the last larval sku-,. The pupal case partially protruded from the body of the host as a rounded capsule. The end cap of this . )
puparium was pushed off by the mature male prior to its escape from the
host.
The existence of parthogenesi s was discussed by Pierce ( 1 9 1 8) and
Schrader (192*0 . It appears, however, that bisexual reproduction Is more
common
Do ri lai dae (Dipte r a
Dorilaidae (Pipuncul idae of authors) was the only family of Diptera
recorded as internal parasites of leaf hoppers. The larval stage occurs
in the host.
Dorilaidae were recorded from the homopterous families C icadel 1 idae,
Fulgoridae and Cercopidae (Hardy, 1 9^3) in various parts of the world.
Curran ( 1 365) included the unconfirmed observation that the hemipterous
family Mir idae also served as a host.
Hardy ( 1 9^3) , in his Nearctic treatment of this family, indicated
that these flies are common in grasslands. Almost any grassy area yielded
one to many species of adults.
The biology of certain of these insects affecting leafhoppers has been
dealt with by Douglas (l8go), Keilin and Thompson (1915), Severin (1924) and
Knowlton (1924). These works and that of Hardy ( 1 9^3) should be consulted
for life, history information. Few American species have been reared and
their life histories studied. Perkins (1905, 1906) associated 15 of 26
Dorilaidae species with their leafhopper hosts in Australia and Hawaii.
Hardy ( listed several i 3^3) genera and species with which he associated proven or suspected leafhopper hosts. .
The larvae are the life forms affecting the leafhopper directly.
Hardy (l3'+3) and others listed above describe such life forms. The
larvae are oblong. The posterior end is rounded slightly whereas the
anterior end is slightly pointed. Body segmentation is indistinct.
No externa] mouthparts are present; each larva has a pair of heavily
sclerotized mandibles or "mouth hooks". These appear in the first
instar and are shed with the last larval molt. The larvae are amphi-
neust ic.
Larvae develop from eggs deposited in both nympha! and adult leaf-
hcppers. Larvae develop within the abdomen of the host. They may com-
pletely fill the abdomen upon maturing, and extend into the posterior
thoracic segments. The mature larva breaks out of the host's abdomen
(Hardy, IS'^3), usually at its junction with the thorax (Oman, 13^9), or
at other points along the abdomen (Parker, 1 367) - This emergence kills th<
host
Pupation may occur on the soil surface, in the soil, or on plant
parts (Hardy, 13^3). The puparium is formed from the last larval skin.
These parasitic forms are not as easily detected as Dryinidae or
Hal ictophagidae. Distention of the infected host's abdomen and its slug-
gish behavior may be clues to parasitized conditions (Oman, 1 9^3) . How-
ever, a gravid female may have an appearance similar to that of a para-
sitized form (Hardy, 1943).
This family contains species which probably exert some control of
leafhopper populations. Until more biological studies are carried out, however, their importance cannot be estimated adequately. -
Henderson (1355) discussed possible methods for dissemination of
these three internal parasites of leaf hoppers based on collections of
parasitized beet leaf hoppers , Ci rcul i fer tenel lus (Baker). He attempted
to determine what effects p3rasi t izat ion had on distance traveled in their
annual migrations. It appeared that, even though wind dispersion accounted
for some adult movement of these parasites, dispersion was primarily de-
pendent on the movements of the larvae within the host. This was particular-
ly probable in the case of Dryinidae and Strepsiptera . Dorilaidae also
depended on this method for dispersion, but adults, being good fliers, may
have been carried by the wind.
MATERIALS AND METHODS
l fir assland Col lections
A survey of Kansas grasslands, involving 36 collections from 55 coun-
ties, was conducted between August 23, 1367, and September 2k, 1 367
Grasslands were sampled for leafhoppers using a round sweep net.
Each survey collection consisted of 120 sweeps. One sweep of the net was made approximately every three feet, or average walking stride. The
sampling was made in a square quadrat of 30 sweeps per side. Each sample
included as many of the existing grass types as noted in the collection ares. No attempt was made to determine population densities of either
leafhoppers or parasites.
Collected material from each site was placed in a small paper sack,
labeled and stored until separated. Parad ichlorobenzene (PDB) was added
to each sack to insure kill of any arthropod material, especially .
dermestid larvae and spiders. The sweep net bag was reversed after each collection to minimize contamination from the prior collection.-
At each site the predominant grasses were recorded, particularly those which had wel 1 -developed seed heads at the time of the survey.
Ho recording of specific forbs was made; such plants were recorded per collection as "forbs". Almost none of the collection sites was devoid of forbs.
Further collections were made involving four reseeded upland pastures at Fort Hays Branch Experiment Station, Hays, Kansas. Collections were made at irregular intervals from 13^7 through 1968.
The pastures collected contained the following grass species:
1. A native mixture pasture of Andropogon gerardi
(big bluestem), A_. sc opar i us (little bluestem),
Eouteloua curt i pen dula (sideoats grama),
8_. 9£a_cj_]js_ (blue grama), Buch loe dactyloides
(buf falograss) , and Pan icum v? rga tum (switchgrass).
2. A pasture of Andropogon intermedius (Caucasian bluestem).
3- A pasture of Ag ropyron smi thl i (western wheatgrass)
h. A pasture of primarily Panicum vi rgatum (switchgrass).
These pastures had been seeded initially in April, 1957. Grazing studies were in progress on these pastures and had been carried out in previous years. Each pasture had an area fenced off from "the yrazing area. Collections were made in these areas also. Due to the low in- cidence of leafhoppers in the Caucasian bluestem pasture, this pasture was excluded during the 1968 collecting season. 10
Two types of collections were obtained from the Fort Hays area. One
100-sweep collection of dead leafhoppers which included the adult para-
sites was mailed to Manhattan, Kansas from Hays, Kansas in pint card-
board ice cream cartons from three of the pastures; one 50-sweep collec-
tion was taken from the Caucasian bluestem pasture. One "live" collec-
tion per pasture was mailed at the same time in quart cardboard ice cream cartons lined with wet paper towels. A small amount of vegetation from
each site was placed in each carton. This vegetation provided a surface for
the leafhoppers to rest upon and tended to keep them off the wet paper
towel. The number of sweeps per "live" collection varied during the study
but was never less than 100.
Various collections were also made during this study in Donaldson
Pasture, a research pasture near Manhattan (Riley county), Kansas. Areas
of buf falograss , blue grama, bluestem and switchgrass were collected.
The grassland insect collections made by W. H. Arnett during his
study of grasshoppers in Donaldson Pasture ( 1 960) were made available
for studying the contained leafhopper material. Material from the re- search collection of the Kansas State University Entomology Department was also used.
Procedure for Handling t_he Dead Parasitized Leafhopper s
The dead leafhopper material was removed and stored dry in empty plastic vials. They were viewed under a binocular microscope. The externally-parasitized leafhoppers were removed and mounted.. External examination was sufficient to detect late larval Drylnldae and adult.
Hal ictophagJda.e. . .
11
it was necessary to clear a sample (approximately half) of each
collection (minus the externally-parasitized materials) in order to find
internal larval stages of parasitic Dorilaidae and early stages of
Dryinidae and Hal ictophagidae
Samples were cleared in a cold potassium hydroxide solution for one
hour, then immersed in glycerine and viewed under a dissecting micro-
scope. Any parasitized leafhoppers were removed and stored in glycerine
in covered deep-well slides for future species determinations.
Since there are no keys to the larval stages of Dryinidae or
Dorilaidae, only the host species (if discernible) and the stage of the
parasite were recorded. The stage of the host, type of parasite, para-
sites per host, location in the host and the effects of parasi t izat ion,
if any, on genital ic structures were noted. The stylopized leafhoppers
were forwarded to R. M. Bohart for determination of the parasite species.
Re arl ng Cages
Cellulose nitrate cages were used for the rearing of larval Dryinidae
to adults. Each cage (Fig. 1 ,a) was kS mm long and 29 mm in diameter.
Two of the cages were cut from each cellulose nitrate centrifuge tube
(Lusteroid, Curtin Co., catalogue number 659) 29 mm in diameter and 105 mm
long
Each cage was fitted with a foam rubber plug (13 mm thick and 30 mm in diameter) at each end (Fig. 1, b and b ) to prevent the escape of the
leaf hopper. A slit was made in each plug from its center through the outer edge to facilitate insertion of a grass blade into the C3ge. EXPLANATION OF FIG, 1
a = rearing cage
= rubber plug b, , b, foam
d in c - cork in hole in cage
e = metal support rod
f - rubber band
g = six-inch clay pot containing grass 13
Fig. 1 . Rearing cage. H
A small hole (Fig. 1 ,c) , made with a standard paper hole-punch, was positioned in the center of the cage side. Introduction and removal of the insect was accomplished by means of a six-inch straight tube glass aspirator inserted through this hole. The hole was plugged with a
microvial cork (Fig. 1 ,d) .
Each cage and its included plant and leafhopper were held in an up-
right position, attached by means of a rubber band (Fig. 1 ,f) to a metal
rod (Fig. 1 ,e) . The metal rod was then inserted into the soil to a depth of about three inches.
Live Leafhopper Material ?1PS^- rJL I°L-. Handl ing
In order to obtain identifiable adult material, attempts were made to rear dryinids from their leafhopper hosts. The parasitized leafhoppers were collected primarily from the buffalo and bluestem grasses in Donaldson
Pasture and from the western wheatgrass pasture in Hays, Kansas.
Live material was aspirated from the sweep net and placed in cardboard ice cream cartons containing a small amount of vegetation and lined with wet towsls. Care was taken to prevent direct sunlight from striking the containers, as this often resulted in death to the leafhoppers.
This 1 ive materia! was taken into the greenhouse and transferred to a large glass chimney cage atop a six-inch pot containing the following
grasses: buf falograss , switchgrass, big blutistem, little bluestem and sideoats grama. These plarts were grown from seeds in flats and when they attained healthy growth (about six inches tall) they were transplanted in the six-inch pots. The grass species were so planted in the pots as to be separated from each neighboring gra:;s. .
15
The top of the chimney cage was covered with a piece of cheesecloth to
prevent escape of the leafhoppers. A small hole was cut in the cheesecloth
to facilitate introduction of an aspirator tube for removal of individual
parasitized leafhoppers. The hole was plugged with cotton.
Leafhoppers were maintained In this cage for 2h hours, at the end of which time observations were made to determine the presence of any exter- nally-visible parasites. These leafhoppers were then aspirated from the grass species upon which they were feeding and placed individually into
the cellulose nitrate cages attached to the same grass species. The cages were situated at the upper level of the plant growth and labeled with the collection locality (or the predominant grass type from which collected) and consecutively numbered.
They were observed daily, noting the development of the larvae, emergence of the mature larvae, and/or the death of the host.
The remaining leafhoppers in the chimney cage were allowed to remain undisturbed for one week. This allowed the previously undetected dryinid
larva to become externally visible on the host. Such hosts were removed and treated as above. Few groups of leafhoppers lived more than a week.
At the end of this period of time the remaining leafhopper material and the grass in the cage were destroyed. Fresh grass was used for each large collection of leafhoppers.
Individually parasitized leafhoppers from grasslands Were treated as above when there was doubt as to the possible feeding host. However, those which were collected from relatively pure stands of grass, such as buffalo- grass, were placed on the corresponding grass in the individual rearing cages 16
Win, studies wer. carried out under both ^ ^^ ^ 9rCenh0USe ^' a - " «'»« -Perature. — rBlatiw h„id ity or day length were enforced.
Observations were mademaa^ at^«- -r.„ approx.mately 9 :00 AM, , 2 .-co and 5:00 PM to in- jure removal of any mature larva, before the,'«> ppupated.UP 3ted Uno Upon eirtergence each »ast instar larva was removed from the • cageJje usinausing a smallSma ll moistened camel 's- hair brush. The dead host was removed fromfr« m then, cage, pinned and numbered for reference and specific identification.
^ach emerged larva was p,aced,ndividua,ly „ in a pupatfon! , vfa , to Observe pupation procedures. Those dryinid adults reared in ,*7 were -owed to pupate in one-dram glass vials filled one-half fun wit h white S nd Ati9 ' rubber stopper was used —— to sea, the mouth - ; the vials. Each vial * was numbered corresponding to the host. The •arvae were allowed to P upate and observations of this process were -de using a binocular dissecting microscope.
This method proved satisfactory for pupation of numerous of the '-- -ever, it did not al,ow for detection of pupal developments within the cocoon since th« l„,„. • ' arVae '""""Porated sand particles , nto , --pact, opa qu e cocoon. These cocoons were ,oose in the sand The Un,aa reared to aduIts ,„ , 968 _ ^ ^ ^ _ Cram ^ vials containing no sand ThSThe | a ruaa ' arVae were otherwise treated as in ••967. In the absence of sand t-h. t "" ' arVae ^"ructed on ly s „ ke „ cocoons wMch adhered to the sides and/or bctt™ of tneth, via „.„is. TheseT„ cocoons al- lowed for observations of Prenr--npupal,UDa ] nil „ , pupa l and re p -e,,ergcd adult activity ^e pupae were left undisturbedea until the<-h„ emergence of the adult drytnfd$. Upon emergence they were V -re transferredtr-n-f from the pupation vials 17
into cellulose nitrate cages. Each cage was numbered corresponding to
the host.
All the emerged adults were supplied with, and consumed, a sucrose- water solution. This solution was introduced twice daily into the cages
using a hypodermic syringe. A medium-sized drop of this solution was placed
directly on the sponge rubber floor of the cage and was usually consumed within one to two hours. That which was not consumed dried to hardness on the cage floor. Dryinid adults, especially females, were frequently observed to "lick" the hardened sugar spots. The sugar solution was
placed in the same area of the cage to minimize danger of the parasite
becoming immobilized in the sticky solution. This event did occur with
two male dryinids which became stuck by their wings. The wingless
females were not bothered.
After death, each adult was mounted and labeled with the host
number. The cocoon was removed from the pupation vial, placed in a
small gelatin capsule and pinned with the host. The sand-encrusted cocoons were easily removed from the vials. However, those cocoons which were
formed against the empty vial surface were difficult to remove without damage. A mounted minuten-nadeln was used to tease the edges of the cocoons from the glass surface.
Occasionally a larva formed its cocoon on the cork stopper or on
the foam rubber cage floor. In such cases the cage was left undisturbed until the adult dryinid emerged. The adult was removed and placed in a clean cage and the host was removed and pinned. The cocoon was then re- moved with that portion of the substrate upon which it was formed, placed
in a gelatin capsule and pinned with the host. 18
Preparation of Adult Dryinid? for Identification
The method of Richards (1339) for preparing dryinid adult heads and
chelae for specific identification was used. The time necessary for
adequate maceration of the structures depended on the size of the insect.
Two hours was adequate for the largest specimens without causing deteriora-
tion of membranous areas.
Dissections of mouthparts were made under a binocular dissecting micro-
scope at 60X. The heads, due to their small size and fragility, were dif-
ficult to secure on a glass slide surface. Therefore, a microscope dis-
section platform was made.
A deep-well ground glass slide was converted into a suitable dissecting
platform. The small end of a microvial cork was glued with the flat side
up in the center of the deep-well. The height of the cork was such that
when the deep-well was filled with glycerine the head atop the cork was
completely covered. The cork provided a suitable surface upon which
to immobilize the head. One mounted minuten nadeln through the foramen
secured the head to the cork surface while the mouthparts were teased from the
head with another mounted minuten nadeln.
The labium and maxilla were dissected either separately or
plete unit. The mouthparts and left prothoracic leg were placed in a
separate drop of glycerine on a clean glass slide and covered with a glass
coverslip. This temporary preparation was studied under 3 compound micro-
scope at 100X and 'tOOX. The mouthparts of all the identified specimens, unless obvious duplicates from the same collection, were dissected com- pletely from the head. 19
After studying the dissected mouthparts and chelae, these
structures were removed from the glass slide and stored in the manner
described by Gurney et a). ()36h) for insect genitalia storage.
RESULTS AND DISCUSSION
Kansas Collect ion Materi a 1
Table 1 summarizes the results of field collections made during
1967 and 1968, various collections from Hays, Kansas, miscellaneous
materials from museum collections at Kansas State University and from
Arnett's collection from Donaldson Pasture. These data are listed
alphabetically by genera and species.
The data indicate each single date on which genera and species
were collected and with what parasite they were infected. In addition,
the collection locality and the predominant grass(es) present in the
collection area are also listed.
Positive identification of the nymphal leafhoppers parasitized is
difficult. Nymphs and females were tentatively identified by com-
paring the specimens with identified males.
Parasites of 27 leafhopper genera were collected. Of the 27 genera,
18 contained specimens of St reps iptera , probably all belonging to the
family Hal ictophagidae (Table h) . Dryinid larvae were the most frequent
parasites occurring in r all 27 genera. Only amily identification is pos-
sible since no key to larval stages exists. Only the larvae of Dryinidae were found in five of the genera. 27 These are: Del tocephal , us Dl craneura , fmpoasca, Norvellina and Scaphytopius. Dorllaldae larvae (Pipuncu i idae . .
20
of authors) were found in 11 genera. Larval stages cannot be identified to
species because no keys are available.
Seven of the 2/ leafhopper genera contained specimens parasitized by
all three parasitic groups, possibly because more specimens of these
genera were collected. These are: Athy sanel l la, Flexamia, G rami ne la ,
Laevicephalus , Macrosteles , Mocuel 1 us , and Parabolocratus.
Of the 2'/ genera, 11 contained parasites of both Dryinidae and
Hal ictophagidae. These include: Aceratagal 1 ia , Balclutha, Dorycephalus,
Draecul ace phala, Driotura , £nd_rij, Mesamia , Paraphlepsius , Polyamia,
Sttre 1 lu s , and Xeroph 1 ea
Four of the 27 genera were parasitized by both Dryinidae and
Dorilaidae larvae. These include: Chloro tett ix, Exi t ian us, Gilletiella
and Psammo t et t i x
Streps iptera males were either emerged or unemerged. Unernerged speci-
mens were pupae or pre-emerged adults, still contained within the un-
opened pupal cases. Pre-emerged adult forms could be easily dissected
from the puparia after the leafhopper host had been slightly macerated
in cold potassium hydroxide solution. St reps iptera males noted as emerged
indicated those which had emerged from the puparia prior to collection of
the host. Empty puparia were considered evidence of parasitism.
Dryinid larvae were arbitrarily designated as in early or late stages of development. Early stages denoted those which were internal and undetect- able in uncleared hosts. The larval sacs, noted externally on the host, which had only one exuvium in addition to the larva were desionated as early
Hardy ( 1 S^3) indicated that the older larval stages of Dorilaidae are generally larger and darker than earlier stages in the host. Clearing, 21
however, removed much of the color and seemed to cause distortion of the
larvae in this study. Therefore, these larvae were not recorded as early
or late stages.
Also contained in Table 1 is a partial listing of the predominant
grasses noted in each collection area. This study did not include deter-
mination of food or oviposition hosts of any leafhoppers, except for those
from which adult dryinids were reared. Oman (19^9), in his treatment of
Nearctic leafhoppers, gave general categories of host plants utilized by
some of the leafhoppers in this study. These were primarily herbaceous
plant feeders. Added information on this subject was found in Metcalf
(1964)
Nymphal and adult stages of several of the leafhopper genera were
subject to parasitism (Table 1). Because nymphs cannot be identified
with certainty, some records probably were missed and many included
records were based on uncertain identification.
Dual parasitism, the existence of two different kinds of parasites
within the same host, not different stages of the pame parasite, was
noted in two genera. A Laev icephalus female was collected which con-
tained a strepsipteran female and a Dorilaidae larva. The strepsip-
teran female also contained several triungulin larvae. Also collected was a male Macrosteles fascifrons (Stal) which contained a late dryinid
larva in addition to a Dorilaidae ]arva. This male did not have any apparent modification of the external genitalia.
The effects of parasitism on the host's external genitalia have been indicated by several investigators. Ross and Moore (1957) discussed the effects of dryinid parasitism on the genitalia of the Empoasca fabae 22
complex. !n such specimens, the genital ic structures and the apodemes at
the base of the abdomen failed to develop completely and were often
entirely different from normal specimens. Raatikainen (1966) discussed
the effects of the streps! pteran parasite Elenchus tenuicornis (Kirby)
on the morphology of adult delphacids, Jove sel la pel lucida (F.). Bohart
(19^3) mentioned several references to such effects of streps ipteran in-
fections, primarily in hymenopteran hosts. Fenton (1918) mentioned
modifications of genital ic structures caused by dryinid infections. Such
modifications as a reduction in the size of the leafhopper ovipositor
and the reduction in the number of the male aedeagal processes were noted.
Oman (19^9) stated that, in spite of their large size, the larval Doriiaidae
did not seem to modify the host's genital ic structures. Douglas (1890)
stated that the action of the larvae on the genitalia of the host did pro-
duce alteration and atrophy of these structures and abortion of function.
Parker ( i So/) noted variations in the shape of the head and in the male
genitalia of Opsius stact ogalu s Fieber due to parasitism by a Doriiaidae.
Tomosvaryella frontata (Becker), in France. Blocker (1967) noted no
apparent genital ic modifications in species of Athysanel la which were para-
sitized by Doriiaidae larvae.
Males of several genera or species of leafhoppers collected in Kansas
grasslands had modified genital ic structures. These modifications probably
were due to the presence of the parasitic forms, although similar modifi-
cations may also occur in specimens containing no detectable parasite forms.
One male of Athysanel la sp. , Graminella mohri DeLong, and Laeviceph-
3 S and two L_. _ JiiL P-' males of sp. (probably parvulus (Gillette)) were x
23
noted to have aberrent genital ic structures. Each of these males con-
tained one Dorilaidae larva. One male Flexamia ref lexa (Osborn and Ball),
containing both an unemerqed male and a female St reps iptera , also had
altered external genitalia.
Location of Dryinid Larval Sacs jn Leaf hopper Hosts
Table 2 indicates the locations of dryinid larval sacs in the leaf-
hopper hosts collected during the 55 county grassland survey. The
number of included genera (16) is therefore less than that of Table 1.
Dryinid larvae were found in various locations in the host's thorax
and abdomen. Specimens of Dor . y LephaJ_us_ s p , Ex it i anus exitio sus (Uhler),
Flexamia sp. F. , and abbreviata (Osborn and Ball) had larval sacs be-
tween different thoracic segments, as well as between abdominal seg-
ments. The other genera in this study had larval sacs between abdominal
segments.
The orientation of the larval body in the host did seem to be more
consistent. Most of the larvae were situated with the head and posterior
abdomen (due to the larva's U-shaped body) inserted into the venter of
the host's thorax or abdomen. The larval body and any adhering exuviae
adhered to the lateral surface of the abdomen. A few larvae were in-
serted dorsal ly rather than ventral ly.
D ce J - -?_Q'. 5 S nymphs had the larval sacs P.b^li . , P- located on the venter of the thorax. Exltianus exitiosus (Uhler) and nexamj3_ sp. had the
larval sacs located on the ventrolateral surface of the thorax. 2k
Although there was usually one dry in id larva per host, exceptions to
this condition were frequent. Males and females of Athysa nella arge nteola
(Uhler) contained one or two larval sacs per host. Usually there was a
size difference between the two larvae. The earlier nymphal stages of this
species contained only one larva. A similar condition existed in
Kocuellus collinus (Boheman) , which had females containing one or two
larval sacs. Females of Parabolocratus sp, are larger and frequently
contained up to three larval sacs. Difference in size was noted in exter-
nally visible sacs; internal early larval stages did not always show a
marked size difference. It is possible that, due to its larger size,
more than one larval parasite per host is normal. Fenton (1318) states
that when there is more than one larva per host, the last one deposited
would probably not reach maturity due to a competition for food. Mul-
tiple parasitism seemed to be a frequent condition in specimens of
Macrostftles fasc if rons (Stal) parasitized by the dryinid Pachygona topus
minimus Fenton. Only one parasite survived to maturity in these cases
(Barrett et al., I365). However, this may not be the case with dryinid
parasites of Parabolocratus sp. The larval parasite stages in this host
regularly occurred singly between consecutive abdominal tergites.
Most of the larval sacs were brown i sh-grey to black. Mo attempt
was made to draw conclusions as to the possible number of parasite
species per host based on color difference of the larval sacs. However,
EmpjDasca sp. did have a very distinctive sac. They were long, often
extending from the point of extrusion to the tip of the host abdomen. . s
25
The sacs were located under the wings on either side of the host. Their
most striking feature was their light to deep green color, often the same
shade of green as the host.
Frequently the adult host contained dry in id larvae positioned beneath
the wings. This often caused a prominent displacement of the wing.
Position of Strepsiptera Parasites in the ir Hos t
Table 3 indicates the positions of Strepsiptera parasites in their
hosts. These parasites seemed more restricted to the host's posterior
abdominal segments than the dryinid larval sacs (Table 2).
A persistence in location of the sexes of these streps ipteran para-
sites was noted in the hosts. Females were found on the venter of the
host abdomen. Male puparia were usually noted extruded between dorsal
segments on the host's abdomen; only occasionally were they extruded
ventral ly
instances of more than one life form of strepsipteran parasite were
noted. These were noted in both male and female hosts, whereas nymphs
normally contained only one such life form. Several combinations of life
forms were noted. Two female parasites, either on the same or opposite
sides, were often noted in the same host. A female and male (emerged and/or
unemerged) sometimes occurred within the same host. Also two male para-
sites (emerged and/or unemerged) occurred in the same host.
Summary of I dent i fj ed_S trepsi ptera from Kansas Hosts
Table k lists the identified Hal ictophagus C-jrtis species which were collected from Kansas hosts. Hal ictophagus and its included species, i
26
recorded from the homopterous families Glcadel 1 idae, Membracidae,
Fulgoridae and Cercopidae, have been discussed by Bohart (1941, 1943, 1962) in
his studies of the North American fauna. A list of hosts for these para-
s ites is included in these works. No records of Hal ictophagidae were re-
corded from Kansas in his earlier work (19^1); however, he did record
arcericanus 1 H' Perkins f rom Aceratagal ?a he lveola Oman in Garnett, Kansas
(Bohart, 1943).
Six species of Hal cto phagus were represented in this collection.
H. acutus Bohart was represented by a female parasite in Draeculacephala
mo! 1 ipes (Say). _H. bidentatus Bohart was found parasitizing four leaf-
Hopper genera: F i 1 exam a , GramineUa , Athysanel la and Parabolocratus .
Six leafhopper genera were parasitized by H_. mack ay? (Bohart).
11- oman I Bohart 'was identified from an unidentified species of Acerata-
Si-LUiL- A male specimen of Parap hle psius irroratus (Say) was parasitized
by a female of H. u hler i (pierce). HL insularum (Pierce) was identified
from a male Dorycephalus platy rhynchus Osborn.
A possible new species of Hal ictophagu_s_ was recorded from Stirellus
bicolor (Van Duzee) . More specimens of this form need to be collected
in order to verify this.
The genus Flexamia contained species which were parasitized by either
bidentatus II- Bohart or HL mackay i (Bohart). The majority of the col-
lected Flexamia species contained the latter.
?-. M. Bohart (personal communication) indicated that the host species and the locality records for these Hal lci:ophag us species are new
Kansas records. )
27
Notes on a Male Dry in id (tr i be : fiona topod ? n i in its Late Larval to Adult Development
The stages in development of this specimen were similar to the stages
in those specimens indicated by asterisks in Table 7(b). The length of
time per stage varied among these specimens. The stages in development
were observed in the cocoons formed In the empty pupation vials. This
specimen pupated in the cellulose nitrate cage.
A male ("DP-1 03") was collected as a late larva in a male Balclutha
negl ecta (DeLong and Davidson) on 6 June 1 S68 . The host was caged on
buffalograss.
The larva emerged from the dead host on the morning of 8 June. It was
orange-colored and had a light yellow area behind the head region. The
larva was initially crcam-cclcred and turned orange as it finished con-
suming the liquified body contents of the host. This process began toward
the rear of the host and proceeded anteriorly. The larva worked its
anterior body segments into the host's body cavity as the meal continued.
The mature larva crawled about on its dorsum on the cage floor.
Microscopic observations revealed that the larva possessed what appeared
to be several retractable protuberances on the lateral, dorsal and ventral
body aspects. Each protuberance seemed to possess one short spine.
The larva moved about by use of peristal ic body movements. The un- dulating movements proceeded primarily posterior to anterior, although
the reverse was noted. The he-^d was partially retractable into the anterior larval body segments. Light prodding with a mounted minuten nadeln resulted in such retraction. .
28
The larva emerged about 8:30 AM and began spinning its initial
cocoon on the side of the cage at 5:15 AM. The outer, loose cocoon was
spun first. The larva stopped on the cage side and began depositing
strands of a silken material. Deposition of this material from side to
side and anterior to posterior provided a loose, imperfect covering over
the larva. It appeared that each strand was laid down separately; each
end was attached to the cage side.
The very active larva moved about within the confines of this imperfect
cocoon and reinforced it with additional silken material. The deposition
of material was done with the larva on its venter. The deposition of silk
strands from anterior to posterior points along the length of the cocoon
necessitated the larva's bending upon itself in a U-shape.
Very little silk was laid down on the cage side which formed the
"floor" of the cocoon. It was through this "floor" that microscope obser- vations were made.
Silk deposition continued for about three hours. At this time the outer cocoon was practically completed and had almost a papery-slick
texture when viewed from above.
The larva then became scill for about kS minutes. Only occasionally did it move about in the cocoon as if to repair weak spots with added
silk deposit ion
The larva then began to deposit another layer of silk inside the outer cocoon. A similar manner of deposition was followed as before, except that the silken strands were attached to the cage side much closer to the larva. This resulted in the inner, tight-fitting cocoon in which pupation occurred. This Inner cocoon more or less fit the contours of the extended larva. 29
An interesting point was observed in these procedures. The larva
was observed to deposit increased amounts of silk at a point inside the
inner cocoon. This material was so positioned as to occur at the junc-
tion of the thorax and abdomen of the later-developed pupa and pre-
emerged adult. This same occurrence was noted in other observed larvae.
This "safety belt" was possibly used later to aid the adult in emerging
from the cocoon.
After the completion of the inner cocoon (in about four hours) the larva
again became quite still, only occasionally moving in the inner cocoon. It
contracted itself to about one-half of its original length at the end of
the cocoon. All points of the body, except the head, were in close con-
tact with the cocoon sides.' This position was maintained for up to five
minutes; then the larva extended itself to add silk to the cocoon. This
done, it retreated as before often to the opposite end of the cocoon.
This procedure was repeated for about four hours, at the end of which time the larva moved to one end of the cocoon and again contracted
to about one-half its original body length. During the stage of con-
traction only slight head movements were noted.
At this point the rough dimensions of the inner and outer cocoons
were: 3 mm x 1 mm (inner cocoon) and 6 mm x 3 mm (outer cocoon).
Only slight larval activity was noted for the next kS hours. On
10 June the larva was observed extended to about three-fourths the length of the inner cocoon. It did not move except when disturbed by a bright microscope light. The heat of the light could have caused the movement. 30
By 8:30 AM, 11 June, the prepupal stage was complete and the larva
was extended the entire length of the inner cocoon. No body movement
wais noted. However, a constriction was observed to be forming at what
wou Id be the juncture of the thorax and abdomen of the pupal stage.
At 9:00 AM, 12 June, further changes were noted in the prepupal
stage. A dark area appeared in the center of the abdomen and leg parts
appeared to be forming. Also noted was a darkening in the "head" region
posterior to the larval mouthparts. This darkened area later proved to be
the ocular area of the adult.
Pupal formation was completed on 13 June after the last larval skin was
shed and pushed to the posterior end of the inner cocoon. The larval mandi-
bles were seen in the shed skin. At this time the pupal antennae appeared,
to be. well formed, as were the legs and mouthpart palpi. These structures
were cream-colored, not dark. Also noted was a much enlarged and darker
ocular area than was observed on 12 June.
A gradual darkening of the pupa from a yellowish color to black and
an enlarged and darkened ocular of the formed head were noted externally
In the pupal stage between 13 and 17 June.
The first movements of pre-emerged adult appendages were noted on
17 June. The male emerged on the same date through a round openinq cut
at the anterior end of the cocoon, where the inner and outer cocoons
were closely appressed. The male seemed to push with its body and
legs against the "safety belt" during its emergence.
At this point the male was allowed to reach a stage of active flight,
it was then transferred to a clean cellulose nitrate cage and fed sugar- water solution. !t died on 22 June 1968. 31
Rearing of Dryinids from Exitianus exitiosus (Uhler) from Bermudagrass
Three nymphs and two females of Exitianus exitiosus (Uhler) were
collected from bermudagrass in Manhattan, Kansas during July and August
of 1368. These specimens contained visible larval dry i n id parasites.
These larvae were reared to adults: Neogon atopus (two specimens) and
Gonatopus (three specimens). The larval sacs of Neogonatopus were located
-J between abdominal segments 3 *; the Gonatopus sacs between segments k~S.
Table 5 indicates the lengths of larval, prepupal and pupal stages for each
specimen reared. The length of adult life was also recorded.
Two of the larval dryinids formed their cococns on the surface of a
bermudagrass blade. The other three specimens pupated on the foam-rubber
cage floor.
Parasitised nymphs did not molt from the initial caging until the death of the host resulting from the emergence of the mature larva. Caged
leaf hopper specimens fed readily on the bermudagrass supplied. Adult dryinids fed on the sugar-water solution supplied.
Drtyinids from Col 1 actions and Rearing Studies
Table 6 lists the genera of. dryinid females collected from grassland areas in Kansas. These genera were tentatively identified using keys to female forms (Perkins, 1905; Richards, 1933)- No males were recorded from the field collections.
Specific identifications were not made for these specimens even though kesys to certain species exist (Perkins, 1305; Kieffer, 1914; Fenton, 1918;
Richards, 1939). Since many of the identified species listed in the liter- ature are based on unique specimens, it is not certain just how many of .
32
these species will be destined to synonymy if a detailed study of all
available material is made. Changes in the color patterns of dryinids
due to age would seem to compound the problem (Perkins, 1306).
Seven genera of dryinid females were identified (Table 6). The majority of the females belonged to Gonatopus and Pseudogonatopus . Accord- ing to Barrett et al (1965). . Pachyfronatopus is possibly congeneric with
frtgonatopus . If this is the case, the number of genera recorded would be reduced to six.
The majority of females were collected during August and September 1967. This gives no real indication of their occurrence in the field because this heavy collecting resulted from taking the state survey. Each area would need to be collected through several seasons to get a valid idea of annual field populations.
Table 7 (a 6 b) summarizes the life history data on dryinid females and males reared from leafhopper hosts collected in Donaldson Pasture. Hosts were collected primarily from buffalograss but some collections from bluestem and switchgrass were also made.
Both female and male specimens reared from Donaldson Pasture as material, well as those reared from the Hays, Kansas site to be dis- cussed later, belong to one tribe of dryinids, i.e., Gonatopod in ? Richards (1939), Fenton (, l8) and 9 Perkins (l 9 12) discuss the characteris- tics of this tribe and give keys to the other tribes in this family. The 133 individual leafhopper hosts containing dryinid larval sacs in various stages of development were collected and caged. Two larvae ("DP-96", . W -,12") were recovered from thfi transpopt ^^^ ^ matur ^ emeroed 3- ,- Isrva no "°^hn«<-cts w-,--. r -„ j _i r • "' Wcr,i 'horded for these specimens. 33
Due to the techniques used during 13o7, the length of time (days)
which the dryinid specimens spent in various stages within the cocoon
could not be ascertained. Therefore, the table notation "cocooned
specimen" refers to the length of time between the larva spinning the
cocoon and its ultimate emergence as an adult. The refinement in method
used in 1 968 did allow for observations of the various intra-cocoon
stages.
This group of leafhoppers yielded 37 adult dryinids: 33 females and
h males. Seven of the hosts yielded mature larvae which, after forming
norma! -appearing cocoons, did not pupate. These larvae shriveled and
occupied one end of the cocoons. Three hosts yielded larvae which died
before forming cocoons; one larva attempted to form a cocoon but failed
to complete it before dying.
Perkins (1505) discussed factors which might cause the shriveled
condition in dryinid larvae and a method by which partial recovery of
these larvae might be achieved. He suggested placing the cocoons in
cooler and damper conditions, but this did not aid in the recovery of
the seven larvae mentioned above.
The majority of the material from which parasites were reared were
nymphs. Of 30 nymphs, possibly Athysanelia, 25 were female Gon_atopus,
three female Epfgonatopus and two male Gonatopod ini .
One. nymph of Parabolocratus yielded a female Chalcogonatopus . A female leafhopper, possibly A thysanelia texana (Osborn) , yielded 3 female
Ep'gonatopus. Another °.f female of this same genus yielded a maie of the
tribe Gonatopod In! . . -.
34
A female Gonatopus was recovered from a male Flexamla prairiana
DeLong. This host also contained a lata larval dry in id which was com-
pletely internal and was not noted until the host was cleared. A male of
Endria i nimica (Say) yielded a female Gonatopus.
One male dryinid, tribe Gonatopodini , was reared from a male
Balclutha neglecta (DeLong and Davidson). An Aceratagall ia_ nymph yielded
a female of Pachygonatopus
The two emerged larvae found in the transport cartons were females
of Gonatopus and Meogonatopus.
Life history data for dryinid adults reared from leafhopper hosts in
a western wheatgrass pasture in Hays, Kansas are given in Table 8 (a & b)
During the summer of 1967, 55 hosts were caged. From these, 12 larvae were
recovered and reared to adults. Five other larvae emerged, formed normal
appearing cocoons but did not pupate. Two genera were represented in
this leafhopper material. Each host possessed one larval sac, with the
exception of one Athysanel la nymph which had two sacs. Only one larva
emerged from this specimen. Ten Athysanel la nymphs yielded ten specimens
of female Go natopus . An unidentified nymph yielded a male dryinid. One
female £x?tianus exitlosus (Uhler) was collected from which a female
-i^iL^R^ was re-a r ed.
parasitic Wqrms , Noted in Kansas Leafhoppers
Hair worms (Phylum Nematomorpha) , or Gordiacea, are cited by Meglitsch
as (1967) parasitizing certain arthropods. Immature forms of the order Gordioidea, in particular, parasitize insects. The adults are free-living and aquatic. Severin (1924) reported such a form from the body of a sugarcane leafhopper (Del phacidae) , Perkmsiella sacctiartcida Klrkaldy 3 5
from the Hawaiian Islands. Apparently such findings of these worms in
leaf hoppers are uncommon.
Three leafhopper specimens were collected during this study, each
containing one unidentified pale-yellow worm. fJo positive identifica-
tion of the worm has been made. The probability exists that these
worms belong to the above group.
Two nyrnphal stages of Mocuel lus coll inus (Boheman) were collected
from western wheatgrass pasture at Hays, Kansas on 8 September 1967. In
one nymph the worm's body was partially extruded between abdominal seg-
ments $-k. The other worm was partially extruded between abdominal seg-
ments 7-8 in the host.
One female _BaJcJ_u_th_a_ nejjjicta_ (DeLong and Davidson) collected
July 3 1368 contained a similar worm form. This worm was partially
extruded between abdominal segments 6-7.
SUMMARY AND CONCLUSIONS
Parasites of leaf hoppers were studied from 36 sweep net collections
from tail grass and mixed prairie grasslands, in 55 Kansas counties in
late summer of I367 and other collections in 1268. Material from 1957-
Ip66 was also included. Adult parasites of Halictophagidae (Strepsiptera) and Dryinidae (Hymenopteraj were identified from both collected and reared material. Representative specimens of leafhoppers with their associated parasites are deposited in the insect collection of Kansas State University,
Manhattan, Kansas.
Specimens from leafhopper 27 genera were collected with either larval or adult parasites of Diptera Dorilaidae (Diptera), Dryinidae (Hymenoptera) 36
and/or Hal ictophagfdae (Streps iptera) . Specimens from 18 genera contained
immature and/or adult Ha lictophog idae. All 27 genera contained dryinid larvae. Dorilaidae larvae were found in 11 leafhopper genera.
Seven of the genera were parasitized by species belonging to all three
families. Parasites of both the families Dryinidae and Hal ictophagidae
were found in 11 of the genera. Five genera contained only dryinid larvae. Four genera were parasitized by both Dryinidae and Dorilaidae.
Hal ictophagidae (Streps iptera) were found in the hosts as triungulin
larvae, males (emerged or unemerged) and/or females. Other strepsipteran larval stages were not observed. Dryinidae and Dorilaidae occurred as larvae within the hests.
Leafhopper nymphs and adults were parasitized by the above forms. Dual parasitism was found in L aevicephal us sp. (with Hal ictophagidae and Dorilaidae) and MacrosteTes fascifrons (Stal) (with Dryinidae and
Dori la idae) .
Modifications of external genitalia of the hosts, possibly due to parasitism, were found. Aberrant genitalia were found in males of
Aphysanella Graminella and , Laevicephalus which contained Dorilaidae
larvae, as well as in males of Flexamla reflexa, (Osborn and Ball) con- taining Hal ictophagidae.
The dryinid larvae in 1 genera 5 of leafhoppers were most often found
in the abdomen. These did not occur with regularity between particular segments, except for Empoasca^sp. in which the larvae occurred regularly between abdominal segments 3"4. Specimens of Doj^cjspfc^ sp. and Endria_ irmnica (Say) had larvae between the mesothorax and metathorax. Larvae were found between the prothorax and mesothorax of ExIHanus. exftlosus (UhJer) 37
and Flexgflita ^>brev?ata (Osborn and Ball). Usually each host had one larval parasite, although more than one were noted in Athysanella
a rgenteola (Uhler), Mocuellus collier- (Boheman) and Parabo locratus sp. Strepsipteran parasites were localized posteriorly in the host's
abdomen. The females appeared to locate i^ore ventrad in the hosts;
the males more dorsad. Both sexes were found in the same host.
Six species of HaHctophaqus turtis (Streps iptera) were found in a total of )k genera of leaf hoppers . One possible new species was found in StireHus bjcolor (Van Duzee) . The . host and locality records for these
parasites are all new for Kansas.
Parasitic worms of the order Gordioidea were found in two nymphs
of Mo^eJJ^ cpJUnus (Boheman) and one female of Balclutha neglects
(DeLong and Davidson). These worms were partially extruded from the
host 's abdomen.
Developmental history of one male dryinid was recorded as observed through an exposed portion of the cocoon. The length of larval, pre P u P al, pupal and adult life stages was recorded. Larval behavior, cocoon forma-
tion and activities within the cocoon were recorded. Similar information
was recorded for all the dryinids reared from leafhopper hosts.
Various dryinid genera were reared from larval stages, the hosts
being kept alive on grass in cellulose nitrate cages until the larva emerged. Pupation occurred in one-dram glass vials. Adults reared and those collected from the grasslands provided the identifiable material in this
StUdy M° St f the ' ° hosts contained only one larva. If more were present only one emerged and developed. 33
Seven genera of drylnlds were collected as females from grasslands with no host information. The majority of specimens belonged to the genera Gonatopu s and fseudogonatoput. Numerous hosts collected in Donaldson Pasture near Manhattan, Kansas, yielded 37 adult dryinlds representing five genera. The majority of these were of the genus Gonatopu* from Athysanella nymphs. Gonatopus was the only genus reared from a small number of Athysanella nymphs and one focitfanus. ejcftlosus (Uhler) female from western wheatgrass from Hays, Kansas. 39
ACKNOWLEDGMENTS
The author acknowledges Dr. H. Derrick Blocker for his guidance,
suggestions and encouragement during the course of this study, and for identifying leafhopper hosts, and Dr. R. J. Elzinga and Dr. L. C.
HuJbert for their review and criticism of the manuscript. The author gratefully acknowledges Dr. R. M. Bohart for identifying the strep-
sipteran parasites.
The author is grateful to Dr. Thomas L. Harvey for supplying
leafhopper material from the Fort Hays Branch Experiment Station,
Hays, Kansas. The help of Mr. Donley H. Johnson in identification
of leafhopper hosts and collection of material is hereby acknowledged, as is that of Mr. George Ameel in preparation of material. The author acknowledges Dr. M. W. Nielson for the use of original rearing cages as patterns for the ones used in this study.
» Dr. Herbert Knutson, Head, Department of Entomology, Kansas State University, is acknowledged for his initiative and interest in support of this research.
The author gratefully acknowledges the patience and encourage- ment of his wife, Cris, and her criticisms of the manuscript. Acknowledgment is also given to Mrs. Helyn Marshall for typing the manuscript. J»o
LITERATURE CITFD
AinS C ]S2 N° te ° n ^EOatOEus ombrodes . a parasite of .assids]i^fd ; to(Hymen.,°-uHomop.).? Entomoj . News 31:169-173, I87-I9O.
R"^° n5e f Acrid!d ^pr^Mc"; \?°' ° W'-t'on. to rangeland r 9e S ' teS ' UnpUbUshed Ph ' D ' "-rftlSn. Kansas sSte Un?vers!ty?
Barrett C. F P. H Westdahl and H. P. Richardson. 1965. Biology of Fent (Hymenopte^Dryinidae) ^SS^r °: a pj J ? I o
8Oha M - eViSi ° n f tHe ° *'1»'*™- CI If. Pub. Ento:o | 7 (6^i-,60: *•*
• New species of I9J3. Hal ictophagus with a key to the s i ' "^ "i^»*,«-) c BnLo, rcM 3 Efts
2, 9 "ew strepsfpteran parasite en T str»n -U3 ii ? ? „ Coreidae """P'^refd..). Entomol. Soc. Wash., Proc. et(2Us\ 3 l
Blocker H. D. 1?67 . Athysanella attenuata and a cicely related new
COOk C 1 1 - ^ ^ '"«»PPir- U.S.Dep.Agrf. Par's. Bull. l386. -,l f
H ] 6 -' T m and - lL'« Genera of North American Dipter* '"^nd-nd eded. "hH. Tripp,T P ' ^Woodhaven,^ N. Y. TlTpT ^
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APPENDIX . * N — ' - ^ —.
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CL -C u- TO cr in TO «**i r» cm -a- -3- ; co <-r\ > O r-~- r>- la vo O - si !! TO 'c 1> L. in '.-0 in 4-1 TO TO H> -C 3 >- •- in .—v i. O Q> TO 1) 4-1 1- 10 o in O TO 1/1 01 l/l in ^ 4-1 CJ < in >n n TO C !_ < < < < < • < < < TJ TO TO in — ~ >» 1- TO O 3T *-> SI m <=! — (U j: l. -~" 0) a. '-+- in o — i(J ~! O. -O E CO AJ O < tl en in _D CM LA O CM CM O ! CM E I 1 -Q i i s 3: TO 3 3 3 3 3 3 3 3 PARASITES IN LEAFHOPPERS FROM KANSAS GRASSLANDS (HGKOPTERA :C ICADELL I DAE) by ROBERT SMOOT BALDP.iDGE B. S., Baylor University, I966 AN ABSTRACT OF A MASTER'S THESIS submitted in partial fulfillment of the requirements for the degree MASTER OF SCIENCE Department of Entomology KANSAS STATE UNIVERSITY Manhattan, Kansas 1S69 . Parasites of leafhoppers were studied from 96 sweep net collections from tall grass and mixed prairie grasslands in 55 Kansas counties in late summer of 1 S 6 7 and in ,*8, and fro. miscellaneous collections made from 1357-1366. Adult parasites of the families Hal ictophagidae (Streps iotera) and Dryinidae (Hymenoptera) were identified from collected and reared material . Mature forms of Drying, Hal ictophagidae and Dorilaidaa (Diptera) (fipunculidae of authors) were collectively recorded fron 27 leafhopper genera la six subfamilies: Del tocepha I ,„„ 20 ( ) , Udrinae (I), Agallinae (0, Hecalinae (I), Dorycepha. inae (2) and Typhlocybinae (2). Specirnens fro. ,8 genera contained i ramature and/or adult Hal ictophagidae. All 27 genera contained urymiadrvinid larvap rv,-M~tj . larvae. Donlaidae larvae were found in 11 leaf- hopper genera. Seven leafho er PP genera were parasitized by species belonging to all three families. Parasites of both Dryinidae and Hal ictophagidae were found in of 11 the host genera. F ive leafhopper genera "contained only drvinid larvae. Four genera were parasitized by both Dryinidae and Dorilaidae larvae. Hal ictophagidae were found in the hosts as trfungulin larvae, males (emerged or unemerged) and/or females. Other larval instars were not observed. Dryinidae and Dorilaidae occurred as larvae within the hosts. Both leafhopper nymphs and adults were parasitized by the above forms • Dual wiciticni i.,-,* c^. < , parasitism was found in laevlcephalus sp. (with Hallcto- Pnagidae and uorilaidae) and Hacrostejes fascjfron, (SU„ (with Dryinidae and Dori laidae) . , Modifications of externa! genitalia of the hosts, possibly due to parasitism, were found infrequently. Aberrant genitalia were found in the males of Athysanella , Graminella and Laevicephalus, all containinq Dorilaidae larvae, as well as in males of Fl exam la reflexa (Osborn and Ball) containing Hal ictophagidae The dryinid larvae in 15 genera of leafhoppers were most often found in the abdomen. These did not occur with regularity between particular segments, except for Empoasca sp. in which the larvae occurred regularly be- tween abdominal segments 3-k. Specimens of D orycephalus sp. and Endria nlmIca (Sa V^ had dryinid larvae . between the mesothorax and metathorax, f. while dryinid larvae in Exitianus exitiosus (Uhler) and Fjexamla jbbrevtata (Osborn and Ball) occurred between the prothorax and meso- thorax. Usually each leafhopper host had one larval parasite, although more than one were noted in Athysanella argentepla (Uhler), Mocuellus doll Inus (Boheman) and Parabo locratus sp. Hal ictophagidae parasites were localized posteriorly in the leaf- hopper's abdomen. The females appeared to locate more ventrad in the host; the males more dorsad. Frequently both sexes were found In the same host. Six species of Halictophagus Curtis (Hal ictophagidae) were found in l|l genera of leafhoppers. One possible new species was found in Stirellus bjcolor (Van Duzee). The host and locality records for these parasites are new for Kansas. Developmental history of one male dryinid was recorded as observed through an exposed portion of its cocoon. The length of larval, prepupal pupal and adult life'forms was recorded. Larval behavior, cocoon formation and activities within the cocoon were recorded. Similar information was recorded for all the dryinids reared from leafhopoer hosts. Various dryinid genera were reared from larval stages, the hosts being kept alive on grass in cellulose nitrate rearing cages until the larva emerged. Pupation occurred in one-dram glass vials. Host hosts contained only one larva. If mor e were present only one emerged and developed. The majority of dryinids reared were of the genus Gonatopu, from Athysanella nymphs. Seven genera of dryinids were collected as females from grasslands with no host information. The majority of thiese specimens belonged to the genera Gonatopus and Pseudogonatopus. Parasitic worms of the order Gordioidea were found in nymphs of ^OCueUus COJUnus (Boheman) and one female of Balclutha peqlecta (DeLong and Davidson). These worms were partially extruded from the host 's abdomen. 4-1 > -w | . f<) • 4J Q. CD C CD 1 'o -Sept. -Sept. O. CO CD C 4-> C a CO Q. s: -Aug. 3 3 3 !*"» CD CM 3 3 Q. o co -> | •* i -3- i to OO .J- i oo CO CM o --a- o oo ro I CM CM CO CM CM CM CM •J" 2, 00 CM CM Csl CM CO CM 2