Masthead Logo

Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations

1968 Distribution and composition of populations of symbiotic on necrophilous Harold Allison Borchers Iowa State University

Follow this and additional works at: https://lib.dr.iastate.edu/rtd Part of the Entomology Commons

Recommended Citation Borchers, Harold Allison, "Distribution and composition of populations of mites symbiotic on necrophilous beetles " (1968). Retrospective Theses and Dissertations. 3532. https://lib.dr.iastate.edu/rtd/3532

This Dissertation is brought to you for free and open access by the Iowa State University Capstones, Theses and Dissertations at Iowa State University Digital Repository. It has been accepted for inclusion in Retrospective Theses and Dissertations by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. This dissertation has been microfilmed exactly as received 69-9843

BORCHERS, Harold Allison, 1935- DISTRIBUTION AND COMPOSITION OF POPULA­ TIONS OF MITES SYMBIOTIC ON NECROPHILOUS BEETLES.

Iowa State University, Ph.D., 1968 Entomology

University Microfilms, Inc., Ann Arbor. Michigan DISTRIBUTION AND COMPOSITION OF POPULATIONS OF MITES

SYMBIOTIC ON NECROPHILOUS BEETLES

by

Harold Allison Borchers

A Dissertation Submitted to the

Graduate Faculty in Partial Sulfillment of

The Requirements for the Degree of

DOCTOR OF PHILOSOPHY

Major Subject; Entomology

Approved:

Signature was redacted for privacy.

In Charge of Major Work

Signature was redacted for privacy.

Chairman of Major Department

Signature was redacted for privacy.

Iowa State University Ames, Iowa

1968 ii

TABLE OF CONTENTS Page

INTRODUCTION 1

MATERIALS AND METHODS 5

RESULTS 12

Host Beetles and the Morphological Distribution of Their Symbionts 12

Carabidae 12

Harpalus pennsylvanicus DeGeer 12

Anisodactylus (Anisodactylus) agricola Say 12

Histeridae 12

Saprinus assimilis Paykull 12

Saprinus lugens Erichson 13

Margarinotus hudsonicus Casey 13

Hister abbreviatus Fabricius 13

Hister circinans Casey 13

Silphidae 13

Silpha noveboracensis Forster 14

Silpha americana Linne 14

Silpha surinamensis Fabricius 17

Silpha lapponica Herbst 17

Nicrophorus tomentosus Weber 18

Nicrophorus orbicollis Say 19

Nicrophorus pustulatus Herschel 22

Prionochaeta opaca (Say) 23 iii

TABLE OF CONTENTS (Continued) Page

Staphylinidae 23

Staphylinus maxillosus Linne 23

Staphylinus maculosus Gravenhorst 25

Philonthus politus Linne 25

Philonthus validus Casey 25

Philonthus cyanipennis (Fabricius) 26

Ontholestes cingulatus (Gravenhorst) 26

Trogidae 26

Trox unistriatus Beauvois 26

Trox suberosus- Fabricius 26

Geotrupidae 26

Geotrupes semiopacus Jekel 27

Geotrupes splendidus (Fabricius) 27

Scarabaeidae 27

Onthophagus hectate Panzer 28

Onthophagus janus Panzer 28

Copris tullius Olivier 28

Dermestidae 28

Dermestes vulpinus Fabricius 28

Dermestes caninus Germar 28

Nitidulidae 28

Stelidota geminata (Say) 28

Glischrochilus fasciatus (Oliver) 28 iv

TABLE OF CONTENTS (Continued) Page

Mite Symbionts and Their Occurrence on Host Beetles 29

DISCUSSION 37

CONCLUSIONS 56

LITERATURE CITED 58

ACKNOWLEDGMENTS 63

APPENDIX A: PHOTOGRAPHS OF MORPHOLOGICAL REGIONS 64

APPENDIX B: PHOTOMICROGRAPHS OF MITES NOT IDENTIFIED TO 77

APPENDIX C: ABBREVIATIONS USED 90

APPENDIX D: REGIONAL DISTRIBUTION OF MITE SYMBIONTS ON HOST BEETLES 91

APPENDIX E: FIELD COLLECTION DATA 120 1

INTRODUCTION

Symbiotic relationships between mites and involve a variety

of associations including , parasitism, and prédation. Per­

haps the most common association is phoretic commensalism where ambula­

tory mites may feed on integumentary exudates or bits of food material

clinging to the surface of the host. Furthermore, ambulatory

mites which feed on debris on insect bodies may also feed on the same

substrate as the insect host, thus utilizing the insect as a source of

transportation from one food supply to another.

A highly specialized form of phoresy involves a morphologically

unique form of mite deutonymph called the hypopus. These hypopi are

found in the suborder Astigmata (families Acaridae, Saproglyphidae,

Glycyphagidae, Labidophoridae and Anoetidae). Feeding does not take

place in that the mouth parts are much reduced. In addition the ventral,

posterior part of the opisthosoma is equipped with a battery of sucker

discs or with a pair of clasping plates enabling the hypopus to adhere to

the host. In many species setae on the legs of hypopi are modified into

suckers and adhesive type organs. The hypopus is apparently formed under

unfavorable conditions and does not moult until suitable environmental

conditions are present for tritonymph formation. Presumably when insects

carrying hypopi encounter favorable conditions of temperature and

humidity the hypopi moult to the non-phoretic tritonymph stage.

In addition to providing a means of finding favorable conditions for further development, this type of adaptation provides an excellent 2

mechanism for dispersal.

The adults and other trophic forms of hypopus-forming mites feed mainly on decaying vegetation, stored grain, and in forest litter and soil. Insects commonly occurring in these often accumulate infestations of hypopi and transport them until the tritonymph stage is formed, often in an environment similar to the one in which the hypopi were formed.

Deutonymphs of some Uropodidae (suborder ) attach to insects by means of a quick-hardening substance secreted by anal glands.

The result is a strong pedicel securing the mite to the surface of the insect. These pedicellate deutonymphs, like the astigmatid hypopi, lack feeding apparatus and are transported wherever the insect travels, or until a moult takes place.

Most of the studies involving acarine-insect symbioses have been concerned with Hymenoptera and Odonata. Very little has been done in analyzing acarine symbioses of other insect orders; however, a tremendous amount of literature exists listing mites and their insect hosts.

Linn/ (1736, 1746, 1758) reported the mite Acarus insectorum coleoptratorum from scarabaeid beetles and Acarus muscarum from house dies. He included all the Acarina in one , Acarus, and often mites took the specific epithet of the host, e.g. Acarus muscarum from museid . Pabricius (1805) reported taking what Linnet apparently had named Acarus coleoptratorum from a scarabaeid beetle. Ecological 3

criteria continued to be used as a basis for classification until the

late 1800's. Schaupp(1881) referred to louse-like parasites on

Nicrophorus tomentosus; undoubtedly he was referring to one of the com­

mensal mites common to many necrophilous beetles. Williams (1928), in his study of the hydrophilid affinis, reported observing red mites under the elytra of the beetles.

The literature is replete with listings of mites taken from various insects and other , but only a few workers have recorded enough data for one to even conjecture as to distributional patterns of mites on insect bodies.

European workers beginning in the late 1800's were involved primarily with descriptive and nomenclatural problems, as exemplified by Canestrini

(1882), Gudemans (1903), Michael (1901, 1903) and Leonardi (1900). Most workers failed to indicate the stage of development of the mite symbiont collected, on what part of the host's body the mite was located, and how many mites occupied the site. In light of the forenamed types of com­ mensal ism, these criteria are indispensible in a symbiotic study.

The order Coleoptera is among the least studied orders of insects involving mite symbioses. It has the largest number of described species, and its members have a biological range from phytophagy to parasitism.

Many of the larger beetles (, , ,

Carabidae, Staphylinidae, , ) frequently have an acarine fauna of symbionts when collected In routine entomological studies, and are well suited to a comparative study of commensal ism.

During preliminary collections of insects and their acarine symbionts u

I found representatives of the families mentioned above, especially the

necrophiles, to be common hosts of mite symbionts.

Examination of several representatives of Silphidae and Geotrupidae

indicated what appeared to be consistent distributional patterns of sessile mites, and perhaps some host specificity of the respective mite faunas. As a result of these apparent patterns and preferences, a three-year study was undertaken to determine the distribution and com­ position of populations of mites symbiotic on necrophilous beetles. 5

MATERIALS AND METHODS

This investigation was begun in mid-July, 1965, after preliminary

collections of beetles in light traps and pitfall traps yielded a high

proportion of necrophilous beetles with mite populations. Collections

were made from July through September 1965, July through October 1966,

and April through September 1967.

It was decided to limit the study to mites symbiotic on necrophilous

beetles in order to restrict the of the host beetles as much as

possible and yet make a diverse collection of the several families of

Coleoptera frequenting decomposing carrion.

Two collection sites were used. The first was located 41°55' N,

93°39' W in the region of an ecotone between an oak-hickory forest com­

munity and grassland on the Iowa State University Science Farm

2 miles south of Ames, Iowa. After two collections the site proved to

be unsatisfactory, so a new site was selected in an undisturbed area of the Thomas Judge Farm, 42°2' N, 93°40' W, % mile northeast of Ontario,

Iowa. This site was also in an ecotone region between oak-hickory forest and grassland.

Necrophilous beetles were collected from carrion baits. Shallow pits were excavated so as to remove vegetation in order to easily see

beetles and also to permit covering the carcasses with pieces of plywood.

The whole area was then camouflaged with branches and bits of litter.

The camouflaged covers helped to keep the carrion out of sight of curious 6

hikers who sometimes passed through the area.

It was difficult to obtain dead in sufficient quantity to maintain a constant supply of fresh carrion, so when a number of dead animals were obtained they were frozen in a large freezer which main­ tained -20°C. Frozen carcasses were allowed to thaw prior to place­ ment in a set.

A wide variety of animal species was used for carrion baits. The total for the three-year study is as follows: 25 chickens (Gallus domesticus), seven turkeys (Meleagris gallopavo), 53 barn pigeons

(Columba livia), one calf (Bos taurus), two sheep (Ovis ovis), five swine (Sus scrofa), two cats (Pelis domestica), one raccoon (Procyon lotor), 18 dogs (Canis familiaris), three guinea pigs (Cavia porcellus),

36 fox squirrels (Sciurus niger rufiventer), about 100 laboratory mice

(Mus musculus), and one woodchuck (Marmota monax).

Due to the difficulty in procuring carcasses, no definite pattern was maintained for carrion sets. The entire surface of each carcass was carefully examined daily. Only adult beetles on the carcass or directly under it were collected. Specimens were segregated by species and placed into plastic grinder jars with a bit of moist paper toweling, and lids were lightly screwed on. Jars were then stored briefly in a laboratory refrigerator at approximately 40°F. Carrion baits were not allowed to accumulate in the study area. A few days after the carcasses became desiccated, they were buried.. 7

For each collection were recorded in a field record book date,

type of carrion, duration of , and number of specimens of

beetles collected (Table 9, Appendix E). Each beetle collection was

numbered in a series for the year of the study. For example the tenth collection in 1965 received the number 65-10.

After cooling in the refrigerator, individual beetles were placed

into covered stender dishes. Each specimen was examined live under a dissection microscope in order to note any movements of mites on the external surfaces of the beetle. The specimen was then killed by adding a piece of filter paper saturated with chloroform to the covered stender dish.

The identity of each beetle was recorded by adding a third numeral to the appropriate collection symbol. For exançle, the first beetle examined from collection 65-10 would receive the number 65-10:1; the second 65 -10:2 and so on.

Beetles were removed from stender dishes and examined under a binocular dissection microscope with the aid of fine jeweler's forceps.

One pair of forceps was used to hold the beetle stationary while the other pair was used to manipulate body parts. The ventral surface of the beetle was examined beginning with the head. All sclerites and tagmata were manipulated in order to see all external surfaces. Legs were manipulated so as to examine the surface of all three pairs of coxal cavities. After careful examination of the venter, the dorsum was examined beginning with the head and proceeding posteriad. The last areas to be examined were the elytral concavities, wings, meso- and 8

metanota and tergites of the abdomen. Examination of these areas was

accomplished by unhooking the elytra and carefully unfolding the wings.

Macrophotographs were made of morphological regions and mite distribu­

tions whenever possible with an Asahi Pentax Spotmatic coupled to ex­

tension bellows. Some photographs were made with the camera mounted

on an A. 0. Spencer dissection microscope. Illumination was by A. 0.

Spencer No. 1036 and No. 735 illuminators. Macrophotographs were ex­

posed on Kodak Plus-X Pan film.

Mites were removed from the host beetles with fine tungsten wire

needles and loops mounted in wooden applicator sticks. The following

data were recorded: number of mites, bilaterality of distribution, and

region of beetle morphology. Mite specimens then were stored tempor­ arily in microvials containing Oudemans' solution (Evans et al., 1961).

Each vial was labeled with the above data and the collection number of the beetle. The external morphological regions of beetles were cate­ gorized as follows: (1) Venter - head, cervix, prosternum, propleuron,

procoxal cavities, mesosternum, mesopleuron, mesocoxal cavities, meta- sternum, metapleuron, metacoxal cavities, abdominal sternite I dorsal to coxae, abdominal sternites I - III excluding dorsal to coxae, and

abdominal sternites IV to apex of abdomen; (2) Dorsum - head, cervix,

pronotum, mesonotum, elytral concavity, metanotum, ventral to scutellum, wings, abdominal tergites I - III, abdominal tergites IV to apex of abdomen, legs I, legs II, legs III. Any ambulatory mite not attached to the surface of the beetle by means of chelicerae, anal pedicel, or opisthoso- 9

mal suckers was designated as XF (external free) even if the mites were encountered under the elytra.

After examination of beetle specimens, their wings were folded under the elytra and the latter were repositioned in repose. Beetles were pinned and stored in Schmitt boxes for: reference.

All mite symbionts were mounted on microscope slides prior to identification. Mite specimens were transferred by pipette from the microvials of Oudemans' solution to 27-mm diameter U.S. Bureau of Plant

Industry watch glasses. Nesbitt's clearing solution (Evans et al.,

1961) was added in sufficient volume to cover the specimens. Small squares of plastic were placed over the watch glasses to prevent evapo­ ration and crystalization of the clearing solution. Clearing was al­ lowed to take place at room temperature, and for times varying with the amount of sclerotization of the mite. Lightly sclerotized specimens such as hypopi usually required one to two days, whereas heavily sclero­ tized specimens such as uropodid deutonymphs required as much as nine days to render the integument translucent.

Cleared specimens were mounted ventral side up in methyl cellulose mountant (Evans et al., 1961) on microscope slides. A 12-mm diameter, number one coverslip was placed on each slide preparation. Slides were marked with the host beetle number and the morphological region of the host and placed in a drying oven. After the mountant had dried to the degree of hardness desired, coverslips were ringed with Zut slide ringing compound^ to prevent absorption of moisture by the mounting

^Bennett's Paint Co., Salt Lake City, Utah. 10

medium. When large numbers of mites were taken from a region, they were mounted in lots of ten and the several slides were numbered in a series so that the total number of mite specimens could be determined readily.

Beetle specimens were identified to family based on the classifica­ tion of Crowson (1955) and Arnett (1968). Identifications to genus were made by use of Arnett's comprehensive work, Dillon and Dillon (1961), and Jaques (1951). Several coleoptera specialists made determinations of some beetle specimens. Mr. Kenneth L. Esau^ identified the speci-

2 mens of Carabidae, Mr. William V. Miller identified several species of 3 Philonthus (Staphylinidae) and Dr. Rupert L. Wenzel determined many of the .

References used for identification of Coleoptera to species were;

Arnett (1944, 1946, 1947), Casey (1900), Dillon and Dillon (1961),

Hatch and Ortenburger (1930), Howden (1955, 1964), Howden and Cartwright

(1963) and Jaques (1951).

Identification of mite specimens presented a special problem in that the of most of the groups included in this study is un- stabilized. Grandjean's (1932, 1935, 1954) classification was followed at the ordinal level. Family determinations were made with the aid of

^Esau, Kenneth L., Department of Zoology and Entomology, Iowa State University of Science and Technology, Ames, Iowa 50010. Determination of Carabidae. Private communication. 1968. 2 Miller, William V., 57 Arlena Terrace, Ramsey, New Jersey 07446. Determination of Staphylinidae. Private communication. 1968. 3 Wenzel, Rupert L. Curator of Insects, Chicago Natural History Museum, Roosevelt Road and Lake Shore Drive, Chicago, Illinois 60604. Determination of Histeridae. Private communication. 1968. 11

Baker et al. (1958), Baker and Wharton (1952), Evans (1957) and

Strandtmann and Wharton (1958). Identification to genus, and whenever possible to species, was accomplished with the aid of Turk and Turk

(1957), Scheucher (1957), Krczal (1959), Karafiat (1959), Griffiths

(1964), Hughes and Jackson (1958), Hughes (1961), Zakhvatkin (1941),

Michael (1901, 1903), Nesbitt (1945), Vitzthum (1930), and Krantz (1962).

Dr. Preston Hunter^ made several determinations of mesostigmatid mites.

All mite specimens were viewed with an A. 0. Spencer Series 10 phase-contrast microscope. Those that could not be determined to species were photomicrographed with an Asahi Pentax Spotmatic, Kodak Plus-X

Pan film and A. 0. Spencer photo-tube mounted on the above microscope

(Appendix B).

The composition and distribution of mite symbionts on host beetles are presented in Table 8, Appendix D.

Hunter, Preston E., Department of Entomology, The University of Georgia, Athens, Georgia 30601. Determination of mesostigmatic mites. Private communication. 1968. 12

RESULTS

Host Beetles and the Morphological Distribution of Their Mite Symbionts

A total of 459 beetles representing nine families were collected

from carrion baits. Of these 223 were infested with a total of 5,816

mite symbionts.

Mite populations and their regional distribution on each host beetle

are presented in Table 8 in the Appendix.

Carabidae

Carabid beetles are not one of the principal members of the necro­

philous fauna. Only two specimens of carabids were taken in the entire

study.

Harpalus pennsylvanicus One specimen was taken. A single speci­

men of Imparipes sp. (Scutacaridae) was collected from the right meta-

coxa (Figure 21).

Anisodactylus (Anisodactylus) agricola One specimen was

collected. No mite symbionts were noted.

Histeridae

Histerid beetles are common members of the necrophilous beetle

fauna. Their role in the carrion community is principally prédation.

Uisterids can be observed preying on the larvae of other insects such

as calliphorids.

Saprinus assimilis Forty-seven specimens were collected with six beetles ha.ing mites on their bodies. All mites were located 13

in the concavities formed by the first three abdominal tergites beneath the wings and elytra. Figure 1 shows the dorsal abdominal concavity

of S. assimilis with elytra and wings removedA, total of nine mites was taken from the six beetles; one laelapid (Figure 22), six halo­

laelapid deutonymphs (Figure 23), and two unidentified mesostigmatid 1 deutonymphs (Figures 24 and 25).

Saprinus lugens Two specimens were collected. One dead specimen of Hypoaspis (Laelapsis) vitzthumi (Laelapidae)^ was taken from a mycelial mat on the mesosternum of one of the beetles. An 2 attempt to identify the fungus was unsuccessful.

Margarinotus hudsonicus One specimen was collected. No mites were found.

Hister abbreviatus One specimen was collected. No mites were found.

Hister circinaris Nineteen specimens were collected. A total of four halolaelapid deutonymphs were found in the concavities of the first three abdominal tergites on two beetles. One specimen of H. circinans had three mites; the other had one.

Silphidae

The silphids, or carrion beetles, are truly necrophagous. They feed upon the decomposing carcass, which is used also for oviposition and as a source of food by the . However, as mentioned earlier

^Hunter, Preston E., op. cit.

^Tiffany, Lois H. Department of Botany, Iowa State University of Science and Technology, Ames, Iowa 50010. Identification of fungi. Private communication. 1967. 14

only adult specimens were examined in this study.

Silpha noveboracensis Of 46 specimens collected, 42

had mites associated with the external morphology. Table 1 presents

the composite compilation of mite populations by morphological regions

for all infested specimens of S- noveboracensis.

It can be seen from Table 1 that hypopi of Histiostoma sp. A

(Figure 32) and H. sp. C (Figure 34) were widely distributed over several

regions of Silpha noveboracensis. The highest populations of hypopi were

encountered in the elytral concavities and on the first three abdominal

tergites. The mites on the elytra were all distributed on the lateral

margins of the elytral concavities. The marginal surface is smooth as

compared to medial surfaces. Figure 2 shows the surface of an elytron.

Figure 3 shows the bilateral distribution of Histiostoma sp. C on ab­

dominal tergites I-III. The most abundant hypopus was Histiostoma sp.

C., which was intermixed with specimens of H. sp. A. The only external,

ambulatory mites were deutonymphs of necrophori.

Silpha americana Sixty-two specimens were collected. Of these, 42 had mites associated with their external morphology. Table 2 shows the distribution of mite populations by morphological regions for

all specimens of S. americana with mites.

Mites were widely distributed on several regions of S. americana.

The primary regions having consistently high populations were the elytral concavities and abdominal sternites I-III. Figure 4 shows the marginal distribution of hypopi on an elytron. The morphology of the elytral sur- 15

Table 1. Distribution of mites by morphological region on Silpha noveboracens is

Regions occupied No. beetles No. mites

Venter: Etosternum 6 6 Hst. C. 1 Hst. A.

Propleuron 6 12 Hst. C.

Procoxal cavities 1 2 Hst. C.

Mesosternum 3 2 Hst. G. 1 Hst. A.

Mesocoxal cavities 1 1 Hst. C.

Abdominal sternite I dorsal to coxae 7 11 Hst. C. 1 Hst. A.

Abdominal sternites I-III excluding dorsal to coxae 4 8 Hst. C.

Abdominal sternites IV-apex 1 1 Hst. c.

Dorsum: Pronotum 1 1 Hst. c.

- Elytral concavity 16 228 Hst. C. 8 Hst* A*

Abdominal tergites I-III 31 1495 Hst. C. 149 Hst. A.

Abdominal tergites IV-apex 2 6 Hst. c.

Legs: I 1 1_ Hst. C on Left Coxa

III 1 1 Hst. C. on Left Coxa

External, ambulatory 20 74 Poec.. nec. DN

No. of beetles Total mites on beetles with mites 42 160 Hst. A. 1775 Hst. C. 74 Poec. Nec. DN 16

Table 2. Distribution of mites by morphological regions on Silpha americana

Regions occupied No. beetles No. mites

Venter; Head (R compound eye) 1 1 Hst. C.

Prosternum 1 1 Hst. C.

Procoxal cavities 2 1 Hst. C. 13 Hst. A.

Mesosternum 1 2 Hst. C. 5 Hst. A.

Mesocoxal cavities 2 1 Hst. C. 1 Hst. A.

Abdominal sternite I dorsal to coxae 4 8 Hst. C. 1 C£. A.

Abdominal sternites I-III excluding dorsal to coxae 1 1 Hst. c.

Dorsum: Elytra1 concavity 19 57 Hst. c. 23 Hst. A.

Wings 1 1 Pyemotes sp.

Abdominal tergites I-III 11 27 Hst. c. 21 Hst. A. 1 Mc. A?- Abdominal tergites IV-apex 1 1 Hst. c.

Legs; I 1 2 Urp. tibia

III 3 1 Hst. C. coxa, 1 Cg^. A. coxa, 3 Urp. tibia

External, ambulatory 34 415 Poec. nec. DN

Total beetles Total mites with mites 42 42 63 Hst. A. 99 Hst. C. 3 C^. A. 1 Pyemotes sp. 1 Mc. A? 415 Poec. nec. DN 5 Urp. 17

faces of this species is quite similar to that of Silpha noveboracensis

(Figure 2). A total of 415 ambulatory Poecilochirus necrophori deuto-

nymphs were collected. Three specimens of Galoglyphus sp. A (Acaridae)

(Figure 37) were taken from abdominal sternite I, abdominal tergites

I-III, and the metacoxa. Figure 5 shows the abdominal tergites of

Silpha americana. A single specimen of Pyemotes sp. (Pyemotidae)

(Figure 26) was taken from the right wing of a beetle. -This was the only instance of a mite being attached to wings in the entire study.

Five uropodid deutonymphs (Figure 27) were attached to the legs of four beetles. One Macrocheles sp. A adult ? (Figure 28) was attached by chelicerae to the dorsum of the abdomen of one beetle.

Silpha surinamensis Thirty-three specimens were col­ lected. Of these, 16 beetles had mites associated with them. Table 3 shows the composite compilation of mite populations by morphological re­ gions for specimens of S. surinamensis with mites. Specimens of S. surinamensis had a low incidence of infestation of mites as compared with other silphid beetles, and the regions which harbored mites were fewer by comparison. The highest hypopial populations were encountered in the elytral concavities. Only six hypopi were taken from the ab­ dominal tergites. It should be pointed out that the dorsal surfaces of sub-elytral segments of the abdomen are membranous with little if any sclerotization (Figure 6).

Silpha lapponica of six specimens collected, one beetle had mites. These were distributed as follows: one Histiostoma sp. C. 18

Table 3. Distribution of mites by morphological regions on Silpha surinamensis

Regions occupied No. beetles No. mites

Venter: Abdominal sternites I-III excluding dorsal to coxae 1 2 Hst. C.

Dorsum: Pronotum 2 1 Hst. C. 1 Hst. A.

Elytral concavity 11 198 Hst. C. 2 Hst. A.

Abdominal tergites I-III 2 6 Hst. C.

Legs: I 1 1 Urp. R femur

III 2 2 Urp. L tibia

External, ambulatory 7 10 Poec. nec. DN

Total beetles with Total mites mites 16 3 Hst. A. 207 Hst. C. 3 Urp. 10 Poec. Nec. DN

in the right elytral concavity, four H. sp. C equally distributed on abdominal tergites I-III, and one Poecilochirus necrophori ambulatory deutonymph.

Nicrophorus tomentosus Beetles belonging to the genus

Nicrophonis are referred to as the burying or sexton beetles because of their habit of burying small animals, an act which undoubtedly delays the decomposition of the carcass and prevents oviposition by flies. 19

These beetles are encountered also on larger carcasses. Nicrophorus tomentosus is the most common member of the genus in central Iowa.

Fifty-one specimens were collected; 44 had mites. Table 4 shows the composition of mite populations by morphological regions for specimens of N. tomentosus. Of hypopi, only Histiostoma sp. A was present, a few being taken from scattered regions; however, the consistently localized populations occurred in mesocoxal cavities. The hypopi were distributed only on the outer smooth region of the cavities, never on the inner roughened surface and never on the coxa proper. Figure 7 shows the above- described textures of the left mesocoxal cavity of N. tomentosus. Six specimens of Macrocheles sp. A adult?? were collected. Four were at­ tached by their chelicerae to prothoracic spiracles just under the spirac- ular lobe of the pronotum (Figure 9). More will be said about this re­ gion in the discussion section. Only one hypopus of Histiostoma sp. A was collected from the elytral concavities, and none from the abdominal tergites (Figure 8). Poecilochirus necrophori deutonymphs were abundant and occurred on 23 of the beetles.

Nicrophorus orbicollis Thirteen specimens were collected and 12 had mites. Table 5 presents the distribution of mite populations by morphological regions for specimens of N. orbicollis.

Although mites were distributed over several body regions, two regions had consistent infestations of Macrocheles sp. A adult??. These were the prothoracic spiracles and abdominal tergites l-lll. The pro­ thoracic spiracles of six beetles had M. sp. A adult f? attached. The 20

Table 4. Distribution of mites by morphological regions on Nicrophorus tomentosus

Regions occupied No. beetles No. mites

Venter: Prosternum 3 1 Hst. A 2 Urp.

Propleuron 2 1 A? on pro- thoracic spiracles

Procoxal cavities 3 5 Hst. A

Mesosternum 1 3 Hst. A 3 Mc. A

Mesopleuron 2 1 Hst. A 1 Urp.

Mesocoxal cavities 19 643 Hst. A on smooth surface only Abdominal sternites I-III excluding dorsal to coxae 1 1 Urp. 2 Mc. Aff chelicerae attached to setae

Dorsum: Elytral concavity 1 1 Hst. A

External, ambulatory 23 289 Poec. nec. DN

Total beetles with Total mites mites 33 654 Hst. A 4 Urp. 6 Mc. A?? 289 Poec. nee. DN 21

Table 5. Distribution of mites by morphological regions on Nicrophorus orbicollis

Regions occupied No. beetles No. mites

Venter: Propleuron 6 67 A?? on pro- thoracic spiracles

Procoxal cavities 3 24 Hst. C 6 Hst. A

Mesosternum 1 2 Hst. A

Mesocoxal cavities 2 58 Hst. C 1 Hst. A

Abdominal sternite I dorsal to coxae 1 60 Hst. C

Dorsum: Pronotum 2 1 Urp.

Metanotum 1 34 Hst. A

Abdominal tergites I-III 10 270 Mc« A $9- in mem­ branous depressions of abd. tergites I-III Abdominal tergites IV-apex 1 93 Hst. C

External, ambulatory 11 59 Poec. nec. DN

Total beetles with Total mites mites 12 43 Hst. A 235 Hst. C 337 Mc. A 59 Poec. nec. DN 22

spiracular lobe of the pronotum extends ventrocaudad in such a way that there is a well-protected area between the prothoracic spiracles and the lobe of the pronotum. Figure 10 shows the region described above. Figure 11 shows four Macrocheles sp. A adult attached to the left prothoracic spiracle of a specimen of Nicrophorus orbicollis. The largest numbers of mites were taken from the abdominal tergites I-III, which on N. orbicollis form a membranous pocket (Figures 12 and 13).

Hypopial deutonymphs on the dorsum were restricted to the metanotum and abdominal tergite IV. These latter regions are smooth and somewhat sclerotized in N. tomentosus (Figure 8), whereas tergites I-III are very membranous and forms a depression in N. orbicollis.

Although only two beetles had hypopi attached to the mesocoxal cavities, it is interesting to note that a similar condition of smooth and rough areas occurs in the cavities. The smooth surface is medial whereas the roughened surface is lateral. Figure 14 shows 23 Histiostoma sp. C attached to the smooth surface of the mesocoxal cavity. Figure 15 shows the same cavities with mites removed. Poecilochirus necrophori deutonymphs were taken from all but two beetles.

Nicrophorus pustulatus Three specimens were collected, all of which had mites. N. pustulatus is not common to Central Iowa but is more western in its distribution (Jaques, 1951). Two specimens of Histiostoma sp. C were taken from the pronotum of one beetle, and a total of 47 specimens of Poecilochirus necrophori ambulatory deutonymphs were taken. 23

Prionochaeta opaca Although Arnett (1968) stated that

this species is commonly found with other silphids and staphylinids on

carrion, I collected only one specimen of P. opaca. No mites were found.

Staphylinidae ' -

Staphylinid beetles are common members of the necrophilous beetle fauna. Their role, like that of the Histeridae, is chiefly one of pré­ dation. Staphylinids are highly cursorial and present a difficult quarry for the collector at carrion baits.

Staphylinus maxillosus This beetle is the most common beetle on carrion in Central Iowa. One hundred and seventy-one beetles were collected; of these, 42 had mites associated with their external morphology. Table 6 presents the composition of mite populations by regions for specimens of S. maxillosus.

A heterogeneous population of mites occurred on Staphylinus maxillosus beetles. Hypopial forms were represented by Histiostoma spp.

A, B (Figure 33), and C; by Caloglyphus spp. A, C (Figure 39), and D

(Figure 40); and by hypopi of Acarus siro. One specimen of Poecilochirus necrophori ambulatory deutonymph was found.

Four morphological regions were represented showing relatively consistent infestation by hypopi. The prosterniim of 12 beetles had from one to five specimens of hypopi attached in the posterior furrow (Figure

16). Mites found on the prosternum always occupied the furrow. Twelve beetles had hypopi in the mesocoxal cavities, the surfaces of which are all uniformly smooth so there was no distributional pattern of hypopi 24

Table 6. Distribution of mites by morphological regions on Staphylinus maxillosus

Region occupied No. beetles No. mites

Venter; Head (gula) 2 4 Hst. C 3 C£. C

Prosternum 12 19 Hst. C 3 Hst. A 1 Ç&. A

Procoxal cavities 4 2 Hst. C 3 Hst. A 1 Acarus siro

Mesosternum 1 1 Hst. C

Mesocoxal cavities 12 10 Hst. C 4 Hst. A 3 Acarus siro

Metasternum 3 2 Hst. C 1 Cg,. A

Abdominal sternites I-III excluding dorsal to coxae 1 1 Hst. C

Dorsum:

Elytral concavity (apical portion only) 24 48 Hst. A 79 Hst. B 22 Hst. C 1 Cg. A 6 eg,. C 1 Cg. D 1 Acarus siro

Metanotum 5 110 Hst. C

External, ambulatory 1 Poec. nec. ON

Total beetles Total mites with mites 42 58 Hst. A 79 Hst. B 171 Hst. C 3 Ç&. A 9 Ç&. C 1 Cg. D 5 Acarus siro 1 Poec. nec. DN 25

as there was in the silphids. There were, however, no hypopi attached

to coxae. The apex of the elytral concavities was the most common

region occupied by hypopi of Histiostoma spp. A, B and C. Caloglyphus

spp. A, C and D (Figures 37, 39 and 40), and Acarus siro were all en­ countered in the apical portions of elytral concavities of 24 beetles.

Five beetles had hypopial populations of Histiostoma sp. C on the metanotum. It is interesting to note that in Staphylinus maxillosus, as well as in most staphylinids, the entire dorsum of the abdomen is exposed because the elytra are short and the wings fold in an intricate pattern between the elytra and metanotum.

Staphylinus maculosus Two specimens were collected yielding no mites.

Philonthus politus Three specimens were collected, two of which had mites. Twenty-two Histiostoma Sp. D (Figure 35) were taken from the mesocoxal cavities, and four H. sp. D hypopi were taken from the apices of the elytral concavities. Two uropodid deutonymphs were attached to the metacoxae of one beetle.

Philonthus validus Of seven specimens collected three had mites. Thirteen hypopi of Histiostoma (2 H. sp. A, 11 H. sp. D) were taken from the gulae of two beetles. Four Histiostoma sp. D were taken from the prosternum of one beetle, and ten H. sp. D were found in the mesocoxal cavities. Five H. sp. C were attached to abdominal stemite two, and five H. sp. C were taken from the metanotum. Eight Histiostoma 26

sp. B were taken from the apices of the elytral concavities. One uro-

podid deutonymph was attached to the right femur.

Philonthus cyanipennis Only one specimen was collected yield­

ing no mites.

Ontholestes cingulatus Twenty specimens were collected, but

only two had mites. One specimen had 86 Histiostoma sp. B hypopi at­

tached to the apices of the elytral concavities. The other beetle had

one H. sp. E (Figure 36) attached to the apex of the right elytral con­

cavity.

Trogidae

The trogids or skin beetles are among the last beetles to arrive

at a carrion set. They feed on the dry skin of the carcass. I did not

continue checking carcasses for prolonged periods, but buried the dried skin and bones several days after the carcass reached the dry state.

Undoubtedly more trogid specimens could have been obtained if collec­ tions were made from dried carcasses for longer periods.

Trox unistriatus Two specimens were collected. No mites were found.

Trox suberosus Two specimens were collected. No mites were found.

Geotrupldae

Geotrupid beetles are found on carrion and dung. Adults burrow under carcasses and bring carrion down into subterranean tunnels where 27

larvae are reared. In order to collect large numbers of these specimens,

special digging techniques must be followed which ultimately disturb

the carrion bait (Howden, 1955). Only those geotrupids on or directly

under carcasses were taken. No attempt at digging into burrows was made.

Geotrupes semiopacus IWenty-one specimens were collected, but

only three had mites. One beetle had 14 hypopi of Galoglyphus sp. B

(Figure 38) attached to the middle of the prosternum, 2 C. sp. B attached to the middle of the mesosternum, and one G. sp. B attached to the left elytral concavity. The other two beetles each had a single specimen of the ambulatory Macrocheles sp. B adult 9^ (Figure 29).

Geotrupes splendidus Fifteen specimens were collected and nine had mites. The principal .region occupied was the elytral concavity where as many as 642 Caloglyphus sp. B were encountered on one specimen (Fig­ ure 17). IVo other beetles had 373 and 300 C. sp. B respectively in . the elytral concavities. A few Caloglyphus spp. were found on scattered regions. Two beetles had one Macrocheles sp. B adult Ç ambulatory mite on the external surface. Figure 18 shows the dorsum of the abdomen of Geotrupes splendidus. No hypopi were taken from the abdominal tergites of either species of Geotrupes. The lack of sclerotization and the dense

pattern of setation on the abdomen may play a role in limiting distribu­ tion of hypopi to the elytra.

Scarabaeidae

Although the scarabaeid beetles taken in this study are primarily coprophiles, they occasionally occur on carrion. 28

Onthophagus hectate Five specimens were collected. No mites

were taken.

Onthophagus janus One specimen was collected yielding no

mites.

Copris tullius One specimen was collected. It had four mites

three of which were uropodid deutonymphs attached to the midregion of

the prosternum, and one was an external ambulatory Alliphis sp.

(Eviphididae)^ (Figure 30).

Dermestidae

Dermestid beetles, like the trogids, are more commonly found in

the last stages of carrion decomposition when the carcass is quite dry.

Dermestes vulpinus Eleven beetles were collected. One beetle

had a population of 75 saproglyphid hypopi (Figure 31) bilaterally dis­

tributed on abdominal tergites I-III (Figure 19).

Dermestes caninus Seven beetles were collected. No mites

were found.

Nitidulidae

Nitidulids are not common to necrophilous fauna. Only two speci­

mens were collected.

Stelidota geminata One specimen was collected. No mites were

found.

Glischrochilus fasciatus One specimen was collected. No mites were found.

^Hunter, Preston E., op. cit. 29

Mite Symbionts and Their Occurrence on Host Beetles

Table 7 presents a tabulation of all mites taken from host beetles.

The number of mites taken from each species of beetle is given and the

number of beetle specimens taken with a particular mite species. Photo­

micrographs of all mites not identified to species are shown in Appen­ dix B, Plates 7 to 12.

Specificity of hosts and morphological regions occupied can be judged from the values presented in Table 7. Table 7. Mite symbionts and their occurrence on host beetles

Mites Host beetles Regions occupied by mites

No. No. Scutacaridae 1 Imparipes sp. 1 Harpalus pennsylvanicus Right metacoxa

1 Laelapidae adult Ç 1 Saprinus assimilis Concavity of abdominal tergites I-III 1 Hypoaspis (Laelapsis) Mesosternum surrounded by vitzthumi 1 Saprinus lugens fungal mycelium 2~

6 Halolaelapidae DN 4 Saprinus assimilis Concavity of abdominal tergites I-III

Halolaelapidae DN 2 Hister circinans Concavity of abdominal tergites I-•III 10

2 Mesostigmatid Kî 1 Saprinus assimilis Concavity of abdominal tergites I-•III

1 Pyemotidae Pyemotes sp. 1 Silpha americana Wing

5 Uropodidae DN 1 Silpha americana Tibiae I and III

3 tt «1 3 Silpha surinamensis Tibiae, femora I and III

4 4 Nicrophorus tomentosus Prosternum, abdominal sternites I--III, mesopleuron

1 t1 1 Nicrophorus orbicollis Pronotum Table 7. (Continued)

Mites Host beetles Regions occupied by mites

No.

2 Uropodidae DN 1 Philonthus politus Coxa III

1 • 1 n 1 Philonthus validus Fémur I

3 tt 11 1 Copris tullius Prosternum 19

Macrochelidae 1 Macrocheles sp. A adult ^ Silpha americana Abd. tergites I-III attached by chelicerae

6 Il tt »l M 2 Nicrophorus tomentosus Prothoracic spiracles, abd. sternites I-III attached by chelicerae

• 1 1» " 10 Nicrophorus orbicollis Prothoracic spiracles, concavity of !L abd. tergites I-III attached by 324 chelicerae

2 Macrocheles sp. B adult^ 2 Geotrupes semiopacus XF, ambulatory

2 " " " " 2 Geotrupes splendidus XF, ambulatory Table 7. (Continued)

Mites Host beetles Regions occupied by mites

No. No. Poecilochims 74 necrophori DN 20 Silpha noveboracens is XF, ambulatory

414 34 Silpha americana XF, ambulatory

9 6 Silpha surinamensis XF, ambulatory

1 1 Silpha lapponica XF, ambulatory

179 34 Nicrophorus tomentosus XF, ambulatory

59 11 Nicrophorus orbicollis XF, ambulatory

47 3 Nicrophorus pustulatus XF, ambulatory

1 1 Staphylinus maxillosus XF, ambulatory

784

Eviphididae 1 Alliphis sp. 1 Copris tullius XF, ambulatory

Saproglyphidae 75 Hypopi 1 Dermestes vulpinus Abd. tergites I-III Table 7. (Continued)

Mites Host beetles Regions occupied by mites

No. No. Anoetidae Histiostoma sp. A 160 Hypopi 16 Silpha noveboracens is Prosternum, mesosternum, abd. sternite I dorsal to coxae Elytral concavity, abd. tergites I-III

63 II II 11 Silpha americana Procoxal cavities, mesosternum, meso- coxal cavities, elytral concavities, abd. tergites I-III

3 U II 1 Silpha surinamensis Pronotum, elytral concavity

656 tr tt 21 Nicrophorus tomentosus Prosternum, procoxal cavities, meso­ sternum, mesocoxal cavities, elytral concavities

43 II II 4 Nicrophorus orbicollis Procoxal cavities, mesosternum, metanotum

58 17 Staphylinus maxillosus Prosternum, procoxal cavities, meso­ coxal cavities, elytral concavity

983

Histiostoma sp. B 7 9 Hypopi 2 Staphylinus maxillosus Elytral concavity

8 II n 1 Philonthus validus Elytral concavity

86 II II 1 Ontholestes cingulatus Elytral concavity 173 Table 7. (Continued;

Mites Host beetles Regions occupied by mites

No. No. Histiostoma sp. C 118 5 Hypopi 34 Silpha noveboracens is Prosternum, propleuron,mesosternum, mesocoxal cavities, ^d. sternites I-III, elytral concavity, abd. ter- gites I-III, coxa III

104 20 Silpha americana Prosternum, procoxal cavities, meso­ sternum,mesocoxal cavities, abd. sternites I-III elytral concavities abd. tergites I-III coxa III

207 10 Silpha surinamensis Abd. sternites I-III pronotum, ely­ tral concavity, abd. tergites I-III

5 It fl 1 Silpha lapponica Elytral concavity, abd. tergites I-III

235 1 Nicrophofus orbicollis Procoxal cavities, mesocoxal cavities abd. sternites I-III, abd. tergite IV

2 If II 1 Nicrophorus pustulatus Pronotum

171 26 Staphylinus maxillosus Gula, prosternum, procoxal cavities, mesosternum, mesocoxal cavities, metasternum, abd. sternite I, apices of elytral concavities, metanotum

12 II II 1 Philonthus validus Gula, abd. sternites I-III, metanotum

1921 Table 7. (Continued)

Mites Host beetles Regions occupied by mites

No. No Histiostoma sp. D 23 Hypopi 1 Philonthus politus Mesocoxal cavities

25 tt tt 1 Philonthus validas Quia, prosternum, mesocoxal cavities

48

Histiostoma sp. E 1 Hypopi 1 Ontholestes cingulatus Elytral concavity

Acaridae 73 Acarus siro 1 Staphylinus maxillosus Procoxal cavities, mesocoxal cavities, elytral concavity

Galoglyphus sp. A 1 Hypopi 1 Margarinotus hudsonicus Mesopleuron

3 ft It tt 2 Silpha americana Abd. sternite I, abd. tergites I-III, coxa III

3 t1 tt tt 3 Staphylinus maxillosus Prosternum, metasternum, elytral concavity

9 tt It tt 2 Geotrupes splendidus Tibiae I and II, femora III, coxa III

16 Table 7. (Continued)

Mites Host beetles Regions occupied by mites

No. No. Galoglyphus sp. B 17 Hypopi 1 Geotrupes semiopacus Prosternum, mesosternum, elytral concav ity

1391 " " " 5 Geotrupes splendidus Elytral concavity

1408

Galoglyphus sp. G 9 Hypopi 3 Staphylinus maxillosus Gula, elytral concavity

28 " " " 2 Geotrupes splendidus Prosternum, procoxal cavities, meso- sternum, abd. stemite I, coxae III

37

Galoglyphus sp. D 1 Hypopi 1 Staphylinus maxillosus Elytral concavity

5,816 37

DISCUSSION

Beetle families collected with more than a few specimens will be discussed with regard to size and distribution of mite populations on the beetles.

Histeridae;

Saprinus assimilis and Hister circinans were the two common repre­ sentatives of histerids collected on carrion. Deutonymphs of the family

Halolaelapidae were taken from both species of histerids; two unidenti­ fied mesostigmatids and a single adult female laelapid were taken from

Saprinus assimilis only. All the mites mentioned above were found in the concavities of the dorsal abdominal tergites I-III. The symbiotic relationship of these mites with histerid beetles is unknown inasmuch as there is no information in the literature concerning their ecological niche. It appears that these mites are involved in a phoretic commensal- ism in that none was attached to the integument. All were ambulatory, but were contained within the cavities of the abdominal tergites. The mode of entry into this cavity can only be conjectured because elytra in the normal position fit very tightly, almost as though machined, against the mesothorax and abdomen.

Silphidae;

Silpha noveboracensis: Two families of mites occurred on silphids.

The family Anoetidae was represented by hypopi of Histiostoma sp. A

(160) and H. sp. C (1,775). The family Parasitidae was represented by 38

the ambulatory deutonymph Poecilochirus tiecrophori. The sites occupied

by the hypopial forms were primarily the first three abdominal tergites.

The distributional pattern of the hypopi was typically bilateral, so that for virtually all populations of 10 or more the hypopi were about equally distributed on either side. Figure 3 shows a typical bilateral distribution of hypopi. The pattern on the elytra was unusual in that hypopi attached only to the lateral margins of the elytral concavities.

This surface (Figure 2) is relatively smooth, whereas the remainder is rather rough in texture.

Seventy-four ambulatory deutonymphs of Poecilochirus necrophori were distributed over 20 beetles. These deutonumphs could be seen walking about over all surfaces of the beetle, and were also observed feeding on the surface of the carrion. When disturbed on beetles the mites often moved under the elytra; when off beetles, mites usually moved onto the surface of the nearest beetle.

Silpha americana; Six families were represented by mites taken from these beetles. Anoetid hypopi consisted of 69 Histiostoma sp. A and 99^. sp. C. Three hypopi of Caloglyphus sp. A (family Acaridae) were found. A single specimen of Pyemotes sp. (family Pyemotidae) was collected. One Macrocheles sp. A adult $ of the Macrochelidae, 415

Poecilochirus necrophori deutonymphs of family Parasitidae and five uropodid deutonymphs also were taken. Table 8 in the Appendix shows that the only consistent mite populations occurred in the elytral con­ 39

cavities on the smooth lateral marginal surfaces, and on abdominal ter- gites I-III. Figures 4 and 5 show the regions mentioned above. It should be noted here that the morphology of the elytra of Silpha americana is very similar to that of noveboracensis. Poecilochirus necrophori were very abundant and responded to disturbances exactly as they did on S. noveboracens is.

Silpha surinamensis; Anoetid hypopi of Histiostoma sp. A and H. sp. C were taken primarily from the surfaces of the elytral concavities.

Two specimens of H. sp. C were found on the first abdominal tergite of one beetle. It was the only instance of a hypopus attached to the dorsum of the abdomen of Silpha surinamensis. The first four abdominal tergites lack sclerotization in S. surinamensis to the degree that they would seem to provide an unsuitable substrate for suctorial hypopial attachment

(Figure 6). The tergites of S. noveboracens is and S. americana (Fig­ ures 3 and 5), however, are sclerotized to a degree that a rigid plate is formed on each segment. It is on these sclerites or rigid portions of the tergites that the hypopi were found. Characteristic of all three species of silphids was the prevalence of poecilochirid deutonymphs.

Nicrophorus tomentosus; On 44 beetles collected, Histiostoma sp.

A was the only hypopial representative. Most of these (643 out of a total of 654) were attached to the smooth outer, lateral posterior part of the mesocoxal cavity. Figure 7 shows the inner roughened surface and the outer smooth surface of the right mesocoxal cavity of N. tomen­ tosus. Mites were not found on the coxae. It appears that surface 40

textures of the coxal cavities were the main factors affecting distribu­

tion of the hypopi. Hypopi occupying the inner roughened surfaces would

not be afforded a uniform substrate to which suckers could be applied.

Furthermore the action of coxae as they rotate in the cavity would very

likely have some grinding action upon any object on the roughened sur­

face.

Three specimens of Macrocheles sp. A adultwere attached by

their chelicerae to the setae in the middle of the mesosternum of a

single beetle, and two Macrocheles sp. A adult to the abdominal setae

of sternites, while another M. sp. A adult^ was found attached to the

membrane of the prothoracic spiracle of a beetle. Figure 9, showing the

prothoracic spiracle with the spiracular lobe of the pronotum, indicates that much of the surface of the spiracle is exposed. Poecilochirus necrophori deutonymphs were common.

Nicrophorus orbicollis was not collected in abundance, but of the

12 beetles (of a total of 13 collected) that had mites, some interesting distributional patterns of mites were found. The most striking condi­ tion is the mode of attachment of Macrocheles sp. A adult . Of all the beetles examined, six yielded 67 M. sp. A adult attached to pro­ thoracic spiracles by their chelicerae. Figure 10 shows a prothoracic spiracle, and a pronotal spiracular lobe extending posteriacaudally over most of the spiracle. Figure 11 shows four M. sp. A adult attached to the prothoracic spiracle. It appears that the additional protection offered by the added covering of the spiracular lobe makes 41

this site more suitable to M. sp. A than the more exposed condition found in Nicrophorus tomentosus. Certainly the niche formed by the lobe covering the spiracle is a fairly secure one.

The other region occupied by Macrocheles sp. A adult was the membranous concavity formed by the first three abdominal tergites, in some ways resembling the niche occupied by ambulatory mites on histerids.

Of all Nicrophorus orbicollis examined, ten yielded 370 M. sp. A adult

Ç? attached to the membrane by their chelicerae. Figure 12 shows 145 M. sp. A adult occupying the concavity, and Figure 13 shows the exposed membranous concavity after removal of the macrochelids. Note the bi­ lateral distribution of hypopi of Histiostoma sp. C on abdominal tergite

IV. No hypopi were encountered on the first three membranous tergites even on those beetles which had no macrochelids present. When hypopi were encountered on the dorsum, they occurred on abdominal tergite IV or on the metanotum.

Although only two Nicrophorus orbicollis had hypopi attached to the mesocoxal cavities, the distributional pattern on the cavity surface

Ls worth noting. One beetle had only one Histiostoma sp. A attached; the other specimen had a total of 58 H. sp. C attached to the outer margin of the cavities. The mesocoxal cavities of Nicrophorus orbicollis ire similar to those of N. tomentosus in that the inner surfaces are roughened to a degree that attachment by hypopi would likely be insecure and the chances of grinding action of coxal movements would be increased.

Figure 14 shows the distributional pattern of Histiostoma sp. C in a 42

mesocoxal cavity, and Figure 15 shows the region with mites removed.

The procoxal cavities of this same beetle yielded 24 H. sp. C. It ap­

pears that with the high population of H. sp. C attached to this single

beetle optimal sites of attachment were overfilled and consequently would

shift distribution to secondary sites.

Staphylinidae;

Staphylinus maxillosus: These staphylinid beetles had a hetero­ geneous population of hypopial forms of Anoetidae; Histiostoma spp. A,

B and C; and of Acaridae; Acarus siro and Caloglyphus spp. A and C. A few hypopi were scattered regionally, but most appeared consistently on two main sites of attachment. These were the apices of the elytral con­ cavities and the furrow of the prosternum. Figure 16 shows the pro- sternum with the posterior furrow occupied by two Histiostoma sp. C.

When hypopi were attached to the presternum, they always were attached in this furrow. There is no evident reason as to why beetle 65-19:1 had the high population of H. sp. C located on the metanotum. This region was occupied on only four other beetle specimens with one H. sp. C found on each. A possible explanation is that populations shift regionally

Erom less optimal to more optimal regions. This has been shown experi­ mentally by Hunter and Mollin (1964) and Hunter and Davis (1965).

Laboratory studies involving marked specimens would be needed to answer this question. An attempt was made to study the spatial relationships of the folded wings to the elytra, metanotum, and dorsum of the abdomen, but no conclusions could be drawn. Very likely there is more space 43

toward the apex of the elytra than there is basally.

Only one specimen of Poecilochirus necrophori was found on a staphylinid beetle. I am inclined to believe that it was an accidental

occurrence as a result of my hasty pursuit of the beetle. Possibly the poecilochirid was picked up with the beetle from the surface of the carrion.

The remaining staphylinids showed similar distributional patterns of the hypopial populations; the apices of the elytra were the primary sites occupied by hypopi of Hst. sp. B only. Some hypopi occurred on the prosternum, but the pattern of distribution was more or less random.

No morphological features similar to the posterior furrow found on the prosternum of Staphylinus maxillosus are present on the prosternum of the other staphylinids.

Geotrupidae;

The morphology of Geotrupes semiopacus and G. splendidus is similar.

Consequently these two species will be discussed together. Only acarid hypopi were found on the geotrupids. Galoglyphus spp. A, B and G were taken from scattered body regions. Located in the elytral concavities were high populations of hypopi all of which were Galoglyphus sp. B

(Figure 38). Figure 17 shows a population of 315 Galoglyphus sp. B attached to the elytral concavity of Geotrupes splendidus. In no in­ stance were hypopi found attached to the dorsum of the abdomen or to the metanotum under the elytra. The dorsal surface of the abdominal tergites 44

is membranous and the metanotal region is quite hairy. Both of the pre­

vious conditions appear to be poo: siLes Cor hypopial attachment. Fig­

ure 18 shows the dorsal surfaces of the metanotum and abdomen.

Several groups of beetles which were represented by only a few specimens have been deleted from this part of the discussion.

In some ways it is easier to make negative statements concerning distributional patterns of mites on beetles. For example, several re­ gions were consistently free of mite infestations. Only a few instances of mites attached to the head were observed. One hypopus was taken from a compound eye of a specimen of Silpha americana; the other few instances involved hypopi attached to a relatively smooth expanse of the gulae of

Staphylinus maxillosus and Philonthus validus. As mentioned earlier in the results and discussion, hypopi were never encountered on interseg­ mental membranous areas. No hypopi, for example, were found on the cervix. The prosternum was often occupied by hypopi. The best example of a consistent pattern of hypopial attachment on the prosternum was pro­ vided by Staphylinus maxillosus in which all hypopi on the prosternum were encountered in the posterior furrow.

Other primary sites of attachment on the venter were pro- and mesocoxal cavities. With some beetle species, for example Nicrophorus tomentosus and N. orbicollis, hypopi occupied only the outer smooth sur­ faces of the mesocoxal cavities (Figures 7, 14 and 15). Other beetles with uniformly smooth coxal cavities had mites occurring in no particular pattern. It should be mentioned here that all of the beetles examined 45

in this study had metacoxae held so tightly against the cavities that no site for hypopial attachment was afforded.

Nicrophorus tomentosus and N. orbicollis were the only beetles col­ lected which had mites attached to the prothoracic spiracles. Most

Macrocheles sp. A adult were taken from N. orbicollis which has a more protected area resulting from more coverage by the spiracular lobe

(Figures 9, 10 and 11). A larger sample size would be needed to draw definitive conclusions, but it appears that the region of the prothoracic spiracles is one of the main sites for attachment of Macrocheles sp. A.

Only a few mites were encountered ventrally on regions posterior to the metasternum.

Legs were the primary sites of attachment for the uropodid deuto- nymphs. These mites were always restricted to a more exposed site, and were never encountered in protected regions such as under the elytra or in coxal cavities.

Like the intersegmental membranes, wings were not a site of attach­ ment for mites. Only one mite, Pyemotes sp. (Figure 26), was taken from the wing of Silpha americana. It, and the scutacarid from the carabid were

probably the only examples.of parasitic mites taken in this study.

The elytral concavities and the first three abdominal tergites har­ bored the majority of mites taken in this study. On some beetles such as Silpha noveboracensis and S- americana, hypopi attached both to the elytral surface and to the surfaces of the tergites.. Some beetles such as Geotrupes semiopacus, G- splendidus and Silpha surinamens is, which had 46

membranous or hairy surfaces, had few or no hypopi attached to the dorsum even when hundreds of hypopi were attached to the elytral concavities.

Future laboratory experiments are planned to test the movements of vitally stained hypopi to various regions. Three different patterns of distribution were found on the elytral concavities. Beetles with uni­ formly smooth elytral concavity surfaces such as Geotrupes semiopacus

G. splendidus (Figure 17), and Silpha surinamensis had hypopi attached to the midregions of the elytra. Silpha noveboracensis and americana which had similarly shaped elytra had hypopi restricted to the smooth, lateral, marginal edges of the elytra (Figures 2 and 4).

An unanswered question remains regarding Nicrophorus tomentosus and

N. orbicollis - that is, why was the region of the elytral concavity virtually free from mite populations? Of 44 beetles of N. tomentosus, only one beetle was found with a single specimen of Histiostoma sp. A attached to an elytral concavity. None of the 12 Nicrophorus orbicollis had mites present on the elytral concavities, although one beetle had high populations of hypopi on the dorsum of the abdomen and another had high populations on the metanotum.

An additional difficulty is encountered in this analysis when one is aware that the same species of hypopial mites occur on elytra of other silphids, staphylinids and geotrupids. Furthermore the same species of hypopi, e.g., Histiostoma spp. A and C (Figures 32 and 34) were found on other regions of these two species of beetles, e.g., mesocoxal cavities.

The abdominal tergites and notai surfaces of all the Nicrophorus 47

tomentosus were free of mite populations of any kind, yet the tergites

as shown in Figure 8 are smooth and rigid enough to provide a suitable

substrate for hypopial attachment. Two N. orbicollis had hypopial popu­

lations on dorsal surfaces under the elytra. Thirty-four Histiostoma sp.

A were found on the metanotum of one beetle, and 93 H. sp. C were taken

from abdominal tergite IV just posterior to the membranous concavity

formed by abdominal tergites I-III (Figures 12 and 13).

Macrocheles sp. A adultwere taken from the dorsal abdominal

concavity of 10 out of 12 Nicrophorus orbicollis specimens examined.

When we compared the abdominal dorsum of N. orbicollis with that of N.

tomentosus, which also harbored specimens of Macrocheles sp. A adult

on the prothoracic spiracles, the morphological difference is striking,

perhaps more so than between any other two beetle species involved in

this study. Figures 12 and 13 show the membranous concavity of the ab­

domen of Nicrophorus orbicollis and Figure 8 shows the fairly uniform

condition of all the abdominal tergites covered by the elytra. No

Macrocheles sp. A adultwere found on the dorsum of the abdomen of

Nicrophorus tomentosus. Laboratory experiments are being planned to

present a large population of Macrocheles sp. A adult ?? with the al­

ternative of selecting one or the other of the above two species of

beetles. Also regional selection needs to be studied by using dye-marked

mites.

Several observations concerning the regional distribution of hy­

popial mites can be made as a result of this study. With the exception of scattered individuals, large populations of hypopi were located in 48

regions which are thought to be free from the effects of self-grooming.

Also, large populations of h>popi were not found attached to integument

which was rough, membranous or densely setose, hairy or any combination

of these conditions as was demonstrated by the abdominal dorsum of

Geotrupes semiopacus, G. splendidus, Silpha surinamensis, and interior

surfaces of mesocoxal cavities of Nicrophorus tomentosus and N. orbicollis.

When populations of more than three hypopi occurred in a region a

bilateral distribution was evident. Figures 3, 5, 12, 13, and 19 are some examples of the many bilateral distributions of hypopi found in' this study.

Another phenomenon which is apparently associated with population size is that of heterogeneity. Smaller populations of hypopi (less than

30) were sometimes heterogeneous with two or more species of hypopi. The larger populations (30 and more) were homogeneous. This seems to indi­ cate that beetles with smaller heterogeneous populations accumulated the hypopi on several different encounters with small numbers of different species of hypopi. Larger homogeneous populations were likely accumulated as a result of a single encounter with a large homogeneous population of hypopi.

The relationship of the mites to beetles in this study appears to be entirely phoretic, with the exception of the single Pyemotes sp.

(Pyemotidae) and the one specimen of Imparipes sp. (Scutacaridae). No information is available concerning the symbiotic relationships of most of the types of mites found in this study. The most detailed information pertains to the symbiosis of Poeciiochirus necrophori. Many references 49

are made by European workers concerning the occurrence of P. necrophori on beetles. Most beetle hosts are "Suropea-i species of Nicrophorus and

Silpha (Th^odorid&s, 1950, 1955; Vitzthum, 1930; Cooreman, 1943, 1951; and Willmann, 1956). No records exist for Poecilochirus necrophori from staphylinids. The likelihood of the occurrence in this study being ac­ cidental seems great. With the exception of the one mite from Staphy- linus maxillosus, Poecilochirus necrophori was restricted to silphids.

I observed the mites moving about on the beetles and carrion in the field and made several observations in the laboratory. During preliminary studies I reared three Nicrophorus tomentosus in a finger bowl with dead mice. Two beetles died and the whole mite population, some 350 plus, joined the single living beetle. Figure 20 shows the beetle with this tremendous population of ambulatory deutonymphs. These particular mites were observed to feed on the dead mice and on carrion debris which clung to the surface of the beetle. If one considers the possible effects of accumulated dry carrion debris, the "cleaning" effect of poecilochirids could be interpreted as leaning toward mutualism whereby the beetle bene­ fits from the grooming and the mites benefit by having an available food supply. Although I have seen silphids groom their bodies I have never seen any action which could be interpreted as attempted mite removal.

Neumann (1943) made a study of the life history of Poecilochirus necrophori. He stated that most of the earlier life stages take place in the carrion and that the deutonymph is the longest stage of development

(15 to 20 days) with adult stages being very short lived. Prior to moult­ 50

ing, deutonymphs left the beetles. After moulting from deutonymph to

adult, the latter sought out places of concealment prior to mating and

oviposition. Neumann also observed the aforementioned "cleaning" effect

by poecilochirids.

The role of the Macrocheles sp. A adulton Nicrophorus tomentosus 1 and on N. orbicollis must be conjectured. Dr. Gerald Krantz has also

collected these from undetermined Nicrophorus sp.; no studies have been conducted to determine their symbiotic relationships. Only adult females were found, so it is possible that earlier life stages may be found on carrion. Evans and Hyatt (1963) found that many macrochelids symbiotic on coprophilous beetles spend their earlier stages in manure. They stated that the macrochelids are parthenogenetic and display a phoretic associa­ tion with beetles in general. I feel that further studies are needed in order to draw definitive conclusions as to their relationship.

Macrocheles sp. B adult were taken only from geotrupids. In contrast to M. sp. A, these mites were ambulatory and behaved similarly to poecilochirids, in fact, initially they were mistaken for poecilochirids.

Krantz and Mellott (1968) discussed a high degree of host specificity of three species of Macrocheles on three genera of geotrupid beetles from

Florida; however, neither Krantz and Mellott nor Evans and Hyatt furnished any detailed biological information regarding the symbiotic role of macro-

chelid mites or the habitat of immature stages. Krantz (1965) theorized that Macrochelidae appear to be a natural group evolving toward a parasitic existence in that many species are free living predators while others are

^Krantz, G. W., Department of Entomology, Oregon State University, Cor- vallis, Oregon 96331. Occurrence of macrochelids on beetles. Private com­ munication. 1968. 51

phoretic commensals or dependent on particular insect species in some other way.

Halolaelapid and laelapid deutonymphs were restricted to the sub- elytral abdominal concavities of histerid beetles; however, no definitive studies have been made on the mites associated with histerids. Steffan

(1967) reported that Stammer found some Laelapidae, living under elytra of some beetles, to be feeding on the hemolymph. I could detect no evi­ dence of this in my observations.

It is difficult to draw any firm conclusions concerning host specifici­ ty when dealing with such varied sample sizes. Some generalizations can be made, however. Uropodid deutonymphs were found on three families of beetles; Silphidae, Staphylinidae, and Scarabaeidae. It appears that the uropodids exhibit little host specificity. Uropodid deutonymphs have been taken from the teats of cows that have come in contact with these 1 mites.

Hypopi of the family Anoetidae were widely distributed on different hosts. Histiostoma sp. A (Figure 32) was collected in abundance from most of the silphids and from Staphylinus maxillosus. Histiostoma sp. B

(Figure 33), however, was taken only from staphylinids and only from the apices of the elytral concavities. Histiostoma sp. C (Figure 34) was common to most of the same host beetles as H. sp. A with the exception of Nicrophorus tomentosus which had hypopial populations of Histiostoma

1 Hicks, E. A., Department of Zoology and Entomology, Iowa State Uni­ versity, Ames, Iowa 50010. Phoresy of uropodid deutonymphs. Private com­ munication. 1965. 52

sp. A only. Histiostoma sp. D (Figure 35) was found only on Philonthus

validus and P. politus.

Hypopi of family Acaridae, with the exception of two Silpha americana

and one histerid Margarinotus hudsonicus, were found only on staphylinids

and geotrupids. Acarus siro was found only on a single Staphylinus

maxillosus. Caloglyphus sp. B was restricted to the elytral concavities

of Geotrupes semiopacus and G. splendidus.

Following is a brief review of literature dealing with some of the

more important studies of mite insect symbioses exclusive of ectopara-

sitism and similar to the relationships involved in this study.

Although no work has been published on distributional patterns of

mites on necrophilous beetles, there has been some investigation of the

mite fauna of passalid beetles, particularly Popilius disjunctus which is

known to have at least eleven species of mites associated with it. Bishop

(1964) found distributional patterns to be primarily on the venter of the

beetles with primary sites of attachment around the bases of the coxae.

Mollin and Hunter (1964), Hunter and Mollin (1964), and Hunter and Davis

(1965) also worked with Popilius dis.junctus and mite populations. Much

of their work dealt with behavioral observations and experiments with

regional selection. They found that Euzercon latus attached at the bases

of coxae by clinging with claws, whereas Cosmolaelaps passali used

chelicerae for attachment to setae just anterior to legs I on the pro-

sternum. Both mites were found to utilize other sites of attachment if the former sites were painted with nail polish. They observed that if the

polish wore off, or if mite-free beetles were introduced with no paint, 53

the mites would reoccupy the original site. Also if mite-free beetles without painted surfaces were added to a container with painted beetles, mites were found to move onto the "preferred" sites. Another experiment showed that the mites appeared to utilize some olfactory sense in finding proper beetle hosts. Two groups of mite-free beetles were introduced to a population of Cosmolaelaps passali. One group was washed in distilled water and dried. The other group was not treated. The mites attached to the unwashed beetles.

Th^dorides (1955) made an extensive survey of the parasites and com­ mensals of Coleoptera. Much of his work involved protozoans, fungi and . Although many listings of mites and hosts are given, he gener­ ally failed to give regional distributions for the attachment of mites.

He implied that hypopial occupation of subelytral spaces is a result of a rather constant temperature and humidity, a suggestion that I do not sup­ port. He further stated that he found mites on all surfaces of beetles including articulations of the cervix. This is definitely contrary to my findings.

Perkins (1899) reported a chamber or acarinarium on the basal ab­ dominal segments of certain xylocopid bees. Females of the genus Kop- torthosoma were consistently found to harbor a population of large acarid mites.

LeVeque (1930) used the fauna of mites in the acarinarium of xyloco- pids as an aid to separate species. She claimed in her studies that each species of bee in the genus Mesotrichia had a characteristic mite in the acarinarium regardless of geographic range and that very closely related 54

bee species sometimes had the same mite species in their acarinaria.

One of the excellent works on mite-insect symbiosis is that of

Krombein (1961). With the aid of wooden nest traps he was able to study the life histories of solitary wood-boring wasps and their mite symbionts.

Most of the vespid wasps studied had acarinaria similar to those found in female xylocopids. Like the carpenter bees, only female wasps possessed the acarinarium. Most vespids possessed saproglyphid hypopi in shingled rows in the acarinarium. Oviposition movements initiated movement of hypopi into the oviposition site where tritonymphal and adult stages were reached. Reproduction took place in the vespid brood cell. Hypopi moved from vespid pupal skins into the female wasp's acarinarium prior to the wasp's emergence.

The unusual membranous concavity formed by the first three abdominal tergites of Nicrophorus orbicollis may serve the function of an acarinari­ um. Further studies are being planned to test this hypothesis.

Donisthorpe (1927) showed that ants almost always had the same species of mites attached to the same body region. One of the important findings in his report was that of the uniform bilateral distribution of the attachments. Donisthorpe reported having seen ants die as a result of a complete covering of hypopial mites. He misinterpreted the role of the hypopus as one of an actively feeding parasite.

Another group of Hymenoptera, the neotropical army ants, was studied by Rettenmeyer (1961). Like Donisthorpe, he found mites attached to par­ ticular body parts. He considered a single ant as providing a group of 55

ecological habitats for mites. He found 50 different species of hypopial

forms on doryline ants.

Among the Diptera some interesting observations have been made on the hippoboscid flies of the genus Ornithomyia. Hill et al. (1967) made a distributional study of mites on these bird parasites and found

Tyrophagus mites on the thorax and abdomen, Myialges macdonaldi on the abdomen only, and Microlichus avus at the bases of the wings and pleura.

They observed that all the distributions were bilateral and in areas not affected by grooming.

It is apparent from this study that our understanding of mite-insect symbioses is meager. Many questions remain unanswered, but lend them­ selves to careful field and laboratory investigations. 56

CONCLUSIONS

1. Hypopi were not found to occupy the following morphological regions:

intersegmental membranes, dorsal surfaces of elytra, and wings.

2. The main sites of hypopial attachment were: prosternum, procoxal and

mesocoxal cavities, elytral concavities and abdominal tergites I-III.

3. The texture and sclerotization of the morphological regions were

found to influence the distribution of hypopial attachment.

4. Hypopi were not found on rugose, punctate, membranous, or densely

hairy surfaces.

5. Populations of four or more hypopi showed a definite bilateral dis­

tribution for particular morphological regions.

6. The main sites of hypopial attachment above appear to be free from

the effects of grooming behavior.

7- Pedicellate deutonymphs of Uropodidae were found to be attached mainly

to the legs and prosternum of beetles.

8. Macrocheles sp. A adult were attached by their chelicerae to setae

and membranous areas on Nicrophorus tomentosus, and N. orbicollis.

9. The main sites of attachment for Macrocheles sp. A adult were on

prothoracic spiracles under the spiracular lobe of the pronotum on

N. tomentosus and N. orbicollis, and in a membranous concavity of the

first three abdominal tergites of N. orbicollis.

10. Ambulatory Macrocheles sp. B adult were found only on Geotrupes

semiopacus and G. splendidus. 57

11. With the exception of what appeared to be an accidental occurrence

on Staphylinus maxillosus, Poecilochirus necrophori ambulatory deuto-

nymphs were found only on silphids.

12. Feeding poecilochirid mites cleaned accumulated carrion material from

the bodies of silphids.

13. Hypopi of Histiostoma spp. A and C were found on a wide variety of

host beetles and morphological regions. lU. Hypopi of H. sp. B were found to be restricted to the apical portions

of staphylinid elytral concavities.

15. Hypopi of H. sp. D were found only on the staphylinids, Philonthus

politus and P. validus.

16. Hypopi of Caloglyphus spp. were found mainly on staphylinids and

geotrupids. Caloglyphus sp- B was found only on the elytral concavi­

ties of Geotrupes semiopacus and G. splendidus.

17. Acarus siro hypopi were found only on Staphylinus maxillosus.

18. Populations of more than 30 hypopi were found to be homogeneous

whereas smaller populations often included two or more species.

19. With the exception of two mites representing parasitic families

(Pyemotidae and Scutacaridae), the symbiotic role of mites on necro­

philous beetles appears to be phoresy and phoretic commensalism. 58

LITERATURE CITED

Arnett, R. H. 1944. A revision of the Nearactic and Necro- phorini based upon female genitalia. New York Entomological Society Journal 52:1-25.

1946. Coleoptera notes; I. Silphidae. Canadian Entomologist 78:131-134.

1947. Coleoptera notes: II. Silphidae. Canadian Entomologist 79:110-113.

. 1968. The beetles of the United States (a manual for identifi­ cation). The American Entomological Institute, Ann Arbor, Michigan.

Baker, E. W., J. H. Camin, F. Cunliffe, T. A. Wooley, and C. E. Yunker. 1958. Guide to the families of mites. University of Maryland, Insti­ tute of Acarology, Contribution No. 3.

and G. W. Wharton. 1952. An introduction to acarology. Macmillan, New York.

Bishop, M- R. 1964. A study of some mites symbiotic on Popilius dis.junctus Iliiger. Unpublished M.S. thesis. Library, Iowa State University of Science and Technology, Ames, Iowa.

Canestrini, R. 1882. Contribuzione alio studio degli parassiti degli insetti. Atti del la Societa'veneto-Trentina di Scienze Naturali, Padova 7(2):154-178.

Casey, T. L. 1900. A review of the genus Dermestes (Dermestidae: Coleoptera). New York Entomological Society Journal 8:138-165.

Cooreman, J. 1943. Note sur la faune des Hautes-Fagnes en Belgique. XI. Acariens (). Bulletin du Musee royal d'histoire naturelle de Belgique 19(63):1-28.

. 1951. Etudes biospeologiques. XXXIV. Acariens de Transylvanie. Bulletin du Institut royal des Sciences naturelles de Belgique 27(42): 1-15.

Crowson, R. A. 1955. The natural classification of the families of Coleoptera. Nathaniel Lloyd and Co., Ltd., London.

Dillon, E. S. and L. S. Dillon. 1961. A manual of common beetles of Eastern North America. Row, Peterson and Co., Evanston, Illinois.

Donisthorpe, H. 1927. The guests of British ants, their habits and life histories. George Routledge and Sons, London. 59

Evans, G. 0. 1957. An introduction to the British Mesostigmata (Acarina) with keys to families and genera. Linnean Society of London Journal U3(291):203-259.

and K. H. Hyatt. 1963. Mites of the genus Macrocheles Latr. (Mesostigmata) associated with coprid beetles in the collection of the British Museum. Bulletin British Museum of Natural History 9(9):327-401.

, J. G. Sheals and D. MacFarlane. 1961. The terrestrial Acari of the British Isles. Volume I. Introduction and Biology. British Museum, London.

Fabricius, J. C. 1805. Systema antliatorum secundum ordines, genera, species adiectis synonymis, locis, observâtionibus, descriptionibus. Carolum Reichard, Brunsvigae.

Grandjean, F. 1932. Au sujet des Palaeacariformes Tragardh. Museum National d'histoire naturelle Paris Bulletin 4(2):411-426.

. 1935. Observations sur les Acarines s^ie). Museum National d'histoire naturelle Paris Bulletin 7(2):119-126.

. 1954. Etude sur les Palaeacaro'i'des (Acariens, Oribates). Museum National d'histoire naturelle Paris M^oires 7A(3):179-274.

Griffiths, D. A. 1964. A revision of the genus Acarus L., 1758 (Acaridae Acarina). The British Museum (Natural History), Zoology, London Bulletin Volume II, No. 6.

Hatch, M. H. and A. I. Ortenburger. 1930. New records of Coleoptera from Oklahoma and Western Arkansas. University of Oklahoma Biological Survey 2(1):7-14.

Hill, D. S., N. Wilson and G. B. Corbet. 1967. Mites associated with British species of Ornithomyia (Diptera: Hippoboscidae). Journal of Medical Entomology 4(2):102-122.

Howden, H. F. 1955. Biology and taxonomy of North American beetles of the subfamily with revisions of the genera Bolbocerosoma, Eucanthus, Geotrupes and Peltotrupes (Scarabaeidae). The United States National Museum, Smithsonian Institution Proceedings 104(3342) 151-319.

. 1964. The Geotrupinae of North and Central America. Entomolog cal Society of Canada Memoirs No. 39.

and 0. L. Cartwright. 1963. Scarab beetles of the genus Ontho- pagus Latreille north of Mexico (Coleoptera: Scarabaeidae). The United States National Museum, Smithsonian Institution Proceedings ll4(3467):l-333. 60

Hughes, A. M. 1961. The mites of stored food. [Great Britain] Ministry of Agriculture, Fisheries and Food Technical Bulletin No. 9.

and C. 6. Jackson. 1958. A review of the Anoetidae (Acari). The Virginia Journal of Science 9(1):5-198.

Hunter, P. E. and R. Davis. 1965. Mites associated with the passalus bettle. III. Life stages and observations on the biology of Euzercon latus (Banks) (Acarina: Euzerconidae). Acarologia 7(1): 30-42.

and K. Mollin. 1964. Mites associated with the passalus beetle. I. Life stages and seasonal abundance of Cosmolaelaps passali n. sp. (Acarina: Laelaptidae). Acarologia 6(2):247-256.

Jaques, H. E. 1951. How to know the beetles. Wm. C. Brown Co. Publishers, Dubuque, Iowa.

Karafiat, H. 1959. ^Systematik und Okologie der Scutacariden. In H.-J. Stammer. Beitrage zur Systematik und Okologie mitteleurop'aischer Acarina. Band 1, Teil 2. Pp. 627-712. Akademische Verlagsgesellschaft, Geest und Portig, K.-G. Leipzig.

Krantz, G. W. 1962. A review of the genera of the family Macrochelidae Vitzthum (1930) (Acarina: Mesostigmata). Acarologia 4(2):143-173.

. 1965. A review of the genus Neopodocinum Oudemans 1902 (Acarina: Macrochelidae). Acarologia 7(2):139-226.

and J. L. Mellott. 1968. TWo new species of Macrocheles (Acarina; Macrochelidae) from Florida, with notes on their host- specific relationships with geotrupine beetles (Scarabaeidae: Geotru- pinae). Kansas Entomological Society Journal 41(l):48-49.

Krczal, H. 1959. Systematik und Okologie der Pyemotiden. In H.-J. Stammer, Beitrage zur Systematik und Okologie mitteleuropaischer Acarina. Band 1, Teil 2. Pp. 387-625. Akademische Verlagsgesell­ schaft, Geest und Portig, K.-G. Leipzig.

Krombein, K. V. 1961. Some symbiotic relations between saproglyphid mites and solitary vespid wasps. Washington Academy of Sciences Journal 51(6):89-93.

Leonardi, G. 1900. Storia naturale degli^ acari insetticoli. Societa^ Entomologica Italiana Bollettino 32:1-76.

LeVeque, N. 1930. Symbiotic mites used to separate species of a genus of bees. Science 71:607-608. 61

/ Linné, Cari von. 1736. Animalia per Sveciam Observata. Acta literaria et scientiarum Sveciae.

. 1746. Fauna Svecica. Stockholmiae Laurentii Salvii.

. 1758. Caroli Linnaei Systema Naturae. Regnum animale. Editio décima. Hoimiae, impensis direct, Laurentii Salvii.

Michael, A. D. 1901. British Tyroglyphidae. Volume I. Adlard and Son, London.

. 1903. British Tyroglyphidae. Volume II. Adlard and Son, London.

Mollin, K. and P. E. Hunter. 1964. Mites associated with the passalus beetle. II. Biological studies of Cosmolaelaps passali Hunter and Mollin (Acarina:—Laelaptidae). Acarologia 6(3);421-431.

Nesbitt, H. H. J. 1945. A revision of the family Acaridae (Tyroglyphidae), order Acari, based on comparative morphological studies. Canadian Journal of Research [Section] D, 23:139-188.

Neumann, K. W. 1943. Die Lebengeschichte der Kafermilbe Poecilochirus necrophori Vitzthum nebst Berschriebung aller Entwicklungsstuffen. Zoologischer Anzeiger 142:1-21.

Gudemans, A. C. 1903. New list of Dutch acari. Second part. Tijd- schrift voor Entomologie 45:1-52.

Perkins, R. C. L. 1899. On a special acarid chamber formed within the basal abdominal segment of bees of the genus Koptorthosoma (Xylo- copinae). Entomologist's Monthly Magazine 35:37-39.

Rettenmeyer, C. W. 1961. Behavior, abundance and host specificity of mites found on neotropical army ants. International Congress of Entomology, Vienna, Proceedings 11th, Vol. 1:610-613.

Schaupp, P. G. 1881. Description of the larva of Necrophorus tomentosus Weber. Brooklyn Entomological Society Bulletin 4(7-8):37-38.

Scheucher, R. 1957. Systematik und Okologie der, deutchen Anoetinen. In H.-J.Stammer. Beitrage zur Systematik und Okologie mitteleuropaischer Acarina. Band 1, Teil 1. Pp. 233-384. Akademische Verlagsgesll- schaft, Geest und Portig, K.-6. Leipzig.

Steffan, A. W. 1967. Ectosymbiosis in aquatic insects. In Henry, S. M. ed. Symbiosis. Volume 2. Pp. 207-289. Academic Press, New York. 62

Strandtmann, R. W. and G. W. Wharton. 1958. A manual of mesostigmatid mites parasitic on . University of Maryland, Institute of Acarology, Contribution No. 4.

Theodorides, J. 1950. Notes diverses sur les Necrophorus (Coleoptera: Silphidae). Institut royal des Sciences naturelle de Belgique Bulle­ tin 26(52):l-20.

. 1955. Contribution a I'etude des parasites et phoretiques de coleopteres terrestres. Actualités Scientifiques et Industrielles No. 1217.

Turk, E. and F. Turk. 1957. Systematik und Okologie der Tyroglyphiden Mitteleuropas. In H.-J. Stammer. Beitrage zur Systematik und Okologie mitteleuropaischer Acarina. Band 1, Teil 1. Akademische Verlagsgesellschaft, Geest und Portig, K.-G. Leipzig.

Vitzthum, H. 1930. Milben als Pesttrager? Ein Beitrag zu den Unter- suchungen der mandschurischen Peststudienkommission in Harbin. Zoologische Jahrbucher, Abteilung fur Systematik, Okologie und Geographie der Tiere 60(3-4):381-^•28.

Williams, B. S. 1928. Helophorus nanus Sturm in Bedordshire. Ento­ mologist's Monthly Magazine 64:234.

Willmann, C. 1956. Milben ^s dem Natur^chutzgebiet auf dem Spieglitzer (Glatzer) Schneeberg. Ceskoslovenska Parasitologic 3:211-273.

Zakhvatkin, A. A. 1941. Fauna U.S.S.R. Arachnoidea vi (1) Tyroglyphoidea (Acari). Zoological Institute of the Academy of Science of the U.S.S.R., new series No. 28. Original available (in Russian); translated to English by Ratcliffe, A. and A. M. Hughes in 1959. American Insti­ tute of Biological Science, Washington, D.C., U.S.A. 63

ACKNOWLEDGMENTS

I wish to express my sincere gratitude to my major professor. Dr.

Ellis A. Hicks who has been a teacher, counselor, and friend during this

investigation. He has given his time and energy untiringly for counsel­

ing, discussing research plans, and editing manuscripts.

Appreciation is extended to Dr. Rupert Wenzel, Chicago Museum of

Natural History, for his determinations of several histerid beetles, and to Mr. William V. Miller of Ramsey, New Jersey, for his determinations of many staphylinids. Mr. Kenneth Esau of Iowa State University was helpful

in identification of the carabids. Dr. Preston E. Hunter, University of

Georgia, and Dr. Gerald Krantz, Oregon State University, were helpful in the determinations of several mesostigmatid mites.

The greatest appreciation is reserved for my wife who has assumed more than her share of family responsibilities in order that I might com­ plete this study. I thank her for her enduring patience. 64

APPENDIX A: PHOTOGRAPHS OF BEETLE MORPHOLOGICAL REGIONS Plate 1

Figure I. Saprinus assimilis dorsum of abdomen with wings and elytra removed. 16X

Figure 2. Elytral concavity of Silpha noveboracensis showing smooth lateral surface and rough medial surface. lOX

Figure 3. Abdominal tergites of Silpha noveboracensis showing bilateral distribution of hypopi on tergites I-III. 7X

Figure 4. Elytral concavity of Silpha americana showing hypopi attached to the smooth lateral margin of the elytron. 8X 66 Plate 2

Figure 5. Abdominal tergites of Silpha americana showing the bilateral distribution of hypopi on tergites I-III. Sx

Figure 6. The membranous dorsum of the abdomen of Silpha surinamensis. 8X

Figure 7. MesocoJfial cavity of Nicrophorus tomentosus showing the outer smooth surface and inner roughened surface. 40X

Plate 3

Figure 8. Abdominal tergites of Nicrophorus tomentosus showing the sclerotized condition of the tergites. lOX

Figure 9. Prothoracic spiracle and spiracular lobe of Nicrophorus tomentosus. 25X

Figure 10. Prothoracic spiracle and spiracular lobe of Nicrophorus orbicollis showing the partially covered spiracle. 25X

Figure 11. Four Macrocheles sp. A adult attached to a prothoracic spiracle of Nicrophorus orbicollis. 25X

Plate 4

Figure 12. Abdominal tergites of Nicrophorus orbicollis with 145 Macrocheles sp. A adult attached bilaterally to tergites I-III, and 93 Histiostoma sp. C hypopi attached bilaterally on tergite IV. 7X

Figure 13. Abdominal tergites of Nicrophorus orbicollis with specimens of Macrocheles sp. A adult removed to show the membranous concavity of abdominal tergites I-III. 7X

Figure 14. Mesocoxal cavities of Nicrophorus orbicollis showing 23 Histiostoma sp. C hypopi attached to the smooth medial area of the left coxal cavity. 15X

Figure 15. Mesocoxal cavities of Nicrophorus orbicollis with mites and coxa removed to show contrasting surface textures. 2OX 72 Plate 5

Figure 16. Prosternum of Staphylinus maxillosus showing two Histiostoma sp. C hypopi attached in the posterior furrow. 3OX

Figure 17. Three-hundred and fifteen Caloglyphus sp. B attached to the elytral concavity of Geotrupes splendidus. 8X

Figure 18. The abdominal tergites of Geotrupes splendidus showing the hairy, membranous texture of the integument. 2OX

Plate 6

Figure 19. Dorsum of abdomen of Dermestes vulpinus showing bilateral distribution of saproglyphid hypopi on tergites I-III. lOX

Figure 20. Nicrophorus tomentosus with Poecilochirus necrophori ambula­ tory deutonymphs. 3.5X 76 77

APPENDIX B: PHOTOMICROGRAPHS OF MITES NOT IDENTIFIED

TO SPECIES Plate 7

Figure 21. Imparipes sp. (Scutacaridae) ex Harpalus pennsylvanicus. 450X

Figure 22. Unidentified Laelapidae adult. lOOX

Figure 23. Unidentified Halolaelapidae deutonymph. lOOX

Figure 24. Unidentified mesostigmatid deutonymph. lOOX

Plate 8

Figure 25. Unidentified mesostigmatid deutonymph. lOOX

Figure 26. Pyemotes sp. ex wing of Silpha americana. 225X

Figure 27. Uropodid pedicellate phoretic deutonymph. 125X

Plate 9

Figure 28. Macrocheles sp. A adult ^ . lOOX

Figure 29. Macrocheles sp. B adult ^ . lOOX

Figure 30. Alliphis sp. (Eviphididae) adult ^ . 150X

Figure 31. Saproglyphid hypopus. 200X 83 Plate 10

Figure 32. Histiostoma sp. A hypopus. 150X

Figure 33. Histiostoma sp. B hypopus. 250X

Figure 34. Histiostoma sp. C hypopus. 200X

Plate 11

Figure 35. Histiostoma sp. D hypopus. 200X

Figure 36. Histiostoma sp. E hypopus. 250X 87 Plate 12

Figure 37. Caloglyphus sp. A hypopus. 225X

Figure 38. Caloglyphus sp. B hypopus. 200X

Figure 39. Caloglyphus sp. C hypopus. 225X

Figure 40. Caloglyphus sp. D hypopus. 300X mim 90

APPENDIX C: ABBREVIATIONS USED

Cg. A, B etc. Caloglyphus sp. A, sp. B etc. hypopi

Hst. A, B etc. Histiostoma sp. A, sp. B etc. hypopi

Hyp, asp. vitz. Hypoaspis (Laelapsis) Vitzthumi

Me. A, B Macrocheles sp. A, sp. B

Mes.stg. Unidentified mesostigmatid

Poec. nec. Poecilochirus necrophori

Pyin. Pyemotes sp.

Sap. Saproglyphid hypopi

Urp. Uropodid deutonymph

DN Deutonumph

^ Adult female

Ai,., Ag etc. Abdominal segment 4, 5 etc.

Spc;l. Prothoracic spiracle

XF External free

L Left

R Right

Mid Middle

= Approximately equal bilateral distribution

J. F. The Thomas Judge Farm collection site

An. Sc. F. Iowa State University Animal Science Farm collection site 91

APPENDIX D: REGIONAL DISTRIBUTION OF MITE SYMBIONTS ON HOST BEETLES Table 8. Regional distribution of mite symbionts on host beetles

BODY VENTER 1 1 1 III i apex IV excluding dorsal to coxae Abdominal stemite* Abdominal stemite* Metapleuron Metacoxol cavities dorsol to coxae Metosternum

HOST BEETLE Procoxal Mesoeoxal cavities Propleuron Abdominal stemite Mesosternum Prosternum Mesopleuron cavities Head Cervix karabidae Harpalus geims'^ivanicus Anisodactvius lAnisoaactvius) 1 bee :le aericoia 0 mit iS 67-21:2 Histeridae 4: be sties Saprinus tes lies assimilis m: on 6 beel

67-9:5

67-27:3

67-31:3

67-54:1

67-35:5

67-25:7

Saprinus 2 bee- les lugens mi tes on 1 beet le

67-20:9

Margarinotus 1 beei le 1 hudsonicus 0 mit( s 67-3:3 R Hister 1 beel le abbreviatus 0 mit(

Hister 19 bee ties circinans mi tes on 2 beet les

67-42:2

67-46:6 1

! ! 1 1 1 1 ! ! Head

Cervix .

Pronotum Y D O B M

Mesonotum U S R O

Elytral D concavity

Metonotum

Ventral to scutellum

Wings

Abdominal 4' si tergites III" lO l- f 11i Ï-- t w

Abdominal tergites IV-apex

-

r = m g

1

•n X m g l-J o 00 o c 5 &

1 . YDOBRE TNEV 1 sternite» IV apex sternite* 1 III excluding dorsal to coxae Abdominal T SOEHL TEEB Abdominal sternite 1 dorsal to coxae Abdominol cavities cavities Metasternum MetapUuron Metocoxal Procoxal Mesocoxal Mesosternum Proplouron cavities Mesopleuron Prosternum Cervix Head e adihpliS1 w Ji" ee b ft g ^ X) r N5 n> o h" 0\ ^ l;* 1 sB e: ee b 03 w M (5 ru en o 03 2" n r- p 1

3: 01- 56

4: 01- 56

3: 21- 56 h r lo R.

5: 41- 56 N PJ 4: 51- 56

4: 91- 56

5: 91- 56

6: 91- 56

4: 1- 66 p ON ON

5: 2- 66 i

6: 2- 66 i\> lo 7: 2- 66 M7V»> ON ON 00

9: 2- 66

3: 1- 76 "R 1 4=" G\

5: 1- 76

6: 1- 76

7: 1- 76

8: 1- 76 1

Head

Cervix

r , 1 Pronotum Y D O B M

Mesonotum U S

r^Vw R

a ^ O

1 îi D Elytral n "i II1 concavity 1 1 # 'i 0

Metanotum

Ventral to scutellum Wings U) NVi Cï| of N Abdominal ""g 4; "11$ tergites n (n "1 fw. i-in "I "i "1 VO «u Ln Abdominal tergites iV-opex

-

C = m s

£fe = f f% F -n X f" T m ûo I—» H- n- C (0 O. o* g ? 3 3

1 , YDOBRE T NE, V 1 termites l-lll sternites Abdominal s excluding dorsal to coxae Abdominal IV-apex

T SOEHL TEEB sternite 1 dorsal to coxae Abdominal cavities cavities Metapleuron Metacoxol Mesocoxal Metasternum Procoxal cavities Mesosternum Mesopleuron Prosternum Pfopleuron Head Cervix ah oliS s necaroDevon s i • )d eunitnoc(

1: 01-76

3 :61-76

4 :61-76

7 :81-76 t

8 :81-76

/ / 5597 In

9 :81-76 7

0 1:81-76 i 1 1:81-76

2 1:81-76 ro "s 1 o\ 00 w VJ h-» •-» '1 1 f ON 00 1—• 1—•

1 :91-76 ="1 1 o\ vo "sj hj

1: 02-76 ro 1 o> N3 O vj N3

3: 02-76

4: 02-76

2 :32-76

2 :62-76

3 :62-76 i 1 :72-76 1

Head

Cervix ,

Pronofum Y D O B M

Mesonotum U S

TV) R O

Elytral D ^1 "1 "1 4 concavity

Metonotum

Ventral to scutellum Wings win >4 N XvJ N Z-N Ï» Uj f- Abdominal »ii| >jffi tergite* î| ^1 "1 :i# "1 # "1sP Ix -If "1 Mil

Abdominal % tergite* IV-apex

N \

-

= r am

—

|>

4 Tt X f 4'T i" f. f I—• n> 00 h" o c g o s &

1 . YDOBRE T NEV 1 Abdominal sternites Abdominal T SOEHL TEEB sternites 1 III excluding dorsal to coxae IV-apex sternite 1 dorsal to coxae Abdominal cavities cavities Metasternum Metapleuron Metocoxol cavities Mesoeoxol Propleuron Procoxol Mesopleuron Masosternum Prostarnum Hood Cervix

a hpliS to o\ CO (D cr a nacirema jr- H" W g rt (D m (D rt H-" cr (D

7 :21-56 CO Ln I—* Ln o

1 :71-56

2 :91-56

3 :02-56

4: 02-56

0 1:12-56

1 1:12-56

3 1:12-56

01 :22-56

41 :22-56

2 :32-56

2: 42-56

3 :42-56

4: 52-56

2 :82-56 \ 0\ Ln I—» •—»

2 :1-66

3 :1-66

1: 2-66 2>^- 1: 6-66 Head

Cervix

Pronotum

09 Mesonotum o o -< l! Elytral o o concavity i8 Metanotum

Ventral to scutellum Wings

Abdominal ^1 lïfl""

Abdominal tergites IV-opex

f- IX»

?? -

= f- am

1 n A Jf 'g~ 4" 4^ f -n X r 'If 4" f f {. 00 M' t-' (0 n o § 5 6 î

1 - , YDOBRE T NEV 1 Abdominal sternites IV apex Abdominal sternites l-lll excluding dorsal to coxae stemite 1 dorsal to coxae

T SOEHL TEEB Abdominal cavities Metapleuron Metacoxal cavities Metasternum Proeoxal cavities Mesoeoxal Propleuron Mesosternum Mesopleuron Presternum Head Cervix ' a hpliS a nacLcema ) deunitnoc(

1 :6-76

2 :6-76

1: 9-76 N. "g

2 :9-76

3 :41-76

4: 41-76 i

3: 51-76

4 :51-76

5 :51-76

1: 81-76

3 :91-76 i

4: 91-76

5 :02-76

1 :22-76

1: 32-76

1 :82-76

2 :92-76

3 :92-76 N

4 :92-76

1 :03-75 1 :24-76 1 Head

Cervix ,

Pronotum Y D O B M

Mesonotum U S N R N Vll"»* O # A ^ r- ^ Elytral D concavity 4 p "# "1 "1i

Metanotum

Ventral to scutellum

Wings w i è| t "II,s '1 i ISA"*'Abdominal

Abdominal tergites IV-apex

-

= r- mg it H — 1 sL J §k y •n X i" •t*u 1- i" f i' !•" 4 f -JLO__ # M' M' cr 00 s? (t> s w 0 1

1 , YDOBRE T NE, V 1 sternites Abdominal Abdominal IV-opex sternites 1 III excluding dorsal to coxae

T SOEHL TEEB •temite 1 dorsal to coxae Abdominal cavities cavities Metapleuron Metacoxal Mesocoxal Procoxal Mesopleiiron Propleuron cavities Mesosternum Head Prosternum Cervix • a hpliS

w is ee b ltî si snemanirus CO B A fn » se t n o 1 h-"

1 :61-56

2: 61-56

3 :91-56

71 :22-56

91 :22-56

1: 42-66

2: 7-76

3: 7-76

4: 7-76

1: 41-76 1 1: 61-76

2 :81-76

6 :62-76

7 :72-76

3 :03-76

9 :73-76

a hpliS ie CA eb se l

a cinoppal W' O* g O B s et n o 1 h-»

4: 21-56 1 Head

Cervix ,

Pronotom Y

îi '1 D O B M

Mesonotum U S M N R Rg O i Elytral D "i " i i "R concavity

Metanotum

Ventral to ' scutellum

Wingt M Abdominal teijijitei o U) Abdominal tergites IV-apex

- R

= i

h 'X — K;

-n X i- r f 4" 4i i Table 8. (Continued)

BODY VENTER 1

E E 3 3 uu 1 1 X « M *> o # Metapleuron cavities Mataeoxal Procoxal Masocoxal cavities

HOST BEETLE Proplauron Mesopleuron Protternum covities lilii Head U J 1 II!

Nicrophorus 5 , be it les tomentosus m tes on 4 iv be it It S

i-Hi A A. 65-12:1 é / Wf# n:é

65-12:2

65-14:3

65-23:1

65-23:17

65-24:5

/Urp. 65-26:1 Nié

65-27:2

/mé. /mJL 66-2:10 A H • 40UtJi 66-2:11 tué

66-13:1 3/ff^A 9MA 66-13:2 i* IL SU 3imi 66-14:1

66-14:10

lUtf. 66-16:2 mé

66-18:2

66-18:3

66-18:5

66-18:6 47iUA.

66-18:7

66-18:8 1

Head M U S

Cervix R O D ,

Pronotum Y D O B Masonotum

V. Elytral concavity

Metanotum

Ventral to scutellum Wings

Abdominal tergite» 1-111 o U1 Abdominal tergite* IV-opex

—

r = m g

=

(•- I- 'g «1- Tl X i 4' 4" 4"t 4" »-» p 00 cr n o n* a> g o. (D

I RE T NEV. YDOB sternites IV apex Abdominal Abdominal sternites 1 III excluding dorsal to coxae

T SOEHL TEEB stemite 1 dorsal to coxae Abdominal Metapleuron cavities cavities Metasternum Metacoxal Mesocoxal cavities Pfocoxal Mesosternum Mesopleuron Proplauron Prosternum Cervix Haad s urohporciN s usotnemot ) deunitnoc( 9: 81-66

0 1:81-66 ro

1: 91-66 1 "i

1 :02-66

1: 12-66 i

2 :12-66 3 :12-66

4 :12-66

4: 22-66 "1 If

3 :42-66 ll|

1: 52-66 ^ï| 2 :52-66 "1 3 :52-66

4: 52-66 "1 5 :52-66

6: 52-66

4: 82-66 "i

1 :92-66 i 6 :81-76

3: 32-76

4: 32-76 1 ON ON to VI 5 :52-76 "1 1

Head

Cervix ,

Pronofum Y D O B M

Mesonotum U S R O

Elytral D concavity

Metanotum

Ventral to scutellum Wings

Abdominal tergites Mil ovj Abdominal tergites iV-apex

-

V - m g

-

f 1^ % t t, & li' r 4 % NO R 1 -n X f ? f î' r1' f r f 1" i- f 'r" 1^ 00 <0 g cr o g s o & < < < < < < -< -< -< -< -< -< -< -< z z z z z z CD CD CD CD CD CD CD CD H H H H H H O O O O O O O O m m m m m m rti rti rti rti rti rti o o o o o o o o 70 70 70 70 70 70 nol Abdominal Abdominal sternites IV-opex sternites l-lll excluding dorsal to coxoe

T SOEHL TEEB stemite 1 dorsal to coxae Abdomi cavities Metapleuron cavities Metocoxol Metosternum Mesocoxol cavities Procoxal Mesosternum Mesopleuron Prosternum Propleuron Head Cervix

su rohporc1i N W IAJ t" cr (D (0 s illocibro ^ fl> M. H-» M g cr 0 H' S rr rf O

1 :52-56 2 :52-56

3 :52-56 Vu •to g

1: 61-66 IV, v.

2: 22-66 "to B 3: 22-66 •fQ ^

1: 62-66

1 :33-66 • 1: 12-76 S "1

6 :82-76

7 :82-76

1 :13-76 n> CO M- y m h-

s urohporciN m fl) (U oj o h-» S i-* o* P h" p s utalutsup 5 u rr k

1 :01-56

41 :12-56

4: 42-56 a teahconoirP O' (0 (5 (0 i-" t-* 1+ a capo tn o m B H" rt 1

Haad M U S

Cervix R O D ,

=>"1 Pronotom Y D O

4 B Mesonotum

Elytral concavity w

Mefanotum "i Ventral to scutellum Wings

W Abdominal ^1 «1 .i| 4g tergites K» to •w 1o t •K> i-in o 4 4o 40 vo Abdominal Ml| tergites IV-apex

-

r = m s

• =

t.? si® st |R 1 a -n X R 4 1' 4' |n 1? 4' H-» (D 00 rt s? n ë cr § &

1 , YDOBRE T NE, V 1 nal sternites IV apex Abdominal Abdominal sternites l-lll excluding dorsal to coxae stemite 1 dorsal to coxae T SOEHL TEEB Abdomi cavities Metopleuron cavities Metasternum Metacoxal Mesocoxal cavities Procoxal Mesosternum Mesopleuron Propleuron Prosternum Haad Cervix 1 .j yitpatS ea aini O i-" et" ^ su nilyhpatS 1 4=- (A n) (D w D* B rt su sollixam B {+

1 :41-56

1: 91-56

1 :12-56 il l FN) 5 :12-56 kg 8: 12-56

6: 22-56 1 K5 ON Ln (jO h-» O

2 1:42-56

3: 5-66

5 :82-66

5 :03-66

3 :5-76

5 :41-76 n *> 1: 51-76 1 1 ?»

5* 61-76. |n 1 ON ON o> I-»

7 :61-76

9 :61-76 W

0 1:61-76 iig ! 6 :02-76 7 :02-76 1

Head

Cervix ,

Pronatum Y D O B M

Mesonotcm U S

^ 3 R O

A, 1 Elytral D i concavity

till n Metanotum

Ventral to scutellum ; Wing*

Abdominal teraltes i-in

Abdominal tergitas IV-opex

-

n = m s

=

-n X i s ) u d n s e 4 0 1 2 i u u 2 8 4 1 5 3 2 1 1 9 1 8 1 5 1 9 8 7 5 6 l s : n : : : : : : : : : : : : : : : : : : : y o 3 i 7 2 7 6 7 5 9 7 7 7 7 7 6 3 3 0 3 3 3 h l 4 t 2 4 3 3 3 2 2 2 3 2 2 2 2 2 2 2 2 2 2 o T l - - - - n ------a S i 7 7 7 7 7 7 7 o 7 7 7 7 7 7 7 7 7 7 7 7 7 t x O E 6 6 6 6 6 6 6 6 6 6 C 6 6 6 6 6 6 6 6 6 6 S a H L

o\ ( m VJ T t E E

£ B 1 Head

Cervix i I"! tk n n n Prosternum

Propleurofi

Procoxal civities

Mesosternum . Y D

Mosopleuron O B R E

"1 Masocoxal h: p P P n cavities N E V *"1 Metasternum Metapleuron Metacoxal cavities Abdominol sternite 1 dorsal to coxae V Abdominal sternites l-lll excluding dorsal to coxae .

Abdominal sternites IV-opex 1 1 1 Head

Cervix ,

Pronotum Y D O B M

Mesonotum U S

N yu CM s (o R O

r* K Elytral D K r- p ''I concavity i 4 "1 v.

Metanotum

Ventral to scutellum Wings

Abdominal tergites III"

Abdominal tergites IV-apex

-•

1" = m s

=

-n X Table 8. (Continued)

BODY VENTER l-lll

tJ 3 IV-opex sternites excluding dorsal to coxae Abdominal sternites Abdominal sternite 1 dorsal to coxoe Metapleuron

HOST BEETLE Metacoxal cavities Procoxal Mesoeoxal cavities Metasternum Prosternum Propleuron Mesotternum Mesopleuron Abdominal cavities X Cervix

Staphylinus 2 beel les maculosus 0 mit( s

Philonthus 3 beel les politus mi tes on 2 beet les

Z^SÉlÛ. 67-2:1

67-44:2

Philonthus 7 beet les validus mi tes on 3 beet les

66-2:12

iâs SiùàL. 67-17:1 Mk & »tka fOUsLB. 67-25:1 6L Philonthus 1 beel le cyanipennis 0 mite s

Ontholestes 20 be« ties cingulatus mi tes on 2 beet les

67-3:1

67-3:2

Trogidae 2 beet les Trox 0 mite s Unistriatus Trox 2 beet les suberosus 0 oitÉ s 1

Head

Cervix .

Pronotom Y D O B M

Mesonotum U S

en Qo R O

Elytra 1 D "il "il concavity

Metanotum I- ^ Ventral to scutellum Wings

Abdominal tergites M11 U1 Abdominal tergites IV-apex

-

> S G E L = . o x o C —

•n X Table 8. (Continued)

BODY, VENTER l-lll not X

> apex 2 • IV excluding dorsal to coxae Abdominal sternites Abdominal sternites sternite 1 dorsal to coxae HOST BEETLE Metapleuron Metasternum Metacoxol cavities Mesocoxal cavities Procoxai Abdomi Mesopteuron Prosternum Propleuron Mesosternum z U cavities Geotrupidae 2] be( Geotrupes ties semiooacus mi tes on 3 beet les

66-11:2

66-11:3 «#£t£[ 2&ja 66-27:1 ff-j n:J

Geotrupes 15 bet ties splendidus mi tes on 9 beet les

65-24:13 açjx R L 66-8:1

66-9:1 /Cj-A R z&c 7&Ç 66-22:1 H!J Mli in.,4.L

66-30:1

66-30:2

66-30:6

66-32:1

66-5:2

Scarabaeidae 5 jeet les Onthophagus 0 nite s hectate Onthophagus 1 jeet le janus 0 nite s

Copris 1 jeet le tullius 4 nite s

66-33:2 3Utf, n:é 1

Head M U S

Cervix R O D ,

Pronotum Y D O B Mesonotum /V) KO s Elytra 1 g i 'i "g "1 concavity

Metanotum

Ventral to scutellum 1 Wings

Abdominal tergites Mil

Abdominal tergites IV-opex

-

= r ; sm # KK =

-n X !l Table 8. (Continued)

BODY VENTER j l-IM '

TJ apex S ' IV

HOST BEETLE excluding dorsal to coxae Abdominal sternites Abdominal sternites sternite 1 dorsal to coxae Procoxal Metosternum Metapleuron Metacoxal Mesocoxal cavities cavities Propleuron Mesopleuron caviliec Mesosternum Abdominal Cervix X Protternum 1 I Dermestidae Dermestes 1] bei it les vulpinus mi tes on 1 beet le

67-27:15

Dermestes 7 beel les Cânls 0 mit( a

Nitidulidae 2 beel les Stelidota geminata 0 mit< .s

Glischrochilus 1 beel le fasciatus 0 miti.s 1

Head M U S

Cervix R O D ,

Pronotum Y D O B Mesonotum

Elytrol concavity

Metanotum

Ventre 1 to scutellum Wings

Abdominal 4 tergites •p i-in

Abdominal tergites IV-opex 1

- S G E L =

—

-n X APPENDIX E; FIELD COLLECTION DATA

Table 9. Field collection data for host beetle collections

Collection Locality Collection Carcass Duration of number date decomposition (days)

65-10 An.Sc.F. July 13, 1965 Chicken 2 65-12 J. F. July 13, 1965 Chicken 2 65-14 An.Sc.F. July 15, 1965 Chicken 4 65-15 J. F. July 18, 1965 Turkey 2 65-16 J. F. July 20, 1965 Calf 2 65-17 J. F. July 22, 1965 Calf 4 65-19 J. F. Aug. 2, 1965 Calf 15 65-20 J. F. Aug. 12, 1965 Chicken 3 65-21 J. F. Aug. 13, 1965 Chicken 4 65-22 J. F. Aug. 15, 1965 Chicken 6 65-23 J. F. Aug. 18, 1965 Chicken 2 65-24 J. F. Aug. 23, 1965 Chicken 5 65-25 J. F. Aug. 25, 1965 Chicken 7 65-26 J. F. Aug. 28, 1965 Chicken 3 65-27 J. F. Sept. 1, 1965 Chicken 5 65-28 J. F. Sept. 2, 1965 Chicken 6

66-1 J. F. July 22, 1966 Cat 1 66-2 J. F. July 23, 1966 Cat 2 66-3 J. F. Aug. 4, 1966 Pig 2 66-4 J. F. Aug. 5, 1966 Pig 3 66-5 J.,. F. Aug. 11, 1966 Pig 3 66-6 J. F. Aug. 12, 1966 Cat 2 66-7 J. F. Aug. 15, 1966 Pig 7 66-8 J. F. Aug. 17, 1966 Pig 2 66-9 J.]P. Aug. 24, 1966 Sheep 1 66-10 J. P. Aug. 25, 1966 Sheep 2 66-11 J. F. Aug. 26, 1966 Sheep 3 66-12 J. F. Aug. 28, 1966 Sheep 5 66-13 J. F. Aug. 29, 1966 Pigeon 2 66-14 J. F. Aug. 29, 1966 Sheep 6 66-15 J. F. Aug. 30, 1966 Pigeon 3 66-16 J. F. Sept. 1, 1966 Sheep 9 66-17 J. F. Sept. 2, 1966 Sheep 10 66-18 J. F. Sept. 2, 1966 Pigeon 2 66-19 J. F. Sept. 3, 1966 Pigeon 3 66-20 J. F. Sept. 3, 1966 Raccoon 1 66-21 J. F. Sept. 4, 1966 Raccoon 2 66-22 J. F. Sept. 5, 1966 Pigeon 1 121

Table 9. (Continued)

Collection Locality Collection Carcass Duration o£ number date decompos it ion (days)

66-23 J. F. Sept. 5, 1966 Raccoon 3 66-24 J. F. Sept. 6, 1966 Raccoon 4 66-25 J. F. Sept. 7, 1966 Squirrel 2 66-26 J. F. Sept. 7, 1966 Raccoon 5 66-27 J. F. Sept. 7, 1966 Pigeon 3 66-28 J. F. Sept. 8, 1966 , Squirrel 3 66-29 J. F. Sept. 11, 1966 Squirrel 6 66-30 J. F. Sept. 12, 1966 Guinea pig 7 66-31 J. F. Oct. 11, 1966 Dog 3 66-32 J. F. Oct. 13, 1966 Dog 5 66-33 J. F. Oct. 14, 1966 Dog 6

67-1 J. F. April 15, 1967 Dog 6 67-2 J. F. April 16, 1967 Dog 7 67-3 J. F. April 28, 1967 Dog 9 67-4 J. F. April 28, 1967 Pigeon 13 67-5 J. F. June I, 1967 Dog 52 67-6 J. P. June 4, 1967 Dog 1 67-7 J. F. June 11, 1967 Dog 8 67-8 J. F. June 12, 1967 Dog 9 67-9 J. F. June 16, 1967 Dog 4 67-10 J. F. June 26, 1967 Squirrel 1 67-11 J. F. June 28, 1967 Squirrel 3 67-12 J. F. June 29, 1967 Squirrel 4 67-13 J. F. July 1, 1967 Squirrel 6 67-14 J. F. July 1, 1967 Dog 6 67-15 J. F. July 2, 1967 Dog 2 67-16 J. F. July 4, 1967 Dog 4 67-17 J. F. July 4, 1967 Squirrel 4 67-18 J. F. July 6, 1967 Dog 6 67-19 J. F. July 8, 1967 Dog 8 67-20 J. F. July 9, 1967 Dog 3 67-21 J. F. July 17, 1967 Dog 3 67-22 J. F. July 18, 1967 Dog 3 67-23 J. F. July 19, 1967 Dog 4 67-24 J. F. July 20, 1967 Squirrel 1 67-25 J. F. July 21, 1967 Squirrel 1 67-26 J. F. July 21, 1967 Dog 2 67-27 J. F. July 22, 1967 Dog 3 67-28 J. F. Aug. 8, 1967 Dog 1 67-29 J. F. Aug. 9, 1967 Dog 2 67-30 J. F. Aug. 10, 1*967 Dog 1 122

Table 9. (Continued)

Collection Locality Collection Carcass Duration of Number date decomposition (days)

67-31 J. F. Aug. 17 1967 Mouse 2 67-32 J. F. Aug. 17 1967 Dog 2 67-33 J. F. Aug. 18 1967 Dog 3 67-34 J. F. Aug. 18 1967 Dog 3 67-35 J. F. Aug. 24 1967 Dog 2 67-36 J. F. Aug. 25 1967 Dog 3 67-37 J. F. Aug. 26 1967 Dog 4 67-38 J. F. Aug. 27 1967 Dog 5 67-39 J. F. Aug. 29 1967 Woodchuck 1 67-40 J. F. Aug. 30 1967 Dog 2 67-41 J. F. Aug. 31 1967 Dog 3 67-42 J. F. Sept. 1 1967 Dog 4 67-43 J. F. Sept. 4 1967 Dog 7 67-44 J. F. Sept. 5 1967 Dog 8 67-45 J. F. Sept. 6 1967 Dog 1 67-46 J. F. Sept. 7 1967 Dog 2 67-47 J. F. Sept. 11, 1967 Dog 6 67-48 J. F. Sept. 16, 1967 Dog 6 67-49 J. F. Sept. 17, 1967 Dog 7 67-50 J. F. Oct. 2, 1967 Dog 10 67-51 J. F. Oct. 3, 1967 Dog 11 67-52 J. F. Oct. 4, 1967 Dog 12