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Home , Acari, Acaridae, Acromyrmex lundii, Antennophoridae, Antennophorina, Antennophorus

Mites () Associated with the (Formicidae) of Ohio and the Harvester ,

Messor pergandei, of

THESIS

Presented in Partial Fulfillment of the Requirements for the Degree Master of Science in the Graduate School of The Ohio State University

By

Kaitlin A. Uppstrom

Graduate Program in , Ecology and Organismal Biology

The Ohio State University

2010

Master' Examination Committee:

Associate Professor Hans Klompen, Advisor

Associate Professor Susan Jones

Professor Steven Rissing

Copyright by

Kaitlin A. Uppstrom

2010

Abstract

Ants (Formicidae) have long been an group of great interest to the scientific world, whether for their ecological roles, feeding strategies, or social behaviors. They form complex colonies, harboring resources that can potentially be exploited by myrmecophiles (organisms living in association with ants). Myrmecophily has been studied in detail for Coleoptera, but (Acari), the most frequent of ant guests, remain largely unstudied. Previous work has focused primarily on descriptions and has provided little ecological information.

The first study is an effort to provide a more robust list of the often overlooked inhabitants of ant nests focusing on Ohio, a state that has yet to be mentioned in any myrmecophilous studies. A general survey of common Ohio ants was conducted from April 2008-March 2010. Phoretic mites were individually removed from ants and debris in 273 colonies. Mite collections totaled 198 : 151 species phoretic and at least 47 mite species in non-phoretic relationships within the ant nests. Phoretic mites consisted of representatives of the cohort Astigmata ( and ), the cohort (, Pygmephoridae, and Microdispidae), and the suborder (, , and Uropodina). Many mite species were host specific and attachment site specific. An unusually large number of mite species was found to be associated with the ant , possibly the result of social . Post hoc statistical analyses show significantly greater mite diversity in ii colonies when 1) in the ant subfamily , 2) the colony is in the woods, 3) the nest substrate is wood, 4) the colony is populous, 5) are large, and 6) the ant species establishes its nest parasitically.

A second study focused on the seed pergandei and its acarine associates. At least seven mite species are phoretically associated with M. pergandei: Armacarus sp., Lemanniella sp., Petalomium sp., Forcellinia sp., sp., Unguidispus sp., and sp. Most of these species show preference for specific phoretic attachment sites and most preferentially board female alates rather than male alates. Five mite species were found in low numbers inhabiting the chaff piles:

Tydeidae sp., Procaeculus sp., sp., Bakerdania sp., and Tetranychidae sp. The phoretic Petalomium sp. was observed consuming fungus on a dead dealate, but the roles of the other species are still unclear.

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Dedicated to Jana, Kevin, Ariel and Jennifer

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Acknowledgments

I would like to thank my advisor, Dr. Hans Klompen, for his inspired ideas, constant support, enthusiasm, and patience, for the freedom to mold my own unique project, and for the career and life changing moment which began the moment I stepped into the lab as an undergraduate student.

I would like to thank my committee member, Dr. Susan Jones, for her thoughtful guidance, assistance in reducing a growing project into something managable, and for careful editing which allowed me to write what I really intended.

I would like to thank my committee member, Dr. Steven Rissing for his great assistance in providing information and guidance allowing me to arrive safely in Arizona and collect , for corrections and input on my thesis and for asking the hard questions.

I would like to thank Dr. Joseph Raczkowski for advice and insight concerning

Messor pergandei and later the Lasius genus and its habits.

Many thanks to Dr. Dave Walter, Dr. Ron Ochoa, Dr. Cal Welbourn, and Dr.

Barry OConnor, of the Acarology Summer Program, who have provided ecological and taxonomic guidance throughout my project.

I would like to thank the people who assisted with the Messor pergandei project:

Dr. Bob Johnson for information, ideas, and the generous use of his shovel for digging ant craters, Clint Penick for teaching me how to search and rear dealates, Shellie and

Robb Hjellum for welcoming me and my ant specimens into their home.

I would like to thank Bob Bryant and Brenda Bryant for permitting me to collect numerous ant colonies on their property giving me the opportunity to find and Trichocylliba mites.

I would like to thank the members of the Acarology Lab past and present, Monica

Farfan, Luke Kapper, and Sam Bolton for friendship, suggestions, and advice throughout my project.

I am indebted to The Ant Course which allowed me to meet so many interesting people and acquire the knowledge in ant , collecting, and ecology that has been invaluable to my project.

I thank Gary and Holly Coovert for their hospitality, assistance with identification of difficult species groups, and for the invaluable reference, The Ants of Ohio.

Special thanks to my family and close friends: My parents Jana Morse, and Kevin

Uppstrom, for supporting me in all things, school and otherwise, nurturing my love of nature, and for having so many rocks in the backyard that house ants and their mites, my sisters Jennifer Alford and Ariel Uppstrom for support, advice, and for listening to me talk incessantly about bugs, to Gabe Campbell for endless patience, support, the technical skills I lack, and for his mutual interest in ants, to Joshua Bryant, for his company and bravery (particularly for exsectoides) on collecting trips, for his advice on my project, and his patient ear.

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Vita

January, 24, 1985 ...... Born, Hamilton, Ohio, USA

2007...... .S. Zoology, The Ohio State University

2007-2009 ...... Graduate Teaching Associate,

The Ohio State University

2009-2010 ...... Metro School Fellow,

The Ohio State University

1980 to present

Publications

Uppstrom, . and . Klompen. (2005). A new species of Julolaelaps (Acari: Iphiopsididae) from African millipedes. International Journal of Acarology, 31: 143-147. Corriveau, M., K. Uppstrom, and H. Klompen. (2010). Effect of eight storage modes on DNA preservation. In: Sabelis, M. ., Bruin, . (Eds), Trends in Acarology, Proceedings of the 12th International Congress. Springer Science, Dordrecht, pp. 553-556. Fields of Study

Major Field: Evolution, Ecology and Organismal Biology

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Table of Contents

Abstract ...... ii

Acknowledgments...... v

Vita ...... vii

List of Tables ...... xiii

List of Figures ...... xix

Chapter 1: Ants, Mites, and Current Knowledge of Interspecific Associations ...... 1

Family Formicidae: The Ants ...... 1

Ants as Social ...... 1

The Ant Nest ...... 2

Communication in Ants ...... 3

Interactions among Ants ...... 4

Myrmecophiles ...... 6

Mites and Myrmecophily...... 7

Phoresy in Mites ...... 8

Mites Documented from Ant Nests ...... 10

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Scope of these Studies ...... 22

Literature Cited ...... 24

Chapter 2: Mites Associated with Ants in Ohio ...... 38

Introduction: Goals, Questions, and Scope of this Project ...... 38

Materials and Methods ...... 40

Collection Dates and Localities ...... 40

Colony Sampling Methods ...... 40

Sampling Phoretic Mites from Ants ...... 41

Collection and Rearing of Non-phoretic Mites ...... 43

Preservation and Identification of Ants and Mites ...... 44

Results ...... 50

Ant Species Collected and Number Inspected ...... 50

Mite Species Collected ...... 50

Total Number of Mites per Ant Species ...... 51

Host Specificity ...... 51

Attachment Site Specificity ...... 52

Prevalence of Phoretic Mite Infestations ...... 53

Mites on Alates: Prevalence and Sex Preference ...... 53

Ecological Factors Effecting Phoretic Mite Diversity ...... 54

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Estimating Number of Associated Mite Species ...... 61

Non-phoretic Mites from the Nest ...... 62

Specificity and Roles of Non-phoretic Mites ...... 63

Discussion ...... 64

The Significance of Species Specificity and the Issue of Lasius ...... 64

Host Sex Specificity ...... 66

Other Ecological Factors Affecting Mite Species Richness and Abundance ...... 67

The Importance of Rearing ...... 68

Caveats...... 69

Summary of Conclusions...... 70

Literature Cited ...... 104

Chapter 3: Mites Associated with Messor pergandei ...... 107

Interdependence in the Desert: Seed Harvester Ants ...... 107

Associates of Seed Harvesters...... 108

Messor pergandei ...... 109

Natural History ...... 109

The Scope of this Study ...... 110

Methods ...... 111

Locality and collection dates ...... 111

Field Collection ...... 111

Laboratory Observation of Dealates ...... 113

Recovery, Preservation, and Identification of Mites from Ants and Chaff ...... 115

Results ...... 117

Colony Collections ...... 117

Mite Association with Female versus Male Alates ...... 118

Phoretic Mite Species and Abundance ...... 118

Attachment Site Specificity ...... 119

Mites on Dealates ...... 120

Mites in Chaff Piles ...... 121

Voucher Specimens ...... 122

Discussion ...... 122

Yearly Variation and Mite Species Richness ...... 123

Sex and Attachment Site Specificity ...... 124

Alate versus Dealate Mite Abundance ...... 125

Chaff Piles ...... 126

Caveats...... 126

Conclusion Summary ...... 127

Literature Cited ...... 130

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Chapter 4: Synthesis of the Studies, Implications, and Future Research...... 133

Distinctive Qualities of this Study ...... 133

Major Conclusions of the Projects ...... 134

Implications and Suggestions for Future Research ...... 135

Literature Cited ...... 137

Appendix A: Chapter 1 Supplementary Tables ...... 138

Appendix B: Chapter 2 Supplementary Tables ...... 193

Appendix : Chapter 2 Raw Data ...... 206

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List of Tables

Table 1. Results of Simple linear regression between Total ants inspected and response variables...... 54

Table 2. Independent -test for phoretic mites in colonies in the woods and open .

Mesostigmata, Heterostigmatina, and Astigmata have been abbreviated to “Meso”,

“Hetero”, and “Astig” respectively...... 55

Table 3. Independent t-test for phoretic mites in colonies in the wood and substrates.

...... 56

Table 4. Independent t-test for phoretic mites in Lasius colonies founded parasitically or normally...... 57

Table 5. Independent t-test for phoretic mites in colonies founded by ants in the genus

Lasius and non Lasius species...... 57

Table 6. Independent t-test for phoretic mites in colonies of ants in subfamily Formicinae and Myrmecinae...... 58

Table 7. Results of one-way ANOVA (with post hoc Tukey’s LSD method) for phoretic mites in colonies of ants forming small, moderate, and large colonies...... 59

Table 8. Results of simple linear regression showing a significantly increasing trend for all response variables except for the number of Astigmata...... 60

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Table 9. Stepwise multiple linear regression model results for determining the response variables...... 61

Table 10. Astigmata Mites and Hosts. Mite species are shown with their associated host species, number of representative slides, and number of colonies in which the mite species were found for each host...... 71

Table 11. Heterostigmatina Mites and Hosts. Mite species are shown with their associated host species, number of representative slides, and number of colonies in which the mite species were found for each host...... 75

Table 12. Mesostigmata Mites and Hosts. Mite species are shown with their associated host species, number of representative slides, and number of colonies in which the mite species were found for each host...... 78

Table 13. Number of Phoretic Mites. Each ant species is shown with number of colonies and individuals inspected and the resulting number of phoretic mites encountered...... 80

Table 14. Ants and their Associated Mite Species...... 82

Table 15. Astigmata host specificity. All phoretic Astigmatid species are shown with codes designating their specificity levels. 1 = associated with only a single host species,

2= associated with multiple host species in the same genus, 3 = associated with multiple hosts in separate genera...... 89

Table 16. Heterostigmatina Host specificity. All phoretic Heterostigmatid species are shown with codes designating their specificity levels. 1 = associated with only a single host species, 2= associated with multiple host species in the same genus, 3 = associated with multiple hosts in separate genera...... 90

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Table 17. Mesostigmata Host Specificity. All phoretic Mesostigmatid species are shown with codes designating their specificity levels. 1 = associated with only a single host species, 2= associated with multiple host species in the same genus...... 92

Table 18. Astigmata Phoretic Attachment Site Specificity. Number of specimens (slides) collected at each of 8 phoretic locations are shown. Phoretic attachment locations are as follows: 1-thorax/petiole, 2-antennal, 3-head, 4-between legs/coxae, 5-on legs, 6-gaster,

7-wing, 8-unknown...... 93

Table 19. Heterostigmatina Phoretic Attachment Site Specificity. Number of specimens

(slides) collected at each of 8 phoretic locations are shown. Phoretic attachment locations are as follows: 1-thorax/petiole, 2-antennal, 3-head, 4-between legs/coxae, 5-on legs, 6- gaster, 7-wing, 8-unknown...... 94

Table 20. Mesostigmata Phoretic Attachment Site Specificity. Number of specimens

(slides) collected at each of 8 phoretic locations are shown. Phoretic attachment locations are as follows: 1-thorax/petiole, 2-antennal, 3-head, 4-between legs/coxae, 5-on legs, 6- gaster, 7-wing, 8-unknown...... 96

Table 21. Non-phoretic Astigmatid mites, their hosts, and locations where the mites were found in the nest. *designates a deutonymph which was also observed in a phoretic relationship. 1 Indicates a species ( sp2) which was reared in the lab.

Undetermined species (undet) may actually be different species...... 99

Table 22. Non-phoretic Prostigmatid mites, their hosts, and locations where the mites were found in the nest. *designates a mite which was also observed in a phoretic relationship...... 101

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Table 23. Non-phoretic Mesostigmatid mites, their hosts, and locations where the mites were found in the nest. *designates a mite which was also observed in a phoretic relationship...... 102

Table 24. Alate Summary and Sex Preference of Mites. Total number of male (M) and female () alates collected per colony and number with phoretic mites. 1Signifies two colonies collected at the Cox Rd site. All other colonies were collected at the W

McCartney Rd site. “/a” is used when no alates of that sex were found within the colony...... 117

Table 25. Host Sex Preference of Mite Species. Number of individuals of the 6 mite species found in each colony on male (M) and female (F) alates and percentages of the total mite fauna that each species comprises (N=593)...... 119

Table 26. Phoretic Locations. Number of mites for each species found at corresponding locations on the hosts...... 120

Table 27. Mites Associated with Messor Species...... 129

Table 28: Temporary Social Parasitism Interactions among Ohio Ant Species. Data presented are compiled from literature in which mixed colonies were observed or laboratory trials indicated acceptance of the parasitic species. The only social parasite

Formica species collected was F. exsectoides. 1Designates a species which was not collected in Chapter 2’s study, but has been documented in Ohio...... 139

Table 29. Mesostigmata and their Associated ants from Literature…...... 140

Table 30. Heterostigmatina and their Associated Ants from Literature...... 170

Table 31. Astigmata and their Associated Ants from Literature...... 182

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Table 32. Non Mesostigmatid, Heterostigmatid, and Astigmatid mites and their

Associated Ants from Literature...... 189

Table 33. Localities and Colony Collections. Locality codes, full locality names, coordinates, collection dates, colonies collected at each locality (field numbers), and number of colonies and ants are shown...... 194

Table 34. Mite voucher specimens deposited in the Ohio State University Acarology

Collection as slides and their corresponding OSAL barcode numbers...... 199

Table 35. Ant voucher specimens deposited in the Ohio State University Insect

Collection as pinned specimens and their corresponding OSUC barcode numbers...... 203

Table 36. Ohio Ant Collection Summary: Ant species, number of colonies collected (#

COL.), and number of each caste (F=female, M=male) collected are shown in the table.

*Designates a collection made by putting the entire collection on the funnel rather than by the usual method...... 204

Table 37. Raw data for Response variables used in Statisical analyses. # Phoretic indicates total number of phoretic mites found in the colony. “# Meso”, “# Hetero”, and

“# Astig” designate number of Mesostigmata, number of Heterostigmatina, and number of Astigmata respectively associated with the colony. “Prev with” is the prevalence of mite infestations in the colony...... 207

Table 38. Raw data for Predictors used in Statistical Analyses. Codes used for predictors are as follows: Subfamily- 1=Formicinae, 2=Myrmecinae, 3=,

4=, 5=Amblyoponerinae. Parasitic: 1= Parasitic, 2=non parasitic. Lasius:

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1=Lasius, 2=Not Lasius. Colony size: 1=small, 2=moderate, 3=large. : 1=woods,

2=open. Nest substrate: 1=wood, 2=soil...... 214

Table 39. Raw data for Prevalence. Ant species used in statistical analyses of prevalence are shown. "W" designates worker, "F" designates female alate, "M" designates male alate. Mean (M), Standard deviation (SD), and number (N) are calculated in the last 3 rows of the table. “n/a” is used when no castes of a type were found in the nest...... 222

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List of Figures

Figure 1. Simplified Summary of the Classification of Acari discussed in Chapter 1.

1Designates a group with many documented ant associates...... 13

Figure 2. Species Accumulation Curves. Cumulative number of mite species, Sobs (Mao-Tau), per number of colonies collected is shown...... 97

Figure 3. Lasius Species Accumulation Curves. Cumulative number of mite species, Sobs

(Mao-Tau), per number of colonies collected is shown...... 97

Figure 4. Ponera and Solenopsis Species Accumulation Curves. Cumulative number of mite species, Sobs (Mao Tau), per number of colonies collected...... 98

Figure 5. Messor pergandei Species Accumulation Curve. Cumulative number of mite species, Sobs (Mao Tau), per number of colonies collected...... 124

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Chapter 1: Ants, Mites, and Current Knowledge of Interspecific Associations

Ants (: Formicidae) have long been an insect group of great interest to the scientific world, whether for their ecological roles, feeding strategies, or social behaviors (Hölldobler & Wilson, 1990). They form colonies which alter the available nutrients, texture, moisture, and temperature of the surrounding soil to suit their environmental needs (Laasko & Setälä, 1998). Kin recognition is involved in protection of the colony’s stored goods and brood, but myrmecophiles (organisms living with ants) are able to overcome these intrinsic colony defenses and apparently have cracked the ants’ code (Donisthorpe, 1927; Hölldobler, 1971). Although myrmecophily has been studied in detail for some groups of (primarily ), mites (Acari) are among the most frequent of ant guests-yet remain largely unstudied (Kistner, 1982;

Rettenmeyer, 1962a).

Family Formicidae: The Ants

Ants as Social Insects

The Hymenoptera is comprised of , ants, and . Hymenopterans have attracted the attention of scientists largely due their complex social behaviors. There are varying levels of sociality throughout the class Hexapoda, from the lower levels of subsociality, quasisociality, and semisociality to the highest degree of sociality known as (Hermann, 1979). Eusocial organisms satisfy the following criteria: 1) 1 formation of a long-lived colony with nestmate recognition, 2) a seemingly altruistic reproductive strategy in which one or few reproduce, while there are many sterile or non- reproductive workers, 3) polyethism, or division of labor, based on 4) caste differentiation, or different morphological or physiological characteristics for members with different roles, and lastly 5) overlapping generations in which offspring assist parents and siblings with rearing, defense, and provisioning of the colony (Hermann,

1979; Lin & Michener, 1972). Eusocial insects primarily are members of two orders:

Hymenoptera and Isoptera (). Within the Hymenoptera, the majority of species in the family Formicidae, the ants, are eusocial.

The Ant Nest

As eusocial insects, the ants create carefully structured nests consisting of and galleries usually in soil or wood. Within these nests, the ants organize their resources

(brood and stored ) and waste in designated chambers (Hölldobler & Wilson, 1990).

Ants nesting in soil create the most structurally complex nests. Tschinkel (2003; 2004) studied ant nests in the sandy soil of by pouring a slurry of tin and orthodontal plaster down the nest entrance. After excavation he found that nest architecture varied by species and age of the colony. Nests ranged from quite shallow (20-30 cm deep) such as those belonging to Aphaenogaster sp. and Formica pallidifulva to the extensive nests

(almost 4 m deep) of imparis and Pogonomyrmex badius. Nests either were sprawling with many branches or simple with one major interspersed with side galleries.

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Through the construction of their nest, the inhabitants alter the condition of the nesting substrate. The humidity, texture, chemical composition, temperature, and amount of organic materials are drastically altered by the ants (Wagner et al., 2009; Boulton et al., 2003; Beattie & Culver, 1983; Cole, 1994; Laakso & Heikki, 1998). This localized change in the soil (or other substrate) composition has been demonstrated to increase litter by nematodes, mites, collembolans, and microorganisms within the nest (Paris, 2008), as as to increase the overall abundance and biomass of these associated organisms (Beattie & Culver, 1983; Boulton et al., 2003; Wagner, 1997).

Communication in Ants

As eusocial insects, the ants have developed methods of communication which allow the colony to function efficiently and collaboratively in activities such as nest building, foraging, care of the brood, and protection of the colony (Hölldobler, 1971).

Ant colonies can be likened to heavily guarded fortresses to which one must not only look like a member, but also smell and feel like a member in order gain access to resources therein. Much of the information is passed along in the form of chemical cues which elicit responses in fellow nestmates.

Cuticular hydrocarbons are sensed by ants and designate colony membership

(Lahav et al., 1999), role or task designation (Greene & Gordon, 2003), and fertility

(Dietemann et al., 2003; Liebig et al., 2000). Additionally, are used by the ants to alert nestmates of other environmental aspects such as approaching danger or for recruitment to food resources.

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Tactile communication is another method employed by ants. One of the most well known methods of tactile communication is trophallaxis, a method by which an ant shares liquid food stored in its crop with another ant through regurgitation. The receiving ant taps the donor’s body with her antennae which signals for the donor’s attention. The receiver may also tap on the donor’s mouth parts if a regurgitation reflex is not triggered by the initial attempt (Hölldobler & Wilson, 2009).

Interactions among Ants

Generally access to the colony is permitted only to members, but under certain conditions, interspecific associations of ants may be encountered. These interactions can be classified into two broad categories, compound nests (brood from more than one species is kept separate) and mixed nests (brood from more than one species is mixed).

Many of these interspecific associations are aided through chemical camouflage or mimicry on the part of a parasitic ant species (Lenoir et al., 2001).

Plesiobiosis, cleptobiosis, lestobiosis, parabiosis, and xenobiosis are relationships in compound nests as described by Wheeler (1910a) and elaborated upon by Hölldobler and Wilson (1990). In plesiobiosis, two or more species of ants nest in close proximity but participate in almost no direct communication. In cleptobiosis, a small ant species nests near a larger species’ nest and steals food from foragers of the host species as they return to the nest. They may also scavenge on their refuse piles. A similar situation known as lestobiosis involves the smaller ants building their nest in the walls of the larger ants’ nest and stealing resources including brood for consumption. Many ants in the genus Solenopsis are known as “thief ants” and are frequent lestobionts. The two more

4 intimate compound nest associations are parabiosis, in which two species use the same nest and often share odor trails, and xenobiosis in which one species moves freely among the host ants, soliciting food through trophallaxis.

Mixed colony situations usually are due to social parasitism. There are three types of social parasitism in ants as defined by Wheeler (1910a): dulosis (or slave-making), temporary, and inquilinism.

Dulosis is an obligatory dependence of one ant species on another for regular maintenance and labor within its nest. The so called “slave raiders” enter the host ants’ nest, steal pupae, and return with them to their own nest. The ants that emerge from the stolen pupae, then do all of the work in the parasite’s nest (except slave raiding) for the remainder of their life. If the slave-ants are reintroduced to their original host colony, the ants will not recognize each other as kin, despite the genetic relationship. The dulotic ant species is completely dependent on the host ant species, unable to survive without the labor of the workers they enslave and thus necessitating continued slave-raiding trips

(Hölldobler & Wilson, 1990).

Temporary social parasitism involves the invasion of a host nest by a newly mated gyne (female reproductive ant) of a different ant species. The gyne eventually is accepted by the host workers, through force or appeasement, and she kills the host queen.

She then replaces the queen, and the workers care for the offspring she produces.

Eventually the host workers die naturally, and the colony consists entirely of the parasitic species’ workers (Hölldobler & Wilson, 1990). The ant genus Lasius has many social

5 parasites (Raczkowski, 2008; Wing, 1968); Lasius species of particular interest to the study in Chapter 2, and temporary social interactions are in Table 28 in Appendix A.

The final type of mixed colony association due to social parasitism is known as inquilinism. Social parasitism of this type is the most degenerate. The queen of the parasitic species enters the host’s nest and usually does not kill the host queen, thereby ensuring the presence of a continuous supply of host workers. Most inquiline species have lost the ability to make the worker caste, and instead they produce only other reproductives, which then leave the nest to parasitize other colonies (Hölldobler &

Wilson, 1990).

Myrmecophiles

Myrmecophiles are defined as organisms living in close association with ants.

Most of these symbionts are not simply present by accident, but rather are thought to depend on their ant host for all or part of their life cycles. Like socially parasitic ants, myrmecophiles manage to infiltrate the nest, hence gaining access to the abundant resources. Colonial insects such as ants are particularly good hosts for other arthropods as their colonies are long lived and rich in organic materials collected from a larger area than that of the nest (Laakso & Setälä, 1998; Wheeler, 1910a). Additionally, these nests are a source of fresh and nutritious food resources (in the form of the host’s brood), have increased temperature and humidity during all seasons, and are protected by the ants from outside intruders (Wheeler, 1910a). The interactions among the ants and their guests can be quite diverse, ranging from hostility to care (Donisthorpe, 1927; Wheeler, 1910a).

The list of arthropods living in association with ants according to Kistner (1982) consists

6 of 18 groups, including class Arachnida, subclass Diplopoda, class Crustacea, and the insect orders Collembola, Orthoptera, Blattaria, Coleoptera, , Neuroptera,

Lepidoptera, Diptera, and Hymenoptera.

Myrmecophiles are divided into five categories (Hölldobler & Wilson, 1990;

Kistner, 1979; Wheeler, 1910a):

1. Synechthrans- These invaders are treated with hostility by the ants and are

usually predators that force themselves upon their hosts. They evade attempts

on the hosts’ part to eliminate them because of their agility and speed.

2. Synoeketes- These organisms are usually scavengers and predators within the

nest, but are tolerated by their hosts. They are generally slow and presumably

overlooked or of little threat to their hosts.

3. Symphiles- These are “true” guests, accepted by their hosts into the colony

and fed and groomed as if they are true members.

4. Ectoparasites and endoparasites- These organisms are true parasites that live

on or in the body of their hosts, respectively, feeding on bodily fluids.

5. Trophobionts- This group includes arthropods (such as homopterans) that

provide the ants with food in the form of or other nutritive

secretions and in return are protected and transported by the ants. This is a

true .

Mites and Myrmecophily

Mites (Arachnida: Acari) are the most frequently encountered and abundant of the myrmecophiles. Approximately 55,000 species of mites have been described with

7 estimates for total diversity ranging from 500,000 to 1,000,000 species (Krantz, 2009).

Due to their relatively small size, mites are able to utilize microhabitats in a range of environments. Mites can be found in habitats ranging from terrestrial, aquatic, and marine, to small-scale niches such as pores and follicles. The minute size of many mites also allows them to act as hitchhikers on other such as birds, , and insects (Krantz, 2009), a behavior which will be discussed in detail in the following section, “Phoresy in Mites”.

Mites may play the roles of parasites, predators, fungivores, or scavengers within the ant nests. Some of the known behaviors of myrmecophilous mites will be further detailed in the section of this chapter titled, “Mites Documented from Ant Nests”.

Phoresy in Mites

The term phoresy is applied to symbiotic relationships in which one organism attaches to another for dispersal. While dispersing, no harm or benefit is brought upon the host. The phoront (the passenger) ceases all reproductive and feeding activity, benefiting from the interaction by receiving transportation. Acari in particular are well known phoronts, climbing aboard larger animals when their current habitat becomes unsuitable due to overcrowding, inadequate humidity, food limitations, or other unexplored reasons

(Houck & OConnor, 1991). Through this symbiotic relationship, mites are able to exploit spatially or temporally scattered habitats more successfully than they would be able to without external assistance (OConnor, 1982). The process of phoresy requires the phoront to receive cues initiating the phoretic behavior, which encompasses actively

8 seeking a host through attractive signals, attaching to the host’s surface, and then remaining quiescent until signals are recognized to disembark (Kaliszewski et al., 1995).

Depending on the type of mite, it may exhibit a specialized developmental stage in which phoresy may occur. For example, mites in the infraorder Astigmata form modified deutonymphs (the second of the three nymphal instars) which look drastically different from the other instars. These deutonymphs usually have large ventral suckers, short legs, a compact tear-drop shape, and a nonfunctioning gut (OConnor, 1982). In the

Heterostigmatina (which includes families Scutacaridae and Pygmephoridae), the mites are phoretic as morphologically specialized females known as phoretomophs (OConnor,

1982). The largest types of mites encountered in ant nests belong to the suborder

Mesostigmata. Most of the Mesostigmata simply grip their phoretic hosts with their tarsal claws (Hunter & Rosario, 1988). In the suborder Uropodina, however, mites have developed other ways to remain on their hosts such as biting onto the hosts’ hairs, secreting gluey stalks from which they suspend themselves, or having modified body shapes which allow them to secure themselves to specific locations on their host’s body

(Elzinga, 1978; Hunter & Rosario, 1988).

It has been documented that when the host’s sex determines the mites’ continued survival or transport to a habitat, the mites board the sexes differentially. For example,

Naiadacarus arboricola (Astigmata: Acaridae) deutonymphs will not attach to male tree hole (Syrphidae: Mallota sp.) because males do not return to a tree hole after mating

(Fashing, 1976).

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Phoretic mites may also correlate their development with their host’s life cycle.

Species which exhibit cyclical phoretic populations are often highly specific and may be found only in the nest of their host species (Kaliszewski et al., 1995).

Once the host reaches the phoront’s preferred habitat (nests, , dead insects, food storages, fungus gardens, dung, carrion, etc.), the mite disembarks, develops into a feeding instar, and subsequently reproduces (OConnor, 1982). In some cases, the interactions may be mutualistic as in Scutacarus acarorum (Heterostigmatina:

Scutacaridae) which feeds on pathogenic mold in the host bumblebee nest. Alternatively, the phoront may be a parasitoid as in the case of Iponemus (Heterostigmatina:

Tarsonemidae), which remains phoretic until immature bark beetles are available. In many cases, the non-dispersal (feeding) stages are unknown or undocumented

(Kaliszewski et al., 1995).

Mites Documented from Ant Nests

Unlike myrmecophilous beetles (relatively large and easily observed) which have had numerous papers and studies devoted to their biology and habits, mites remain enigmatic and seriously underrepresented in the literature. Few studies attempt to provide any ecological details, rarely stretching beyond basic descriptions of the new species encountered, despite the abundance and diversity of mite associates (Hölldobler &

Wilson, 1990; Kistner, 1982; Rettenmeyer, 1962a). Most of these descriptions have focused on the large mesostigmatid mites, which are more difficult to overlook. In a survey of approximately 150 colonies (primarily spp.), Rettenmeyer

(1962a; 1962b) determined that mites are the most abundant guests, usually more than

10 twice as abundant as the insect myrmecophiles, and sometimes more than 100 times more abundant.

The majority of recent studies attempt to at least identify mites to the family level, but still, many refer to the Acari recovered from the nest as simply “mites” or at another broad taxonomic level which tells readers little about the mites’ ecology or possible roles in the ant colonies (Boulton et al., 2003; Park, 1932; Wagner et al., 1997; Witte et al.,

2008). Likewise, many descriptions of mites do not clearly define the ant host, labeling it simply as “in ant nest” (Delfinado & Baker, 1976; Mahunka, 1970b; Mahunka &

Mahunka-Papp, 1980). Rettenmeyer (1962a) also observed the difficulty in obtaining information about myrmecophiles and urged readers to make every effort “to report accurate observations however insignificant they may seem by themselves”.

Available literature concerning mites and their associated ants is scattered among various journals, many of which are difficult to obtain due to obscurity or age.

Consequently, few have attempted to organize the literature into any manageable form.

The earliest attempt to consolidate the literature was by Berlese (1904) who included a list of more than 80 myrmecophilous Acari and their hosts; however, many of these early descriptions are very brief with poor drawings or are devoid of accompanying drawings, making subsequent identification of these species difficult or ambiguous (Gorirossi-

Bourdeau, 1993). The most recent list was compiled by Hölldobler and Wilson (1990), but it overlooked all but the most cited literature, namely the lists by Kistner (1982), and

Donisthorpe (1927), and Rettenmeyer’s doctoral work (1962a).

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To further complicate the issue, mite taxonomy is in a state of flux; many previously described myrmecophilous mites are no longer in the same genus and frequently are in different families. In the following section are my efforts to unearth the forgotten articles regarding the Acarine inhabitants of ant nests. I have included brief highlights of some of the more informative myrmecophilous mite studies, organized by major mite groups (Mesostigmata, Heterostigmatina, , and Astigmata), and I provided some basic biological information about these groups. A full list of mite species associated with ants, as collected from the literature, can be examined in Appendix A

(Table 29 to Table 32). Hosts have been updated in the list to include the current valid scientific names.

Brief Introduction to Mite Taxonomy

The subclass Acari is comprised of two superorders, and

Acariformes. Ant mite associates belonging to the superorder Parasitiformes can be found in only one of the 4 constituent orders, the Mesostigmata. Within Mesostigmata associates are found in 3 of the 4 lineages (Klompen et al., 2007): Trigynaspida,

Uropodina, and Gamasina but not Sejina. In the superorder ant associated mites can be found in both orders (Krantz & Walter, 2009): (cohort

Heterostigmatina) and Oribatida (primarily cohort Astigmata). A simplified summary of mite classification discussed in this chapter is shown in Figure 1.

12

Trigynaspida1

1 Uropodina Parasitiformes Mesostigmata Gamasina1

Sejina Pygmephoroidea1 Acari Scutacaroidea1 Prostigmata Heterostigmatina Tarsonemoidea Acariformes Oribatida Pyemotoidea (includes Astigmata)

Figure 1. Simplified Summary of the Classification of Acari discussed in Chapter 1. 1Designates a group with many documented ant associates.

Order Mesostigmata

The Mesostigmata are the largest of the Acari (200- 4,500 μm) found in ant colonies and consequently the most thoroughly documented. Most of the Mesostigmata are free-living predators, while others are symbionts or parasites of and arthropods. Relatively few are known to feed on fungi, , and nectar (Lindquist et al., 2009). The most frequently mentioned Mesostigmata in ant literature are army ant associates, Antennophorus species (Trigynaspida: : Antennophoridae), members of the Uropodina, and various species in the family Laelapidae (Gamasina:

Dermanyssina).

Although Rettenmeyer’s (1962a) study of ant mites focused on all mites associated with the army ants of Central and South America, his Mesostigmata observations were the most numerous and thus his work will be summarized in this section and briefly mentioned in subsequent sections for the other mite groups he discovered. 13

Rettenmeyer (1962a) hypothesized that mites associated with army ants fall into two ecological groups: those found on the refuse deposits and those found on the ants or in the nest. To test this hypothesis, Rettenmeyer (1962a) collected bivouacs (a temporary colony formed from living worker ants with their tarsal claws interlocked) in buckets.

The ants were anesthetized with ether, and colony members were individually inspected for myrmecophiles. He acknowledged that this methodology led to the loss of many of the mites, as they often dropped off their hosts and stuck to the sides of the container. He also noted that individual inspection was useful for determining the phoretic position of mites on their hosts, but it was time consuming and thus used occasionally. Rettenmeyer also sampled the refuse deposits of the army ants by means of Berlese funnels and field and laboratory observations.

Since the publication of Rettenmeyer’s thesis, most of the mites he discovered have been described. The total army ant mite fauna comprises 126 described species in

33 genera, with 21 of the genera known only from army ants (Eickwort, 1990; Gotwald,

1996). The vast majority of mites riding on the ants are believed to be phoretic rather than parasitic (Gotwald, 1996), though that assumption is based primarily on cheliceral structure. Supporting behavioral data are rarely available

Army ants are nomadic foragers, never making permanent colonies; instead they move the colony to areas with abundant resources. The constant movement of the host has selected for associates that have methods allowing them to travel with their hosts

(Gotwald, 1995). Many of the mite species evolved some form of morphological modification which allowed for ease of transport and in some cases concealment. Mites

14 in the genus Planodiscus (Uropodina: Planodiscidae), for example, attach themselves to the tibiae of their host ants during phoresy, and SEM photography reveals that they have sculpturing and setal arrangements identical to the host’s leg (Elzinga, 1990; 1991;

Elzinga & Rettenmeyer, 1966; 1970; Gotwald, 1995; Kistner, 1979). Wasmannian mimicry is similarly exhibited by mites in the family Larvamimidae (), also entirely specific to army ants, which imitate the larvae of the ants. Based on the cheliceral structures, these mites are believed to be parasitic on the ant larvae. During emigration, larvamimids are carried by the army ants as if they are the ant larvae

(Elzinga, 1993). Yet another astounding mimic, is the mite rettenmeyeri

(Macrochelidae). This parasite feeds from the pulvillar membrane of tarsi III, and its long, curved rear legs act as a substitute for the normal tarsal claws of the ants. During construction of the bivouac, these mites are, in fact, used as if they are ordinary tarsal claws, thereby holding the bivouac together (Gotwald, 1995; Gotwald, 1996; Krantz,

1962; Rettenmeyer, 1962c). Further morphological modifications have been described in various journal articles by Elzinga, for the Uropodina genera, Coxequesoma (1982a),

Antennequesoma (1982b), Habeogula (1989), Trichocylliba (1981), and Circocylliba

(Elzinga & Rettenmeyer, 1974). All of these mites have their dorsal regions expanded into flanges to form a ventral cavity, allowing them to fit tightly to specific areas of the hosts’ bodies (Elzinga, 1978).

The second most well known mite group associated with ants is the genus

Antennophorus (Trigynaspida: Antennophoridae). These are among the largest of the mites associated with ants, and consequently were avidly observed into the early 1900s

15

(Janet, 1897a; 1902; Karawajew, 1906; Wasmann, 1902; Wheeler, 1910b). Associated with the ant genus Lasius, these mites cling to the underside of a worker ant’s head.

Using elongated forelegs that resemble the ant’s antennae, they vigorously tap the mouthparts or antennae of the host provoking a trophallactic response. Regurgitated liquids are lapped up by the mite. The Antennophorus mites harass their hosts to such an extent that the parasitized workers are no longer able to fulfill their duties in the colony

(such as larval care and nest maintenance). Experiments have shown that mites removed from their hosts quickly climb back onto their hosts. When placed outside the colony,

Antennophorus are in many cases picked up, carried back into the nest by the host, and placed amongst the brood suggesting a cuticular hydrocarbon array similar to that of the ant brood (Franks et al., 1991).

The suborder Uropodina has numerous ant associates. The most morphologically unique of the Uropodina are found with the army ants; however, many more are associated with other ant groups. In a survey of () associated with ants in the southeastern , 17 new species were described, 12 of which were specific to a single ant species (Hunter & Farrier, 1975; 1976). When observed on the host, they were always found on the tibial spur. Donisthorpe (1927) also found an Oplitis species

(=Urodiscella philocenta) clinging to the tibial spur and suggested that the mites feed on substances scraped from the ant during cleaning with the spurs. Lehtinen (1987), however, suggests that the mites are simply dispersing by way of the ants’ tibial spurs.

Two members of Uropodina are clearly parasites of the brood of ants. An undescribed uropodine was found feeding on pupae of Pheidole megacephala, an

16 invasive introduced species in Japan. None of the parasitized pupae survived to adulthood. The same uropodine species was found on the native ant Pheidole noda and appears to have shifted to the non-native Pheidole megacephala (Le Breton et al., 2006).

A second uropodine, sellnicki, from Columbia has been suggested as a possible biological control agent for the invasive crazy ant, Paratrechina fulva. It feeds on the gular region of the ant pupae for its entire development to the extent that the dies soon after the adult mite emerges from the deutonymphal . The mite does not change positions during feeding, and all of the exoskeletons from previous stages can be found beneath the adult mite after (Krantz et al., 2007).

Among Gamasina, genera of Laelapidae (Dermanyssoidea) are most frequently found in ant nests, including Gymnolaelaps (Conway, 2003; Hunter & Costa, 1971),

Cosmolaelaps, formerly called (Gray, 1974; Hull, 1923; Michael, 1891),

Oolaelaps (Donisthorpe, 1927), Laelaspis (Hunter, 1964; Porter, 1985), and

Myrmozercon (Hull, 1923; Hunter & Hunter, 1963; Rosario & Hunter, 1988; Shaw &

Seeman, 2009; Vitzthum, 1930; Walter, 2003). Some such as Gymnolaelaps and

Oolaelaps are found on the eggs and young larvae of the ants and are believed to feed on the secretions applied to the brood by the workers (Donisthorpe, 1927). Others such as

Cosmolaelaps, are believed to scavenge on dead ants (Kistner, 1982).

Suborder Prostigmata: Cohort Heterostigmatina

Eight superfamilies of mites are included in the cohort Heterostigmatina, most of which are small or moderately sized (80-675 μm) (Walter et al., 2009). Three of these are commonly found associated with ants, Tarsonemiodea, Pygmephoroidea, and

17

Scutacaroidea. Although members of the superfamily Pyemotoidea have been observed as parasitoids of ant larvae in cultured situations, they are generally not considered common ant associates in nature (Bruce & LeCato, 1980; Kaliszewski et al., 1995).

Other Heterostigmatina which are clearly associated with ants have bladelike or styletlike chelicerae which are usually used for fungivory. Many display feeding specificity, with phoretic females carrying fungal spores in sporothecae, in order to inoculate their habitat with their preferred food source (Ebermann & Hall, 2003; 2004). In only a couple of cases, Glyphidomastax and Perperipes on army ants (Pygmephoroidea: Microdispidae), have Heterostigmatina been observed feeding on the eggs or immature instars of their host as parasites or parasitoids (Cross, 1965; Kaliszewski et al., 1995).

Most of the work with ant associates in the Heterostigmatina has focused on descriptions of new species encountered. Following up on Rettenmeyer’s discoveries,

Mahunka (1977a; 1977b) identified 22 of the Heterostigmatina species (in families

Microdispidae and Scutacaridae), 20 of which were new to science. Ross and Cross

(1979) provided descriptions of mites in the genus Acinogaster (Pygmephoridae), also based on Rettenmeyer’s army ants. Preferred attachment locations for phoretic

Heterostigmatina on the army ants were between the coxae ventrally (Rettenmeyer,

1962a).

General surveys of heterostigmatid mites were conducted in (Metwali,

1981), Turkmenistan (Khaustov & Chydyrov, 2004), (Mahunka, 1970b), and

Austria (Ebermann, 1979). Delfinado & Baker (1976) described many mites in the family Scutacaridae associated with a variety of animals including ground beetles

18

(Carabidae), rodents, shrews, bees (Halictidae and Adrenidae), leaf cutter ants (Atta), and termites (Reticulitermes).

For most of the descriptions, only information concerning phoretic females is provided due to lack of availability of reared specimens. Biological and ecological aspects have been explored in a couple of studies in which laboratory rearing was successful. Petalomium fibrisetum (Pygmephoroidea: Pygmephoridae), associated with

Lasius flavus, was found to feed on hyphae of different fungi available in ant nests. Egg production of females was dependent on the species and age of the fungi with which they were provided, and lack of food was a cue for phoretic behavior (Ebermann & Rack,

1982). Larvae and males were reared for another Lasius flavus associate, Imparipes brevitarsus (Scutacaroidea: Scutacaridae) (Ebermann, 1981). More recently, Ebermann and Moser (2008) collected 5 species of scutacarid mites associated with red imported fire ants (Solenopsis invicta) in Louisiana. The study focused on alates (the winged unmated reproductive ant caste); the mites were found more frequently on the female alates than the male alates. Numbers of alates and number of individual mites were documented throughout the season; the two were clearly correlated. The authors rule out any parasitoid habits for the mite species described, citing the presence of small pharyngeal pumps which are indicative of fungivory.

Suborder Oribatida

Although the suborder Oribatida now includes the Astigmata, the two are often discussed separately in the literature due to their vast differences in biology, morphology and life cycle. Oribatids typically range in size from 300-700µm. Life history strategies

19 in this group primarily involve low reproduction and long development times, and have resulted in the evolution of many defensive mechanisms in this suborder including camouflage, waxy secretions, defensive glands, thick cuticles, or flaps providing protection to sensitive areas (Norton & Behan-Pelletier, 2009).

The majority of oribatids inhabit soil or litter habitats where they feed on decomposing litter or fungi; relatively few are believed to be associated with ant colonies

(beyond presence in the soil environment) (Ito & Aoki, 2003). Those oribatids associated with ants are on opposing ends of the spectrum of interactions. In some cases, oribatids

(in the families , , , , , and

Scheloribatidae) are preferred prey for ant species (.. for the genera Pheidole (Wilson,

2005) and (Masuko, 1994)) and secondary prey (Collembola are primary) for trap jaw ants in genera such as and Pyramica (Masuko, 1984; 2009).

More specific symbiotic interactions have been shown between the mite

Protoribates myrmecophilus which is groomed (though infrequently) by ants in the genus

Myrmecina. P. myrmecophilus feeds on dead wood and fungal material within the nest.

The host ants feed upon dead mites but they have never been observed eating any live mites. When mites attempted to exit the ant nest, ants were observed bringing P. myrmecophilus back into the nest. Additionally ants were frequently observed carrying the mites with them when moving to a new nest site (Ito & Aoki, 2003). This apparent symbiotic relationship is overshadowed by an earlier study involving Aribates javensis, also found in nests of Myrmecina, which exhibits a clearly mutualistic relationship. A. javensis are not only groomed and cared for by the ants, but they never walk by

20 themselves and rely on the ants for transport despite their well-developed legs. Eggs laid by the mites are collected from the ovipositor by the ants and placed amongst the ant egg cluster where they are cleaned and licked by the ants. In fact, A. javensis can only survive

3 days without the host ants’ care. Similar to Protoribates, A. javensis are eaten once they die, which appears to be their main benefit to ants (Aoki et al., 1994).

Suborder Oribatida: Cohort Astigmata

The Astigmata includes 71 families of generally small to medium sized mites.

Their size and the general lack of Astigmata specialists limited their detection and subsequent documentation in the ant mite literature, relative to other mite groups such as the Mesostigmata. Unlike the other oribatids, the Astigmata exhibit short developmental time and high fecundity making them highly effective at exploiting ephemeral habitats.

Phoretic associations with other arthropods by heteromorphic deutonymphs (sometimes referred to as hypopi) have also allowed the Astigmata to utilize patchy habitats where they usually feed on decaying organic material, fungi, bacteria, or sometimes nematodes.

Most Astigmata have little sclerotization and little or no defensive mechanisms at their disposal (OConnor, 1982; 2009).

Literature regarding the Astigmata associated with ants is scanty and most papers lack details beyond basic descriptions. Rettenmeyer’s collections, yielded some

Astigmata descriptions (Mahunka, 1978; 1979), but Rettenmeyer himself noted that many of the smaller mites were lost during processing (1962a). Forcellinia (Acaridae) is one of the most ant-specific genera with 15 species, all found in ant nests or on the ants themselves (Fain, 1987). Lemanniella (Lemanniellidae, formerly in Acaridae) is another

21 genus in which all species have been described from ant nests. In one of the infrequent rearing and observational studies, all instars of Lemanniella minotauri were described along with this species’ feeding, mating, and dispersal biology (Wurst, 2001). The deutonymphs were found in low numbers (no more than two per ant) riding on the underside of the ants’ () heads. As feeding instars, mites ate black fungus that grew on the wood of the ant nest (Wurst, 2001).

Mites in the family Histiostomatidae have chelicerae modified into feathery filter feeding structures. They inhabit wet substrates and use their mouthparts to filter organic material and microorganisms. Some species of Anoetus remove pathogenic microorganisms from pollen and nectar in the nests of halictid bees in a mutualistic relationship (OConnor, 2009). In a study of the red imported (Solenopsis invicta), mites in the genus Histiostoma were found phoretic on the female alates in the colony. In most cases, they were found primarily on the gasters of these ants (Solokov et al., 2003).

It may be that Histiostoma are simply awaiting the death of their host as phoretic instars, since they have often been found feeding on decaying insect bodies (OConnor, personal communication).

Scope of these Studies

The current study is an effort focused on the overlooked Acarine inhabitants of ant nests. In Chapter 2 the reader will find work focused on the state of Ohio, one which has yet to be mentioned in any myrmecophilous mite studies. The objectives of Chapter

2’s study are as follows: 1) to provide a robust list of the associated mite fauna, 2) to investigate host specificity, 3) to document phoretic attachment sites, 4) to determine

22 preference of mites for sex of the host ant, 5) to determine mite roles and functions within the ant nest whenever possible, and 6) to statistically analyze the possible ecological factors influencing mite abundance, prevalence, and species richness in ant colonies.

Chapter 3 focuses on a smaller, initial study based in Arizona concerning the mites associated with alates and dealates of a single common species of harvester ant,

Messor pergandei. The objectives of this study are similar to those of Chapter 2: 1) to identify associated mite fauna, 2) to investigate attachment site specificity, 3) to determine mite preference for sex of the host, 4) to determine the prevalence of infestations for alates and dealates, 5) to collect and identify mites from chaff piles, and

6) to observe the mites in culture by rearing the dealates.

Chapter 4 provides: 1) a summary of the findings presented in the two studies, 2) implications of the results, 3) distinctiveness of the studies, 4) suggestions for future research, and 5) applications for the study of mite fauna from ant nests.

23

Literature Cited

Aoki, J., G. Takaku, and F. Ito. (1994). Aribatidae, a new myrmecophilous oribatid mite family from Java. International Journal of Acarology, 20: 3-10.

Balazy, S. and J. Wisniewski. (1982). Two species of entomopathogenic fungi on myrmecophilic mite coccinea (Michael, 1891) (Acari: Uropodina). Bulletin de ’Academie Polonaise des Sciences, 30: 81-84.

Banks, N. (1915). Two Mexican myrmecophilous mites. Psyche, 22: 60-61.

Banks, N. (1916). Acarians from Australian and Tasmanian ants and ant-nests. Proceedings of the Royal Society of South , 40: 224-40.

Berghoff, S. M. and N. . Franks. (2007). First record of the army ant Cheliomyrmex morosus in Panama and its high associate diversity. Biotropica, 39: 771-773.

Berlese, A. (1904). Illustrazione iconografica degli Acari mirmecofili. Redia, 1903: 299- 474.

Berlese, A. (1916). Centuria seconda di acari nuovi. Redia, 12: 125-180.

Beattie, A. J. and . C. Culver. (1983). The nest chemistry of two seed-dispersing ant species. Oecologia, 56: 99-103.

Boulton, A. M., B. A. Jaffee, and K. M. Scow. (2003). Effects of a common harvester ant (Messor andrei) on richness and abundance of soil biota. Applied Soil Ecology, 23: 257-265.

Bruce, W. A. and G. L. LeCato. (1980). tritici: a potential new agent for biological control of the red imported fire ant, Solenopsis invicta (Acari: ). International Journal of Acarology, 6: 271-274.

Cole, B. J. (1994). Nest architecture in the western harvester ant, Pogonomyrmex occidentalis (Cresson). Insectes Sociaux, 41: 401-410.

24

Collins, H.L. and G. P. Markin. (1971). Inquilines and other arthropods collected from nests of the imported fire ant Solenopsis saevissima richteri. Annals of the Entomological Society of America, 64: 1376-1380.

Conway, J. (2003). Architecture, population size, myrmecophiles and mites in an excavated nest of the , mendax Wheeler, in Arizona. The Southwestern Naturalist, 48: 449-450.

Cross, E. A. (1965). The generic relationships of the family Pyemotidae (Acarina: ). University of Kansas Science Bulletin, 45: 29-275.

Cross, E. A. (1974). A new genus and two new species of Pyemotid mites phoretic upon insects (Acarina: Tarsonemoidea). Acarologia, 16: 677-683.

Delfinado, M. D. and E. W. Baker. (1976). New species of Scutacaridae (Acarina) associated with insects. Acarologia, 28, 264-301.

Dietemann, V., C. Peeters, J. Liebig, V. Thivet, and B. Hölldobler. (2003). Cuticular hydrocarbons mediate discrimination of reproductives and nonreproductives in the ant gulosa. Proceedings of the National Academy of Sciences, 100: 10341-10346.

Donisthorpe, H. St. J. K. (1927). The guests of British ants their habits and life-histories. London, George Routledge and Sons, Ltd.

Ebermann, E. (1979). Ein beitrag zur erforschung der milben-family Scutacaridae (Acari- Trombidiformes) in Osterreich und angrenzenden Gebieten. Carinthia II, 89: 259- 280.

Ebermann, E. (1981). Imparipes (Imparipes) brevitarsus n.sp. (Acari, Scutacaridae), eine mit ameisen vergesellschaftete neue milbenart aus Österreich. Mitteilungen des Naturwissenschaftlichen Vereines für Steiermark, 111: 183-192.

Ebermann, E. & M. Hall. (2003). First record of sporothecae within the mite family Scutacaridae (Acari, Tarsonemina). Zoologischer Anzeiger, 242: 367-375.

Ebermann, E. & M. Hall. (2004). A new species of scutacarid mites transferring fungal spores (Acari, Tarsonemina). Revue Suisse de Zoologie, 111: 941-950.

Ebermann, E. and J. C. Moser. (2008). Mites (Acari: Scutacaridae) associated with the red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicdae), from Louisiana and Tennessee, U.S.A. International Journal of Acarology, 34: 55-69.

25

Ebermann, E. and G. Rack. (1982). Zur biologogie einer neuen myrmecophilen art der gattung Petalomium (Acari, Pygmephoridae). Entomologische Mitteilungen aus dem Zoologiischen Museum Hamburg, 115: 175-191.

Eickwort, G. C. (1990). Asociations of mites with social insects. Annual Review of Entomology, 35: 469-488.

Elzinga, R. J. (1978). Holdfast mechanisms in certain Uropodine mites (Acarina: Uropodina). Annals of the Entomological Society of America, 71: 896-900.

Elzinga, R. J. (1981). The generic status and six new species of Trichocylliba (Acari: Uropodina). Acarologia, 23: 3-18.

Elzinga, R. J. (1982a). The genus Coxequesoma (Acari: Uropodina) and descriptions of four new species. Acarologia, 23: 215-224.

Elzinga, R. J. (1982b). The genus Antennequesoma (Acari: Uropodina) and descriptions of four new species. Acarologia, 23: 321-325.

Elzinga, R. J. (1989). Habeogula cauda (Acari: Uropodina), a new genus and species of mite from the army ant Labidus praedator (F. Smith). Acarologia, 30: 341-344.

Elzinga, R. J. (1990). Two new species of Planodiscus (Acari: Uropodina), range extensions and a synonomy within the genus. Acarologia, 31: 229-233.

Elzinga, R. J. (1991). Two new elongate species of Planodiscus (Acari: Uropodina) with a key to the known species. Acarologia, 32: 109-114.

Elzinga, R. J. (1993). Larvamimidae, a new family of mites (Acari: Dermanyssoidea) associated with army ants. Acarologia, 34: 95-103.

Elzinga, R. J. (1995). Six new species of Trichocylliba (Acari: Uropodina) associated with army ants. Acarologia, 36: 107-115.

Elzinga, R. J. (1998). A genus and five new species of mites (Acari: ) associated with army ants (Hymenoptera: Formicidae). Sociobiology, 31: 351-361.

Elzinga, R. J. and C.W. Rettenmeyer. (1966). A neotype and new species of Planodiscus (Acarina: Uropodina) found on Doryline ants. Acarologia, 8: 191-199.

Elzinga, R. J. and C. W. Rettenmeyer. (1970). Five new species of Planodiscus (Acarina: Uropodina) found on Doryline ants. Acarologia, 12: 59-70.

Elzinga, R. J. and C. W. Rettenmeyer. (1974). Seven new species of Circocylliba (Acarina: Uropodina) found on army ants. Acarologia, 16: 595-611.

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Estebanes-Gonzalez, M. L. (2000). A redescription of Forcellinia faini Delfinado-Baker and Baker, 1989 (Acari: Acaridae). Folia Entomologica Mexicana, 109: 1-6.

Fain, A. (1987). Observations on the hypopi of the genus Forcellinia Oudemans, 1924 (Acari, Acaridae). Bulletin de L’Institut Royal des Sciences Naturelles de Belgique, 57: 111-120.

Fain, A. and W. Chmielewski. (1987). The phoretic hypopi of two acarid mites described from ants’ nest: formicitorum Volgin, 1948 and Lasioacarus nidicolus Kadzhaja and Sevastianov, 1967. Acarologia, 28: 53-61.

Fashing, N. J. (1976). The evolutionary modification of dispersal in Naiadacarus arboricola Fashing, a mite restricted to waterfilled treeholes (Acarina: Acaridae). American Midland Naturalist, 95: 337-346.

Franks, N. R., K. J. Healey, and L. Byrom. (1991). Studies on the relationship between the ant ectoparasite Antennophorus grandis (Acarina: Antennophoridae) and its host Lasius flavus (Hymenoptera: Formicidae). Journal of the Zoological Society (London), 225: 59-70.

Gorirossi-Bourdeau, F. (1993). Redescription of Oplitis paradoxa (Canestrini & Berlese, 1884), and the description of Oplitis farrieri, a new species (Mesostigmata: Uropodina: Oplitidae). Bulletin et Annals de la Societe Royale Belge d’ Entomologie, 129: 359-394.

Gotwald, W. H., Jr. (1995). Army Ants: The Biology of Social , Ithaca: Cornell University Press.

Gotwald, W. H., Jr. (1996). Mites that live with army ants: a natural history of some myrmecophilous hitch-hikers, browsers, and parasites. Journal of the Kansas Entomological Society, 69: 232-237.

Gray, B. (1974). Associated fauna found in nests of Myrmecia (Hymenoptera: Formicidae). Insectes Sociaux, 21: 289-300.

Greene, M. J., and D. M. Gordon. (2003). Cuticular hydrocarbons inform task decisions. Nature, 423: 32.

Haller, G. (1877). Antennophorus uhlmanni ein neuer Gamaside. Archiv für Naturgeschichte, 43: 57-62, pl. V.

Hermann, H. R. (1979). Insect sociality- An introduction. In H. R. Hermann, (Ed.), Social insects: Vol 1 (pp. 1-33), New York: Academic Press.

27

Hermann, H. R., M. S. Blum, and A. N. Hunt. (1970). Myrmecophilous arthropods associated with the imported fire ant, Solenopsis saevissima (Hymenoptera: Formicidae). Louisiana Academy of Sciences, 33: 13-18.

Hirschmann, W. (1973). Gangsystematik der Parasitiformes Teil 170-172. Acarologie, Schriftenreihe für vergleichende Milbenkunde, 19: 121-124.

Hölldobler, K. (1928). Zur biologie der diebischen Zwergameise (Solenopsis fugax) und ihrer Gäste. Biologisches Zentralblatt, 48: 129-142.

Hölldobler, B. (1971). Communication between ants and their guests. Scientific American, 224: 86-93.

Hölldobler, B. and E. . Wilson. (1990). The Ants, Cambridge: Harvard Press.

Hölldobler, B. and E. O. Wilson. (2009). The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies, New York: W. W. Norton & Company.

Houck, M. and B. M. OConnor. (1991). Ecological and evolutionary significance of phoresy in the Astigmata. Annual Review of Entomology, 36, 611-636.

Hull, J. E. (1923). New myrmecophilous Gamasids. Annals and Magazine of the Natural history of London, 12: 610-616.

Hunter, P. E. (1961). The genus Laelaspis with descriptions of three new species (Acarina: Laelaptidae). Annals of the Entomological Society of America, 54: 672- 683.

Hunter, P. E. (1964). Three new species of Laelaspis from North America (Acarina: Laelaptidae). Journal of the Kansas Entomological Society, 37: 293-301.

Hunter, P. E. (1967). Comments on Hypoaspis (Gymnolaelaps) Berlese, 1916, with descriptions of new species. (Acarina: ; Laelapinae). Journal of the Entomological Society, 2: 99-102

Hunter, P. E. (1993). Two new genera, Crassoseta and Brachysternopsis of the family (Acari, Mesostigmata, Trigynaspida. Zoologica Scripta, 22: 91-99.

Hunter, P. E. and M. Costa (1971). Description of Gymnolaelaps shealsi n. sp. (Acarina: Mesostigmata) associated with the imported fire ant. Journal of Georgian Entomological Society, 6: 51-53.

Hunter, P. E. and R. Davis. (1962). Two new species of Laelaspis mites. Proceedings of the Entomological Society of Washington, 64: 247-252.

28

Hunter, P. E. and M. H. Farrier. (1975). Mites in the genus Oplitis Berlese (Acarina: ) associated with ants (Hymenoptera: Formicidae) in the southeastern United States Part I. Acarologia, 17: 595-623.

Hunter, P. E. and M. H. Farrier. (1976). Mites in the genus Oplitis Berlese (Acarina: Uropodidae) associated with ants (Hymenoptera: Formicidae) in the southeastern United States Part II. Acarologia, 18: 20-50.

Hunter, P. E. and C. A. Hunter (1963). The genus Myrmonyssus with descriptions of two new species (Acarina: Laelaptidae). Acarologia, 5: 335-341.

Hunter, P.E. and R. M. T. Rosario. (1988). Associations of Mesostigmata with other arthropods. Annual Review of Entomology, 33: 393-417.

Ito, F. and J. Aoki. (2003). Bionomics of the myrmecophilous oribatid mite Protoribates myrmecophilus (Acari: Oribatida) in the Oriental tropics. Journal of Natural History, 37: 2383-2391.

Jacot, A. P. (1939). A new Antennophorid mite, rider of the Philippine ant Aenictus martini. The Philippine Journal of Science, 69: 433-434.

Janet, C. M. (1897a). Sur les rapports de l’Antennophorus uhlmanni Haller avec le Lasius mixtus Nyl. Compte Rendus de l’Academie des Sciences, Paris, 124: 583-585.

Janet, C. M. (1897b). Sur les rapports du Discopoma comata Berlese, avec le Lasius mixtus Nylander. Comptes Rendus Hebdomadaires des Séances de l’Academie des Sciences, 124: 102-105.

Janet, C. M. (1902). Sur le Lasius mixtus, l’Antennophorus uhlmanni, etc. Etudes sur les Fourmis, les guêpes et les abeilles Note 13. Limoges, 1897: 84-98.

Kaliszewski, M., F. Athias-Binche, and E. E. Lindquist. (1995). Parasitism and parasitoidism in Tarsonemina (Acari: Heterostigmatina) and evolutionary considerations. Advances in Parasitology, 35: 335-367.

Karafiat, H. (1959). Systematik und ökologie der Scutacariden. In H. J. Stammer (Ed.), Beiträge zur Systematik und Ökologie Mitteleuropaischen Acarina, I: 627-712.

Karawajew, W. (1906). Weitere beobachtungen uber arten der gattung Antennophorus. Zapiski Kievskago Obscestva estestvoispytatelej, 20: 209-230.

Karawajew, W. (1909). Myrmecophilen aus Transkapien. Russkoe Entomologicheskoe Obozrenie, 3: 227-237.

29

Khaustov, A. A. (2005). New species and records of the genus Petalomium (Acari: Heterostigmatina: Pygmephoridae) from Crimea (). Acarina, 12: 173-179.

Khaustov, A. A. and P. R. Chydyrov. (2004). New species of mites of the family Scutacaridae (Acari: Heterostigmatina) associated with ants (Hymenoptera, Formicidae) from Turkmenistan. Acarina, 12: 87-103.

Khaustov, A. A. and J. C. Moser. (2008). Two new species of mites of the genera Petalomium Cross and Caesarodispus Mahunka (Acari: Heterostigmatina: Neopygmephoridae, Microdispidae) associated with Solenopsis invicta Buren (Hymenoptera: Formicidae) from the U.S.A. International Journal of Acarology, 34: 115-121.

Kistner, D. H. (1979). Social and evolutionary significance of social insect symbionts. In H. R. Hermann (Ed.), Social insects: Vol 1 (pp. 340-413). New York: Academic Press.

Kistner, D. H. (1982). The social insects bestiary. In H. R. Hermann (Ed.), Social insects: Vol 3 (pp. 2-244), New York: Academic Press.

Klompen, H., M. Lekveishvili, and W. C. Black IV. (2007). Phylogeny of parasitiform mites (Acari) based on rRNA. Molecular Phylogenetics and Evolution, 43: 236- 251.

Kneissl, L. (1907). wasmanni Kneissl Eine neue myrmekophile milbe. Zeitschrift fur wissenschaftliche Insektenbiologie, 3: 190-191.

Krantz, G. W. (1962). Descriptions of three myrmecophilous Macrochelidae (Acarina: Mesostigmata from Panama, British Guiana and the British West Indies. Journal of the Kansas Entomological Society, 35, 349-357.

Krantz, G. W. (2009). Introduction. In G. W. Krantz & D. E. Walter (Eds.), A Manual of Acarology (3rd ed.), (pp. 1-2). Lubbock, TX: Tech University Press.

Krantz, G. W. and D. E. Walter. (2009). Classification. In G. W. Krantz & D. E. Walter (Eds.), A Manual of Acarology (3rd ed.), (pp. 99-103). Lubbock, TX: Texas Tech University Press.

Krantz, G. W., L. A. Gomez, and V. E. Gonzalez (2007). Parasitism in the Uropodina: a case history from Colombia. Acarology XI: Proceedings of the International Congress, 29-38.

Laakso, J. and H. Setälä. (1998). Composition and trophic structure of detrital food web in ant nest mounds of Formica aquilonia and in the surrounding forest soil. Oikos, 81: 266-278.

30

Lahav, S., V. Soroker, and A. Hefetz. (1999). Direct behavioral evidence for hydrocarbons as ant recognition discriminators. Naturwissenschaften, 86: 246- 249.

Le Breton, J., G. Takaku, and K. Tsuji. (2006). Brood parasitism by mites (Uropodidae) in an invasive population of the pest-ant Pheidole megacephala. Insectes Sociaux, 53: 168-171.

Lehtinen, P. T. (1987). Association of Uropodid, Prodinychid, Polyaspidid, Antennophorid, Sejid, Microgynid, and Zerconid mites with ants. Entomologisk tidskrift, 108: 13-20.

Lenoir, A., P.D’Ettore, and C. Errard. (2001). Chemical ecology and social parasitism in ants. Annual Review of Entomology, 46: 573-599.

Liebig, J., C. Peeters, N. J. Oldham, C. Markstadter, and B. Hölldobler. (2000). Are variations in cuticular hydrocarbons of queens and workers a reliable signal of fertility in the ant Harpegnathos saltator?. Proceedings of the National Academy of Sciences, 97: 4124-4131.

Lin, N. and C. D Michener. (1972). Evolution of sociality in insects. The Quarterly Review of Biology, 47: 131-159.

Lindquist, E. E., G. W. Krantz, and D. E. Walter. (2009). Order Mesostigmata. In G. W. Krantz & D. E. Walter (Eds.), A Manual of Acarology (3rd ed.), (pp. 124-232). Lubbock, TX: Texas Tech University Press.

Mahunka, S. (1961). Contributions to the Tyrogphid fauna of Hungary (Acari). Annales Universitatis Scientiarum Budapestinensis de Rolando Eötvös nominatae, 4: 113- 118.

Mahunka, S. (1965). Zwei neue milben-arten aus der gruppe Tarsonemini (Acari). Zoologischer Anzeiger, 174: 156-160.

Mahunka, S. (1967). Beiträge zur kenntnis der Tschechoslowakischen Tarsonemini- fauna. Věstník Československé Společnosti Zoologické, 31: 240-244.

Mahunka, S. (1968). Beiträge zur Tarsonemini-fauna Ungarns V. (Acari, Trombidiformes). Rovartani Közlemények Folia Entomologica Hungarica, 21: 125-136.

Mahunka, S. (1970a). Two new Scutacarid mites (Acari, Tarsonemina) from Pogonomyrmex occidentalis (Hymenoptera) in the United States of America. Parasitologia Hungarica, 3: 87-96.

31

Mahunka, S. (1970b). Zwei neue Heterodispus- Arten und einige interessante in Ameisenhaufen lebende milben aus Ungarn (Acari: Tarsonemina). Folia Entomologica Hungarica, 23: 313-331.

Mahunka, S. (1970c). Considerations on the systematics of the Tarsonemina and the description of new European taxa (Acari: Trombidiformes). Acta Zoologica Academiae Scientiarum Hungaricae, 16, 137-174.

Mahunka, S. (1977a). The examination of myrmecophilous Tarsonemid mites based on the investigations of Dr. C. W. Rettenmeyer (Acari) I. Acta Zoologica Academiae Scientiarum Hungaricae, 23: 99-132.

Mahunka, S. (1977b). The examination of myrmecophilous Tarsonemid mites based on the investigations of Dr. C. W. Rettenmeyer (Acari) II. Acta Zoologica Academiae Scientiarum Hungaricae, 23: 341-370.

Mahunka, S. (1977c). Neue und interessante milben aus dem Genfer Museum XIX. Einige angaben zur kenntnis der milbenfauna der ameisen-nester (Acari: Acarida, Tarsonemida). Archives des Sciences, 1: 91-106.

Mahunka, S. (1978). The examination of myrmecophilous Acaroidea mites based on the investigations of Dr. C. W. Rettenmeyer (Acari: Acaroidea) I. Acta Zoologica Academiae Scientiarum Hungaricae, 31: 135-166.

Mahunka, S. (1979). The examination of myrmecophilous Acaroidea mites based on the investigations of Dr. C. W. Rettenmeyer (Acari: Acaroidea) II. Acta Zoologica Academiae Scientiarum Hungaricae, 25: 311-342.

Mahunka, S. (1982). Zwei neue Scutacariden-arten aus Mexico (Acari: Tarsonemina). Miscellanea Zoologica Hungarica, 1: 133-138.

Mahunka, S. and L. Mahunka-Papp. (1980). Beiträge zur Kenntnis der europäischen myrmecophylen Tarsoneminen (Acari). I. Folia Entomologica Hungarica, 33: 283-292.

Mann, W. M. (1913). Literature for 1912 on the behavior of ants and myrmecophiles. Journal of Behavior, 3: 429-445.

Mašán, P. (1992). A new species of the Gamasid mite from the genus Hypoaspis canestrini, 1884 (Acarina: Parasitiformes) from Czecho-. Biologia (Bratislava), 47: 383-388.

Mašán, P. (1999). Description of the deutonymph of Trichocylliba comata (Acarina, Mesostigmata, Uropodina). Biologia, Bratislava, 54: 525-527.

32

Masuko, K. (1984). Studies on the predatory biology of Oriental dacetine ants (Hymenoptera: Formicidae) I: some Japanese species of Strumigenys, Pentastruma, and Epitritus, and a Malaysian Lapidogenys with special reference to hunting tactics in short-mandibulate forms. Insectes Sociaux, 31: 429-451.

Masuko, K. (1994). Specialized predation on oribatid mites by two species of the ant genus Myrmecina (Hymenoptera: Formicidae). Psyche, 101: 159-173.

Masuko, K. (2009). Studies on the predatory biology of Oriental dacetine ants (Hymenoptera: Formicidae) II. Novel prey specialization in Pyramica bente. Journal of Natural History, 43: 825-841.

Metawali, S. H. (1981). Studies on Tarsonemini (Acarina) associated with ants in forests in Poland. Entomologische mitteilungen aus dem zoologischen museum Hamburg, 7: 87-100.

Michael, A. D. (1891). On the association of Gamasidae with ants. Proceedings of the Zoological Society of London, 1981: 638-654.

Moniez, E. (1892). Memoirs sur quelques acariens et Thysanoures parasites commensaux des fourmis. Revue biologique du nord de la , 4: 1-12.

Moniez, E. (1894). Sur quelques arthropods trouves dans des fourmilieres. Revue biologique du nord de la France, 6: 201-215.

Neece, K. C. and D. P. Bartell. (1982). A Faunistic Survey of the Organisms Associated with Ants of Western Texas, No. 25. Graduate Studies Texas Tech University. Lubbock, TX: Texas Tech University Press, 36p.

Norton, R. A. and V. M. Behan-Pelletier. (2009). Suborder Oribatida. In G. W. Krantz & D. E. Walter (Eds.), A Manual of Acarology (3rd ed.), (pp. 430-564). Lubbock, TX: Texas Tech University Press.

OConnor, B. M. (1982). Evolutionary ecology of astigmatid mites. Annual Review of Entomology 27, 385-409.

OConnor, B. M. (2009) Cohort . In G. W. Krantz & D. E. Walter (Eds.), A Manual of Acarology (3rd ed.), (pp. 565-657). Lubbock, TX: Texas Tech University Press.

Paris, C. (2008). Litter decomposition and soil organisms within and outside of Camponotus punctulatus nests in sown pasture in Northeastern Argentina. Applied Soil Ecology, 40: 271-282.

Park, O. (1932). The myrmecocoles of Lasius umbratus mixtus aphidicola Walsh. Annals of the Entomological Society of America, 25: 77-88. 33

Porter, S. D. (1985). Masoncus spider: A miniature predator of Collembola in harvester ant colonies. Psyche, 92: 145-150.

Rack, G. (1964). Scutacaridae von Hamburg (Acarina, Trombidiformes). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 61: 185-194.

Rack, G. (1968). Milben als hausungeziefer u. vorratsschädlinge in Nordwestdeutschland. Entomologische Mitteilungen aus dem Zoologischen Staatsintitut u. Zoologischen Museum Hamburg, 3: 1-20.

Raczkowski, J. M. (2008). Phylogeny and founding stage of Lasius (Acanthomyops) (Hymenoptera: Formicidae). Ph. D. Dissertation, The Ohio State University, Columbus, OH. 172p.

Rettenmeyer, C.W. (1962a). Arthropods associated with neotropical army ants with a review of the behavior of these ants (Arthropoda; Formicidae: ). Ph.D. Dissertation. University of Kansas, Lawrence, . 644p.

Rettenmeyer, C.W. (1962b). The diversity of arthropods found with neotropical army ants and observations on the behavior of representative species. Proceedings of the North Central Branch-Entomological Society of America, 17: 14-15.

Rettenmeyer, C. W. (1962c). Notes on host specificity and behavior of myrmecophilous Macrochelid mites. Journal of the Kansas Entomological Society, 35: 358-360.

Rosario, R. M. T. and P. E. Hunter. (1988). The genus Berlese, with descriptions of two new species (Acari: Mesostigmata: Laelapidae). The Journal of Parasitology, 74: 466-470.

Ross, L. J. and E. A. Cross. (1979). A revision of the genus Acinogaster (Acari: Pygmephoridae). International Journal of Acarology, 5: 231-250.

Samšiňák, K. (1960). Ueber einige myrmekophile milben aus der family Acaridae. Časopis Československé Společnosti Entomologické, 57: 185-192.

Saboori, A., A. A. Khaustov, M. Hakimitabar, H. Hjiqanbar. (2009). A new genus and species of larval Erythraeinae (Acari: Prostigmata: ) from Ukraine and the taxonomic state of Zhangiella. Zootaxa, 2203: 22-30.

Sellnick, M. (1926). Alguns novos Acaros (Uropodidae) myrmecophiles e termitophilos. Archivos do Museu Nacional do Rio de Janiero, 26: 29-56.

Sevastinov, V. D. (1965). On the number of mites carried and exterminated by the ant Lasius fuliginosus (in Russian). Zoologicheskii Zhurnal, 44: 1651-1660.

34

Shaw, M. D. and O. D. Seeman. (2009). Two new species of Myrmozercon (Acari: Laelapidae) from Australian ants (Hymenoptera: Formicidae). Zootaxa, 2025: 43- 55.

Silvestri, F. (1912). Contribuzioni alla conoscenza dei mirmecofili. Bollettino del laboratorio di zoologia generale e agrarian della facoltá agraria in portici, 6: 222-245.

Sokolov, I. M., . Y. Sokolova, and J. R. Fuxa. (2003). Histiostomatid mites (Histiostomatidae: Astigmata: Acarina) from female reproductives of the red imported fire ant (Hymenoptera: Formicidae). Journal of Entomological Science, 38: 699-702.

Southcott, R. V. (1989). Larval (Acarina: Erythraeidae) ectoparasitic on a North American ant. Acarologia, 30: 241-248.

Štorkán, J. (1940). Myrmecophilní Acari Čech. Vestnik Ceskoslovenske zoologicke spolecnosti v Praze, 8: 166-172.

Travis, B. V. (1941). Notes on the biology of the fire ant Solenopsis geminata (F.) in Florida and Georgia. The Florida Entomologist, 24: 15-22.

Tschinkel, W. R. (2003). Subterranean ant nests: trace fossils past and future? Palaeogeography, Palaeoclimatology, Palaeoecology, 192: 321-333.

Tschinkel, W. R. (2004). The nest architecture of the Florida harvester ant, Pogonomyrmex badius. Journal of Insect Science, 4: 1-19.

Vitzthum, H. G. (1919). Neue myrmecophiles milben. Zoologischer Anzeiger, 50, 188- 192.

Vitzthum, H. G. (1930). Ein Ameisengast (Acar.). Deutsche Entomologische Gesellschaft, 1: 89-94.

Wagner, D., J. F. Brown, and D. M. Gordon. (1997). Harvester ant nests, soil biota and soil chemistry. Oecologia, 112: 232-236.

Walter, D. E. (2003). A new mite from an arboreal ant (Formicidae: Polyrachis sp.): Myrmozercon iainkayi n. sp. (Mesostigmata: Laelapidae). International Journal of Acarology, 29: 81-85.

Walter, D. E., E. E. Lindquist, I. M. Smith, D. R. Cook, and G. W. Krantz. (2009). Order Trombidiformes. In G. W. Krantz & D. E. Walter (Eds), A Manual of Acarology (3rd ed.), (pp. 233-420). Lubbock, TX: Texas Tech University Press.

35

Wasmann, E. (1897). Über einige myrmecophile acarinen. Zoologischer Anzeiger, 531: 170-173, 346-350.

Wasmann, E. (1899). Weitere nachträge zum verzeichniss der ameisengäste von Holländisch Limburg. Tijdschrift voor Entomologie, 42: 158-171.

Wasmann, E. (1902). Zur Kenntnis der myrmecophilen Antennophorus und anderer auf ameisen und termiten reitender acarinen. Zoologischer Anzeiger, 25: 66-76.

Weber, N. A. (1972). Gardening ants the Attines. Philadephia, The American Philosophical Society.

Wheeler, W. M. (1910a). Ants: Their structure, development and behavior. New York, Columbia University Press.

Wheeler, W. M. (1910b). Two new myrmecophilous mites of the genus Antennophorus. Psyche, 17: 1-7.

Wilson, E. O. (2005). Oribatid mite predation by small ants of the genus Pheidole. Insectes Sociaux, 52: 263-265.

Wing, M.W. (1968) Taxonomic revision of the Nearctic genus Acanthomyops (Hymenoptera: Formicidae). Cornell University Agricultural Experiment Station Memoir, 405: 1-173.

Wiśniewski, J. (1979). Stadien von 2 neuen (Uroseius)- arten aus ameisennestern Polens (Uropodini, Uropodinae). Acarologie, 25: 43-46 + pl 6.

Wiśniewski, J. (1980). Gang einer neuen mit spatulifera (Moniez 1892) verwandten art aus ameisennestern Polens (Trichouropodini, Uropodinae). Acarologie, 27: 8-10 + pl 1.

Wiśniewski, J. & W. Hirschmann. (1983). Stadium einer neuen Dinychus- artaus ameisennest in Polen (Dinychini, Uropodinae). Acarologie, 30: 133-134.

Wiśniewski, J. & W. Hirschmann. (1992). Gangsystematische studie von 3 neuen Antennophorus-arten aus Polen (Mesostigmata, Antennophorina). Acarologia, 33: 233-244.

Witte, V., A. Leingärtner, L. Sabaß, R. Hashim, and S. Foitzik. (2008). Symbiont microcosm in an ant society and the diversity of interspecific interactions. Animal Behavior, 76: 1477-1486.

Womersley, H. (1958). Some new or little known Mesostigmata (Acarina) from Australia, New Zealand and Malaya. Transactions of the Royal Society of South Australia, 81: 115-130. 36

Wurst, E. (2001). The life cycle of Lemanniella minotauri n. sp. and the erection of the new family Lemannillidae (Acari: Astigmata). Stuttgarter Beiträge zur naturkunde, Serie A (Biologie), 621: 1-34.

Zeman, P. (1982). Two new species of Hypoaspidinae (Acari: Mesostigmata: Dermanyssidae) associated with ants. Věstnik Československé Společnosti Zoologické (Praha), 46: 231-237.

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Chapter 2: Mites Associated with Ants in Ohio

Introduction: Goals, Questions, and Scope of this Project

As detailed in Chapter 1 very few publications concerning ant associated mites have provided details beyond basic descriptions. Most studies have focused on specific mite groups or a particular ant species. Further, many mites have been described from

European ants, while North American ants have received little attention. The study in this chapter is an attempt to exceed the breadth of the typical ant mite studies by executing a robust examination of all mites associated with numerous ant species while also including ecological observations and analyses of mite diversity.

The only broad scale survey focused specifically on ant mites in the United States was conducted by Hunter and Farrier (1975; 1976) in the southeastern United States.

They collected Oplitis species (using kerosene extraction methods) associated with 14 ant species, 5 of which are also present in Ohio. Neece and Bartell (1982) conducted a survey in western Texas focused primarily on beetles and large arthropods, but they found approximately 10 species of mites associated with 10 ant species. The majority of these

“associated” species were large mites in the (Trombidiformes:

Prostigmata) a group which does not have any true association with ants but is easily detected as they walk on the soil surface due to bright coloration. The remainder of the specimens was comprised of large mesostigmatid mites, only a couple of which may have

38 true associations with ants (for example an Oplitis species). None of the Astigmata collected were identified and no Heterostigmatina were recovered at all.

Rettenmeyer’s (1962) survey of army ants in Panama stands out as a colossal stride in the knowledge of ant mites and their habits, hosts relations, and diversity, but does not paint a realistic picture of mites associated with most of the ants in North

America, let alone Ohio, which has only 1 rare species of legionary ant (Neivamyrmex carolinensis) recorded (Coovert, 2005). Rettenmeyer’s (1962) work and suggestions therein were very useful when formulating my own ant mite study, however. For example to avoid the issues Rettenmeyer noted concerning loss of small mite species and undocumented attachment site locations, live ants were individually inspected (details provided in “Sampling Phoretic mites from Ants” section).

The state of Ohio resides in the Midwestern United States and acts as an intersection for many ecological zones including the Appalachian Mountains in the southeast, wetlands and beaches to the north along the Great Lakes, and to the west. Although the natural landscape is predominantly deciduous forest throughout the state, urbanization and agriculture have altered and fragmented much of the natural habitat. Ants in Ohio comprise 7 subfamilies, and 128 species belonging to 33 genera

(Ivanov, 2010). The most common genera are Formica, Camponotus, Lasius and

Aphaenogaster. The distribution of Ohio ant species spans much of the Northeastern

United States and many species are common throughout the entire United States

(Coovert, 2005). Some are introduced and common disturbance specialists such as

39

Tetramorium caespitum. Others such as Camponotus pennsylvanicus and are common house pests.

The study which follows gave equal representation to all associated mite lineages and identified them to a level which allowed ecological roles within the nest to be more easily understood. A significant contribution of this study is the fact that it is the first to attempt to explain trends observed in ant mite species richness using ecological factors.

This study focuses on the following: 1) what mites (phoretic and non-phoretic) are associated with Ohio ant species?, 2) are the mites host specific?, 3) are the mites attachment site specific?, 4) are phoretic mites found more frequently on female alates than male alates?, 5) what ecological factors influence phoretic mite prevalence and species richness in ant colonies?, and 6) what roles do the non-phoretic mites play in the colonies?

Materials and Methods

Collection Dates and Localities

Ants were collected from April through September in both 2008 and 2009 from

22 sites in Ohio, 1 site in Illinois, and 1 site in Kentucky. Three colonies were collected from 2 previously sampled localities in March 2010 to fill in knowledge gaps concerning uncommon ant species. Table 33 in Appendix B details the collection localities, dates, number of colonies collected at each locality, and associated field numbers (=colony codes). Collections from the 24 sites were made in open and forested habitats in both urban and undisturbed areas.

Colony Sampling Methods

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Searching for ant colonies at the designated localities involved flipping rocks or other objects on the surface of the soil, turning logs, peeling back the bark of logs or standing dead trees, and digging in mulch, mounds, and soil. All colonies belonging to any ant species were collected when encountered. In cases of medium to large nests

(>500 individuals), a portion of the nest was sampled by filling a 473 ml, 11.43cm diameter and 7.62 cm deep, (16 oz, 4.5 in diameter, 3 in deep) clear plastic container with ants and the natural substrate of the nest. Whenever possible, the larvae were collected along with workers. Alates and dealates were collected when they were found in colonies or at the site. In cases of very small colonies (<100 individuals) the entire colony could often be collected. Sometimes large-bodied ants such as Formica spp. or Camponotus spp. were collected in gallon-sized plastic zip bags in order to get a sufficient number of individuals. Each nest was collected separately, field identifications were written on the lid of the container along with a field number in the format “collector initials-year- monthday-colony number”.

Sampling Phoretic Mites from Ants

Samples were returned to the lab, and worker ants were inspected for mites.

While observing with a stereomicroscope at 25-50x magnification, each live ant specimen was individually placed in a glass well with 95% ethanol. The number of mites and their phoretic attachment sites were documented in a notebook as the ant died. If mites appeared to be different species or were found on different attachment sites on the host, they were collected using a minute wire loop or pipette, placed in separate glass (filled with lactophenol) for clearing prior to slide making and subsequent

41 identification, and labeled accordingly. If they appeared to be members of a series, some were placed in lactophenol while others were stored with the host in 95% ethanol. Wells in which the mites cleared were labeled with the field number (=colony code), host number, preliminary identification (usually to family), and attachment site location. For example, notes on the cover of a clearing mite that read, “KAU-09-0725-15 #7 gaster

Histio,” signifies a mite in the family Histiostomatidae was collected from the gaster of the 7th ant posessing mites from the fifteenth colony collected by the author on July 25th,

2009. At least 30 workers per colony typically were inspected individually, but fewer individuals were available when colonies were not populous. Total number of ants with and without mites for each colony was recorded to calculate prevalence, number infected hosts divided by number of hosts inspected (Margolis et al.,1982).

Alates were inspected individually (as described previously) in addition to the typical number of workers. The caste and sex of the alates was recorded along with the typical details concerning any mite specimens collected.

Dealates were sometimes found at the sites after a mating flight walking on the ground in search of a potential nest location or having just founded a colony with only a small number of workers. They were collected and placed in rearing tubes for potential non-phoretic mite observation. Their mites were generally not collected, but rather were documented as present or absent for the overall total prevalence calculations. Wandering dealates of Tetramorium caespitum were commonly encountered and all dealates collected in the day at the site were given a single field number, despite the fact that they probably came from different colonies.

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Collection and Rearing of Non-phoretic Mites

After sampling of the collection at the laboratory, the plastic holding container became a short-term nest for any remaining ants. Artificial ant food consisting of egg, honey, vitamins, and agar (as described by Bhatkar & Whitcomb, 1970) was provided for the ants on a small piece of aluminum foil as often as twice a week. A higher number of lab nests were cultured in this way in the 2008 season than for 2009.

Non-phoretic mites were collected by examining the nest substrate, ant brood, artificial food, fungus, and dead ants with the aid of a stereomicroscope. During a portion of the 2008 season, any remaining artificial ant food was examined for mites before new food was added. Fungus often had grown on the food, and any mites observed on the fungus were collected. Most of the mites were placed directly into lactophenol for slidemaking, but some rearing attempts were made. Small polystyrene cell wells

(Corning Inc., Corning, ) which contained a substrate of 9 parts Plaster of Paris and water to 1 part activated carbon were constructed for rearing the mites (similar to rearing containers described by Rohde, 1956). Once the substrate had dried, distilled water was used to moisten each well and a single mite (typically Astigmata found on the artificial ant food) was transferred into each rearing chamber. A small amount of ant food was provided for the mite on the substrate. Finally, a of Vaseline® was applied to the topmost edge and a coverslip was placed on top of the chamber to prevent escape.

Rearing chambers were placed into an incubator at 35º C and checked every couple of days for development of offspring. Individuals of all mite stages (larvae, protonymphs, deutonymphs, tritonymphs, and adult males and females), were then collected and placed

43 in lactophenol. Some second generation adults were placed into their own chambers for subsequent reproductive observations. In other cases, wandering Astigmatid deutonymphs were found in the plastic temporary nests (on the natural nest substrate) and were placed in rearing chambers to gain an understanding of subsequent life stages.

Mites walking in the nest were sometimes collected during food checks and placed in lactophenol. Ant larvae and pupae, if present in the nests, were also checked for mites. When colonies appeared to have only a few ants remaining, or if all ants were removed during initial inspection, the contents of the plastic container were put onto a

Berlese funnel, and any remaining arthropods were extracted into 75% ethanol. Only a portion of the Berlese samples were sorted for mites; most remain unsorted and stored in

75% ethanol in vials due to time constraints.

Preservation and Identification of Ants and Mites

After a few days in lactophenol, all cleared mite specimens were mounted on slides in Hoyer’s-Strandtmann’s medium and placed into a slide oven (approximately 45-

47° C) until dry. Using red enamel paint (Glyptal®), two coats of rings were applied to the dried slides to insulate and prevent crystallization of the medium. After rings were dry, mites on slides were sorted by host ant species and the number of associated morphospecies was determined. Characteristics such as shape, arrangement and length of setae, shape and length of gnathosoma, shape of coxal fields (Astigmata), patterning on dorsum or venter, and overall size were used for morphospecies determination.

Morphospecies were cross compared with all other morphospecies belonging to other ant species in order to determine the actual number of unique mite morphospecies and their

44 host ranges. Once all morphospecies were clearly distinguished from one another, each morphospecies was identified to genus using the following keys for designated mite groups:

 Mesostigmata- Krantz & Ainscough’s (1990) key for most Mesostigmata,

and Karg (1989), and Hirschmann (1993) for the Uropodina

 Astigmata- key to the families (OConnor, 2009) and unpublished key to

the genera for the Astigmata (OConnor, 2008)

 Heterostigmatina- Savulkina’s (1981) key for Pygmephoroids, Mahunka’s

(1970c) key for Pygmephoridae, Microdispidae, and Scutacaridae

 Prostigmata (non Heterostigmatid)- keys to the families of soil

Prostigmata (Walter et al., 2009)

Ants which had mites were placed into individual shell vials with 95% ethanol (in some cases spare mites were stored with them), labeled with the field number and ant number then plugged with cotton. Five shell vials containing 5 individual ants from the same sample were then placed into an ethanol filled (20 ml) scintillation vial for storage.

All ants from each colony without any mites were inserted into one shell vial, labeled with field number and “no mites” and stored in a scintillation vial, as well. One to four reference ant specimens were removed from the “no mites” vial for each colony (unless there were no ants in the colony without mites) and mounted on points, given full data labels, and placed in an insect box as a reference collection.

Ants were identified using Coovert’s key to the ants of Ohio (2005), and reference specimens were later verified by Gary Coovert. Outdated names were updated based on

45 the Ohio State University’s “Hymenoptera Name Server, version 1.5” (Johnson, 2007).

Representatives of each ant species are deposited as voucher specimens in the Ohio State

University Insect Collection under code numbers: OSUC0359951 to OSUC0359992 (see

Table 35). Voucher specimens of the mite species are deposited in at the Ohio State

University Acarology Collection. Specimen numbers are listed in Table 34.

Statistical Analyses of Collections

In order to more closely examine the questions posed in the introduction, statistical tests were conducted to determine the effects of ecological and phylogenetic factors on mite species richness and prevalence within colonies. Results were analyzed using PASW Statistics version 18.0 software in conjunction with the Ohio State

University’s Statistical Consulting Service. Independent-sample t-tests, one-way

ANOVA, and simple and multiple linear regression techniques were used to analyze the following phoretic mite response variables:

 Mite species richness- Total number of different phoretic mite species per colony

was tallied and also separated into major mite groups to form the response

variables: # of phoretic mite species, # of phoretic Mesostigmata, # of phoretic

Heterostigmatina, and # of phoretic Astigmata.

 Prevalence- number of infested hosts divided by the total number of hosts

inspected (Margolis et al., 1982).

Predictors include the ecological, phylogenetic and other factors described below:

 Total ants- number of ants inspected for each colony (workers, alates and

dealates). This factor was analyzed using a simple linear regression to determine

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if number of ants sampled determined the patterns seen in the response variables,

and thus whether sampling was inadequate. Lack of significance suggests

adequate sampling.

 Ant subfamily- variables are Formicinae (1), Myrmecinae (2), Dolichoderinae (3),

Ponerinae (4), Amblyoponerinae (5). Only two of the subfamilies had

representatives from more than one species (Formicinae and Myrmecinae), so a t-

test was used to determine statistical differences in mite species richness and

prevalence among colonies belonging to these two subfamilies.

 Social parasitism- ant colonies were coded as parasitic (1) or not parasitic (2).

These data were determined from records of mixed colonies and parasitic

interactions as listed by Raczkowski (2008) and Coovert (2005) (see Table 28). I

hypothesized that social parasitism may be an important factor increasing mite

richness in colonies exhibiting this behavior. As discussed in Chapter 1, several

species in the genus Lasius and one species collected in the genera Aphaenogaster

and also Formica found colonies as temporary social parasites. T-tests were used

for Lasius and Aphaenogaster parasitism because they had the most representative

colonies. I hypothesized that colonies formed by species through social parasitism

would have an increased number of mite species per colony (their own mites in

addition to any mites they accumulated from their ant hosts during colony

foundation).

 Genus Lasius- colonies coded as Lasius (1) or not Lasius (2). As described above,

I suspected that there may be something unusual occurring within the genus

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Lasius due to the fact that several species found their colonies parasitically. Based

on preliminary analyses, the genus seemed to be very influential in the data set, so

a closer examination of the effect of this genus was worth examining.

 Host habitat- colonies coded as woods, such as forest or forest edge (1) or open,

such as lawn, beach, or field (2). These data were gathered for each colony at the

time of collection. A t-test was used to determine differences in mite species

richness and prevalence in colonies in the woods and colonies in open

environments.

 Nest substrate- coded wood such as log, stump, bark or tree (1) or soil such as

lawn, pavement or curb, , or underneath items (2). These data were gathered

for each colony at the time of collection. A t-test was used to determine

differences in mite species richness and prevalence in colonies in wood and

colonies in soil substrates.

 Colony size- small (1), moderate (2), or large (3) number of individuals in a

typical colony as documented by Coovert (2005). Species that form small,

medium, and large colonies have up to 500 members, 500-2000 members, and

>2000 members, respectively. Colony sizes are not documented for some species

in the Coovert publication, so field observations of colony size were used (this

was usually only needed for very small colonies). I hypothesized that larger

colonies (which consequently would have more resources available) would be

more likely to have greater numbers of mite species than smaller colonies. I also

hypothesized that larger colonies would have more Mesostigmatid mites (as they

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are generally predatory and in need of larger food resources than other mite

groups) than smaller colonies. A one-way ANOVA was used with post hoc

comparisons using Tukey’s LSD method to determine differences in mite species

richness and prevalence for small, moderate, and large colonies.

 Host size- average ant size (mm) for each ant species as determined by Coovert

(2005). I hypothesized that larger ants would allow for more phoretic space and

therefore, more associated mites and more frequent mite infestations (prevalence).

Additionally I suspected a decrease in number of phoretic Mesostigmata as ant

size decreased. Very small ants simply may not be capable of carrying large

mesostigmatid mites. A simple linear regression was used to understand the

relationship between ant size and mite species richness and prevalence in

colonies.

All of these variables were tested using step-wise multiple linear regressions for inclusion into models explaining the response variables # of phoretic mites, # of

Mesostigmata, # of Heterostigmatina, # of Astigmata, and prevalence.

Colonies in which 20 or more individual ants had been inspected were used for analyses except 10 individuals were acceptable for Amblyopone pallipes, Ponera pennsylvanica, , punctiventris and Pyramica species, due to their extremely small colony sizes in natural conditions. A total of 175 colonies were used for the analyses unless otherwise indicated in the results section. The raw data used for analyses can be found in Appendix C (Table 37 to Table 39). An alpha level of 0.05 was used for all statistical tests of significance.

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Mite species accumulation curves were constructed from the accumulated number of observed mite species (Sobs Mao-Tau) per colony sampled. Sobs Mao-Tau was calculated using the diversity statistical program, EstimateS (Version 7.5; Colwell, 2005).

Results

Ant Species Collected and Number Inspected

A total of 43 species of ants (9093 individual ants) was collected and examined for mites. The collections comprised 273 ant colonies (78 in 2008, 192 in 2009, and 3 in

2010). Table 36 in Appendix B shows the number of colonies and individuals of each of the ant species observed for mites. Of the 43 ant species, 30 were found to have phoretic mites.

Mite Species Collected

A total of 1255 mite slides were identified to 151 morphospecies. 60 morphospecies (667 slides) belonged to the Astigmata, 56 morphospecies (452 slides) were members of the Heterostigmatina, and 35 morphospecies (135 slides) represented the Mesostigmata.

Almost all of the Astigmata belonged to two families, Acaridae (31 species, mostly and Forcellinia) and Histiostomatidae (27 species, all Histiostoma).

Two specimens looked distinctly different from the other slides, from each other, and did not fit the clear characteristics of Acarids or Histiostomatids, but were not identifiable to family due to poor slide quality. They were labeled Astig undet gen sp3 and sp4. A full list of mites and their associated ants is shown in Table 10 at the end of this chapter.

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Heterostigmatina consisted of three families: Pygmephoridae (9 species in the genera Petalomium and Bakerdania), Microdispidae (5 species in the genera

Caesarodispus and Unguidispus), and Scutacaridae (42 species in the genera Imparipes and Scutacarus). A full list of the heterostigmatid mites and their host species can be found in Table 11 at the end of this chapter.

Mesostigmata were represented by five families: Laelapidae (19 species),

Oplitidae (11 species), Uropodidae (2 species), Antennophoridae (2 species), and a single specimen of Rhodacaridae. The most common genera were Cosmolaelaps,

(Laelapidae) and Oplitis (Oplitidae). Four of the Oplitis species had been previously collected by Hunter and Farrier (1975; 1976) on the same ant species. See Table 12 at the end of this chapter for the full list of species and hosts.

Total Number of Mites per Ant Species

Thirty of the 43 species of ants had at least one phoretic mite species. Lasius umbratus had the highest number of associated mite species (38); mean number (M) of phoretic mite species per ant was 5.30 (SD =7.32, median = 2). Of the 13 ant species without associated mites, 4 species had fewer than 2 individual ants collected, and only 5 had more than 30 individuals collected. None of the species without associated mites had more than 3 colonies collected. Number of mites associated with each of the ant species is shown in Table 13 at the end of this chapter. Actual species identifications of mites associated with each ant (the inverse of Table 10 to Table 12) are shown in Table 14 at the end of this chapter.

Host Specificity

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The majority of mites were found on only one host ant species (68.9%). The percentage of mites associated with multiple species within the same genus was 25.2% and only 5.9% were found on ants belonging to multiple genera. A clearer view of host specificity can be seen when mite groups are analyzed separately. The Mesostigmata showed the most specificity with 91.4% having only one host species. The other 8.6% were found on ants in a single genus; none were found on multiple genera of ants.

Heterostigmatina also had very specific relationships with their hosts: 69.6% was found on a single host species, 26.8% on hosts in the same genus, and 3.6% was found on hosts in multiple genera. The Astigmata were less specific, but still showed fairly strong relationships between mite and host: 55.0% specific to a single host species, 33.3% present on host species in the same genus, and 11.7% found on hosts in multiple genera.

Each species and their host specificity levels are detailed in Table 15 for Astigmata,

Table 16 for Heterostigmatina and Table 17 for Mesostigmata at the end of the chapter.

Attachment Site Specificity

In addition to host specificity, many mites showed a preference for one or a few attachment sites on their hosts. Attachment sites were categorized into 8 distinct locations:

1. Thorax/petiole- dorsal or lateral portions of thorax and all regions of the petiole

2. Antennal- antennal groove, antennal scape, antennal flagella

3. Head- any surface of the head region not included in the antennal category plus

the neck

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4. Between legs/coxae- thorax regions surrounding the bases of the legs, ventrally

between the coxae and on the coxae, but not on any other segments of the legs

5. Legs- all segments of the legs except for the coxae

6. Gaster- regions of the abdomen posterior to the petiole

7. Wing- on the wing (only applicable for alates)

8. Unknown- any unclear or undocumented phoretic site due to lack of observation

or documentation

Number of specimens (slides) documented for each species at each location are shown in Table 16 to Table 18 located at the end of the chapter.

Prevalence of Phoretic Mite Infestations

Mite infestation prevalence was calculated for each colony. The prevalence was subcategorized as “total prevalence”, “worker prevalence”, “female prevalence”, and

“male prevalence”. Dealate prevalence was not calculated separately because only a few species had collectable dealates. Mean mite prevalences for workers, female alates, and male alates were 0.27 (SD = 0.30, N = 7142), 0.46 (SD = 0.37, N = 541, and 0.19 (SD =

0.29, N = 656) respectively. Total mite prevalence (workers, male and female alates, and dealates) was 0.27 (SD = 0.30, N = 8450).

Mites on Alates: Prevalence and Sex Preference

Mean mite prevalences of male and female alates for each colony were compared using an independent-samples t-test. For colonies in which only one sex or no alates at all were collected, the cells were left blank, zeros were only used for cases in which no mites were found on the alates, but alates were present. Female alates were collected from 34

53 colonies and males from 41. The t-test for equality of the means between male and female alates in these colonies was significant (p = 0.001, t(61) = 3.41) for females having a higher mean prevalence of mites (M = 0.46 SD = 0.37) than males (M = 0.19,

SD = 0.29). A total of 24 colonies had both alate sexes present and in 15 colonies, prevalence was higher for females. Only 4 colonies had higher prevalence for males; the remaining 5 colonies were equal with either 0% or 100% for both sexes.

Ecological Factors Effecting Phoretic Mite Diversity

Test for Adequate Sampling: Total Ants

Relationships between total ants (M = 48.29, SD = 36.39) inspected and the response variables were tested using simple linear regression to determine if sampling methods were adequate and/or if they greatly influenced the observed results. None of the response variables showed significant correlations with total ants inspected. In fact,

Astigmata showed no significance at all (p-value = 1.000) for number of ants inspected.

Results are summarized in Table 1.

Response Standardized Mean Std. t-stat p-value Explained F(1,173) to Total ants β (M) Deviation variance (SD) R2 # Phoretic 0.045 2.13 2.78 0.599 0.550 0.002 0.359 # Meso 0.129 0.35 0.71 1.715 0.088 0.017 2.941 # Hetero 0.025 0.98 1.38 0.329 0.743 <0.001 0.108 # Astig 0.000 0.79 1.35 0.000 1.000 <0.001 0.000 Prevalence -0.097 0.27 29.66 -1.281 0.202 0.009 1.640 Table 1. Results of Simple linear regression between Total ants inspected and response variables.

Habitat

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The effect of host habitat (woods vs. open) on # of phoretic mite species, # of

Mesostigmata, # of Heterostigmatina, # of Astigmata and prevalence was tested using independent-sample t-tests for equal means. Results for the t-tests indicated that ant colonies in the woods had a higher mean # of phoretic mite species per colony (Mw =

2.55, SD w = 3.20) than those in the open habitats (Mo = 1.50, SDo = 1.86), t(170.1) =

2.75, p = 0.007). Subsequent tests for # of phoretic Mesostigmata, # of phoretic

Heterostigmatina, and # of phoretic Astigmata were significant for only Heterostigmatina

(Mw = 1.34, SDw = 1.58) in which mite prevalence was significantly higher in the woods habitat (Mo = 0.44, SDo = 0.75, t(159.2) = 5.04, p<0.001). A summary of these results is shown in Table 2. Removal of the predominately forest dwelling ant Lasius umbratus (38 phoretic mite species, and generally highest number of mites per colony) altered p-value for # of phoretic mites from 0.007 to 0.144, and that of prevalence from 0.006 to 0.049.

Significance of the other variables did not change after removal of L. umbratus.

RESPONSE HABITAT N Mean Std. df t-stat p-value (M) Deviation (SD) # Phoretic Woods 105 2.55 3.20 170.1 2.75 0.007 Open 70 1.50 1.86 # Meso Woods 105 0.34 0.65 173 -0.26 0.795 Open 70 0.37 0.80 # Hetero Woods 105 1.34 1.58 159.2 5.04 <0.001 Open 70 0.44 0.75 # Astig Woods 105 0.87 1.56 173 0.87 0.388 Open 70 0.69 0.96 Prevalence Woods 105 0.32 0.30 173 2.79 0.006 Open 70 0.19 0.28 Table 2. Independent t-test for phoretic mites in colonies in the woods and open habitats. Mesostigmata, Heterostigmatina, and Astigmata have been abbreviated to “Meso”, “Hetero”, and “Astig” respectively.

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Nest substrate

Similar results were seen for t-tests with the variable nest substrate (wood vs. soil). # of Heterostigmatina (Mw = 1.29, SD w = 1.56 , t(165.9) = 3.48, p = 0.001) and prevalence (Mw = 0.31, SD w = 0.29, t(173) = 2.17, p = 0.032) had significantly higher means for colonies in wood than soil (Heterostigmatina: Ms = 0.61, SD s = 1.03, prevalence: Ms = 0.2 1, SD s = 0.29). # of Mesostigmata showed the reverse relationship with higher mite richness per colony in the soil, though results were not significant.

Results for Mesostigmata reached borderline significance (t(128.6) = -0.26, p = 0.051) after removal of L. umbratus. Results are summarized in Table 3.

RESPONSE NEST N Mean Std. Deviation df t-stat p-value SUBSTRATE (M) (SD) # Phoretic Wood 96 2.40 3.13 173 1.38 0.167 Soil 79 1.81 2.28 # Meso Wood 96 0.28 0.56 128.6 -0.26 0.151 Soil 79 0.44 0.86 # Hetero Wood 96 1.29 1.56 165.9 3.48 0.001 Soil 79 0.61 1.03 # Astig Wood 96 0.82 1.57 173 0.31 0.759 Soil 79 0.76 1.04 Prevalence Wood 96 0.31 0.29 173 2.17 0.032 Soil 79 0.21 0.29 Table 3. Independent t-test for phoretic mites in colonies in the wood and soil substrates.

Temporary Social Parasitism and the Genus Lasius

Separate t-tests were used to determine the effect of social parasitism (observed in some Lasius and Aphaenogaster colonies, Formica parasitism was not tested due to low numbers of most species). Results for the t-tests for equal means for parasitism in Lasius were highly significant for all five variables being greater in the parasitic colonies (N =

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15, non parasitic: N = 24). These results are summarized in Table 4. Removal of L. umbratus reduced the significance of all p-values, and # Heterostigmatina became non significant (p = 0.338). L. umbratus, however, comprised 11 of the 15 parasitic colonies, so removal of this species would be expected to greatly reduce the strength and reliablility of the test. Simply being in the genus Lasius (a t-test done on the entire 175 colonies) resulted in high significance (p≤0.001) for all variables. Results are summarized in Table 5.

RESPONSE LASIUS N Mean (M) Std. Deviation df t-stat p-value PARASITISM (SD) # Phoretic Parasitic 15 9.13 3.44 37 7.94 <0.001 not 24 2.54 1.74 # Meso Parasitic 15 1.27 0.80 37 4.05 <0.001 not 24 0.38 0.58 # Hetero Parasitic 15 3.87 2.03 18.5 6.04 <0.001 not 24 0.92 1.02 # Astig Parasitic 15 4.00 2.10 16.9 5.74 <0.001 not 24 1.25 0.85 Prevalence Parasitic 15 0.70 0.21 35.7 3.81 0.001 not 24 0.38 0.29 Table 4. Independent t-test for phoretic mites in Lasius colonies founded parasitically or normally.

RESPONSE GENUS N Mean (M) Std. Deviation df t-stat p-value LASIUS (SD) # Phoretic Lasius 39 5.08 4.09 40.69 5.68 <0.001 not 136 1.29 1.43 # Meso Lasius 39 0.72 0.79 53.63 3.37 0.001 not 136 0.25 0.65 # Hetero Lasius 39 2.05 2.06 42.40 4.05 <0.001 not 136 0.68 0.92 # Astig Lasius 39 2.31 1.98 40.31 6.07 <0.001 not 136 0.36 0.64 Prevalence Lasius 39 0.50 0.30 54.60 5.64 <0.001 not 136 0.20 0.26 Table 5. Independent t-test for phoretic mites in colonies founded by ants in the genus Lasius and non Lasius species.

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In a test similar to the Lasius parasitism test, colonies in the genus Aphaenogaster were analyzed. Aphaenogaster has only a single parasitic species present in Ohio, A. tennesseensis. None of the Aphaenogaster tests showed any significant differences between parasitic (N=5) and nonparasitic (N=46) colonies. This lack of significance may be attributed to insufficient collection of the parasitic species (all three non parasitic species had at least 12 representative colonies). The effect of social parasitism in this group, therefore, remains unclear.

Phylogenetic Effects of Subfamily

Phylogenetic effects were tested at the subfamily level. These analyses were restricted to Formicinae and Myrmecinae colonies due to lack of adequate representative species of Dolichoderinae, Amblyoponerinae, and Ponerinae. All of the latter three subfamilies, however, had very few associated mite species and much less than either of the other two. For all response variables p-values were <0.001 except Heterostigmatina which was less significant (p = 0.021). Removal of all Lasius colonies from the analysis greatly reduced significance for all variables, leaving only number of Mesostigmata (p =

0.021) and prevalence (p = 0.051) significant or nearly significant. Results are summarized in Table 6.

RESPONSE SUBFAMILY N Mean (M) Std. Deviation df t-stat p-value (SD) # Phoretic Formicinae 68 3.63 3.75 77.49 4.87 <0.001 Myrmecinae 91 1.33 1.21 Continued Table 6. Independent t-test for phoretic mites in colonies of ants in subfamily Formicinae and Myrmecinae

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Table 6 continued RESPONSE SUBFAMILY N Mean (M) Std. Deviation df t-stat p-value (SD) # Meso Formicinae 68 0.71 0.96 81.29 4.50 <0.001 Myrmecinae 91 0.15 0.36 # Hetero Formicinae 68 1.38 1.83 94.10 2.34 0.021 Myrmecinae 91 0.81 0.95 # Astig Formicinae 68 1.54 1.82 78.21 5.16 <0.001 Myrmecinae 91 0.36 0.61 Prevalence Formicinae 68 0.43 0.35 99.74 5.18 <0.001 Myrmecinae 91 0.19 0.20

Colony Size

Colony size was tested using one-way ANOVA with post hoc Tukey’s LSD method. The tests showed no significant differences in number of species of phoretic mites per colony between small and moderate sized colonies. However, when comparing small and moderate to large sized colonies, results were strongly significant (p<0.001) for all variables except for the Heterostigmatina which were only significant when comparing small to large (p = 0.004). Results are summarized in Table 7.

RESPONSE COLONY N COLONY Mean diff. df (x,y) F –stat Post hoc SIZE (I) SIZE (J) (MI-J) F-stat Btwn groups p-value # Phoretic small 22 moderate -1.13 (2,172) 13.58 0.187 moderate 65 large -1.64 <0.001 large 88 small 2.77 <0.001 # Meso small 22 moderate -0.14 (2,172) 12.54 0.679 moderate 65 large -0.46 <0.001 large 88 small 0.46 0.001 # Hetero small 22 moderate -0.76 (2,172) 5.21 0.063 moderate 65 large -0.28 0.422 large 88 small 1.03 0.004 # Astig small 22 moderate -0.23 (2,172) 12.85 0.738 moderate 65 large -0.90 <0.001 large 88 small 1.14 <0.001 Continued Table 7. Results of one-way ANOVA (with post hoc Tukey’s LSD method) for phoretic mites in colonies of ants forming small, moderate, and large colonies.

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Table 7 continued RESPONSE COLONY N COLONY Mean diff. df (x,y) F –stat Post hoc SIZE (I) SIZE (J) (MI-J) F-stat Btwn groups p-value Prevalence small 22 moderate -0.14 (2,172) 15.31 0.099 moderate 65 large -0.18 <0.001 large 88 small 0.32 <0.001

Host Size

Host size was analyzed using simple linear regressions. Significant positive trends were indicated for the relationship between host size and # of species of phoretic mites (p

= 0.046), # of Mesostigmata (highly significant p = <0.001), # of Heterostigmatina (p =

0.031), and prevalence (highly significant p = 0.001), but not for # of Astigmata (p =

0.924). R2 values were generally quite low indicating that host size alone explains little of the variance observed within this dataset. Results are summarized in Table 8.

Response Standardized Mean Std. t-stat p-value Explained F (1, 173) to Host size β (M) Deviation variance (SD) R2 # Phoretic 0.151 2.13 0.42 2.10 0.046 0.023 4.04 # Meso 0.288 0.35 0.20 3.95 <0.001 0.083 15.61 # Hetero 0.163 0.98 0.23 2.18 0.031 0.027 4.73 # Astig -0.007 0.79 0.01 -0.10 0.924 <0.001 0.009 Prevalence 0.395 0.27 0.12 5.66 <0.001 0.156 32.038 Table 8. Results of simple linear regression showing a significantly increasing trend for all response variables except for the number of Astigmata.

Multiple Linear Regression

Stepwise multiple linear regressions were used to determine which combination of ecological and phylogenetic factors (habitat, nest substrate, parasitic, genus Lasius, subfamily, colony size, host size, and total ants) best explains the response variables. The best model for # of phoretic mites, # of Heterostigmatina, and prevalence included 60 parasitic, genus Lasius, and habitat. Mesostigmata were best explained by parasitic and subfamily, Astigmata were determined by genus Lasius and parasitic. All models included parasitic. Results are shown in Table 9.

RESPONSE MODEL t-stat p-value Explained F VARIABLE COMPONENTS variance df = (x,y) R2 # Phoretic Parasitic -7.934 <0.001 0.517 61.09 Genus Lasius -7.522 <0.001 (3, 171) Habitat -2.426 0.016 # Meso Parasitic -5.819 <0.001 0.265 31.05 Subfamily -2.925 0.004 (2, 172) # Hetero Genus Lasius -4.510 <0.001 0.327 27.71 Habitat -4.449 <0.001 (3, 171) Parasitic -4.427 <0.001 # Astig Genus Lasius -8.516 <0.001 0.494 84.03 Parasitic -6.736 <0.001 (2, 172) Prevalence Parasitic -3.271 0.001 0.452 46.922 Host size 6.350 <0.001 (3, 171) Genus Lasius -6.958 <0.001 Table 9. Stepwise multiple linear regression model results for determining the response variables.

Estimating Number of Associated Mite Species

Species accumulation curves were calculated using EstimateS for selected species of ants to determine if an asymptote was reached at which most of the associated mite species had been collected after repeated colony collections. Curves for Aphaenogaster species (Figure 2), Lasius species (Figure 3), Solenopsis molesta, and Ponera pennsylvanica (Figure 4) can be seen at the end of the chapter. Aphaenogaster species do not appear to be approaching an asymptote indicating further sampling is needed for this group. Lasius neoniger and L. alienus both appear to be reaching asymptotes around 15 total mite species. L. umbratus appears to be approaching an asymptote around 40 or 45

61 species, and L. , sampled only 4 times, appears to be following a similar trajectory as L. umbratus and is in need of further sampling. Solenopsis molesta and

Ponera pennsylvanica have clear asymptotes at 5 and 3, respectively, indicating that adequate numbers of colonies were sampled.

Non-phoretic Mites from the Nest

At least 66 non-phoretic mite species were gathered from soil, ant food, fungus, dead ants, and larvae in the ant nests. Attempts to determine the non-phoretic mites to morphospecies were limited by the fact that many of the non-phoretic mites represented immature feeding instars. For most immature mites there are no keys available, and often only the adult instars have been described. Thus, any mites listed as “undet sp.” may in fact be immature instars of adults already on the list with morphospecies numbers.

Without laboratory rearing, however, linking these stages may not be possible. 256 of the slides were Astigmatid mites (14 morphospecies determined), 138 slides were

Mesostigmata (35 morphospecies determined), 36 slides were Heterostigmatina (13 morphospecies determined), and 4 slides were other non-Heterostigmatid Prostigmata (4 morphospecies determined, all singletons). Of the 66 non-phoretic species 47 were only found in non-phoretic relationships, and 19 were also found phoretically. Table 21, Table

22, and

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at the end of this chapter show the non-phoretic mite associations.

Rearing of Astigmatid mites was highly successful for a few of the mite species attempted. Sancassania sp2 and Schwiebea sp1 were able to reproduce successfully even without access to males of their species. In many cases an immature mite was placed into the culture well and within 4 days, the second generation offspring were observed. In both of these cases, deutonymphs were collected along with the other instars (larvae, protonymphs, tritonymphs, and adult males and females), which created a link between the heteromorphic deutonymphs seen on the ants and the other stages encountered within the colony. Histiostoma females were also used for rearing; however, attempts were less successful. Deutonymphs were not observed, but other instars were collected.

Specificity and Roles of Non-phoretic Mites

Generally most of the non-phoretic mite species were specific to only one or two ant species’, as previously observed for phoretic mites. The Astigmatid deutonymphs and

Heterostigmatina were commonly found on the dead ants in their usual phoretic positions

(perhaps developing into a feeding instar or else still anticipating a ride). Feeding stages of Astigmata, such as Histiostoma and Schwiebea, and Mesostigmata, such as

Cosmolaelaps, were frequently found actively feeding on the dead ants or fluids on the ants. The Heterostigmatina (especially ) were often found walking on the old ant food and feeding on the fungal hyphae growing from it. Antennophorus species were observed on two species of Lasius eliciting trophallactic, responses from their hosts by waving their forelegs rapidly across the ants’ mouth, as previously documented

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(Franks et al., 1991; Janet, 1897). Trichocylliba sp1 (possibly comata) was collected only a couple of times from larvae in Lasius umbratus colonies. Its highly curved dorsum allowed it to hold tightly to the larvae and avoid removal by the host. One was observed walking around on a and appeared to be eliminating a white substance (similar in color to the ant larvae). This may indicate intake of larval hemolymph; however, I did not observe any dark feeding scars upon the hosts, and when not walking, any activity under the mite’s dorsum is not observable. Imparipes sp1 (Scutacaridae) was observed multiple times on larvae of Aphaenogaster species as well, but feeding scars were not observed.

Chelicerae in this species do not seem well adapted for puncturing a larva’s cuticle, and may instead be used for cleaning the larvae or for ingesting the secretions provided to the larvae by the workers.

Discussion

The Significance of Species Specificity and the Issue of Lasius

Ants are generally very protective of their nests and allow access only to members. Ant colonies can be likened to islands which have little or no interactions with each other. This lack of interaction may be the main factor influencing host specificity, which was observed in this study. Ant-associated mites have little to no access to any modes of transportation aside from their ant hosts, and thus they have little opportunity to move between colonies beyond vertical transmission through dispersal on alates.

Interestingly, colonies formed by the genus Lasius have a greater number of mite species per colony than non-Lasius species. Furthermore, within Lasius, greater numbers are observed on the parasitic ant species. The variables parasitism and genus Lasius

64 appeared in almost all of the models predicting mite species richness and prevalence. The data suggest that parasitic Lasius species acquire mites from their hosts through horizontal transmission. This is further supported by a collection I made of a colony with a documented parasitism history (KAU-09-0725-1). The colony was first observed by G.

Coovert when it was a Lasius umbratus colony and then as a mixed colony of L. umbratus and L. speculiventris (a hyperparasite). At the point of my collection, there were only L. speculiventris workers present. All 10 mite species I collected from the L. speculiventris workers were also collected from other L. umbratus colonies; none were unique to L. speculiventris.

In fact, a close examination of the full list of mites shows that 26 species of mites are shared among the Lasius ants, 8 are shared among Aphaenogaster ants, and 3 are shared among Formica ants. It is worthwhile considering this in context. Only 47 of the

151 species of mites observed during this study were found on more than one host species (38 specific to genus, and 9 collected from multiple genera). All but one (a single specimen on a Camponotus sp.) of the 38 mite species observed on multiple ants in the same genus may be accounted for by horizontal transmission by ants belonging to the 3 parasitic genera. As a result, the percentage of non-specific mite species (unaccounted for by those with parasitic hosts) is just 5.9%. It appears that in situations such as social parasitism, where the original host becomes unavailable, mites will climb aboard the parasitic species in its absence.

In sharp contrast, are situations where two different species of ants live close together without social parasitism associations (including plesiobiosis, cleptobiosis,

65 lestobiosis, parabiosis, and xenobiosis; see Chapter 1). In these cases members of each ant species were given “a” and “b” designations at the end of their field numbers.

Cohabiting colonies collected include: Tetramorium caespitum x Camponotus chromaiodes, Lasius neoniger x Solenopsis molesta (3 instances), L. alienus x Ponera pennsylvanica, Formica integra x S. molesta, Paratrechina faisonenesis x Ponera pennsylvanica, Aphaenogaster fulva x Ponera pennsylvanica, Formica exsectoides and S. molesta (2 instances), F. subsericea x S. molesta x T. caespitum, T. caespitum x S. molesta, F. nitidiventris x S. molesta, Aphaenogaster sp. x Ponera pennsylvanica, x

Myrmecina americana, M. americana x Ponera pennsylvanica, Paratrechina parvula x

S. molesta, C. subbarbatus x Pyramica laevinasis, L. umbratus x M. americana. In none of these instances of cohabitation were any mite species observed on both of the host species available to them. This suggests that when given the option of more than one host species, mites preferentially remain on their original hosts, but when given no other alternative, as in social parasitism (after the host species is dead), mites will board another ant species.

Host Sex Specificity

The result of the t-test for alate sex preference in mites was highly significant for higher prevalence on female alates. Male ants do not live long after mating and generally do not return to a nest. Strong selective pressure exists for the mites to differentially board female alates rather than males. However, necrophagy is common in ants, thus it would still be possible for mites on dead males to make it back into a nest (Hölldobler &

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Wilson, 1990). Reliance on necrophagy is a risky option for mites boarding males, but if no female alates are produced by the colony, this may be the only option for dispersal.

Other Ecological Factors Affecting Mite Species Richness and Abundance

Host size is an important factor determining prevalence and # of phoretic mites, especially for mites in the Mesostigmata. It is logical that a larger host would have more space available on which more mites could cling and subsequently would allow more mite species. There also was an apparent trend between mesostigmatid mites and host size. Small ant species do not have correspondingly smaller mites, and probably are incapable of even transporting a mesostigmatid mite without it becoming a hindrance

(and easily detected).

Habitat (woods or open) significance ranges from strongly correlated (as in

Heterostigmatina for woods habitats) to not clearly correlated (but preferring soil as in the Mesostigmata). Habitat was highly correlated (Pearson Correlation = 0.806, p<0.001) with nest substrate (wood or soil) which was anticipated as most of the logs or other wood substrates were found only in the woods. It may be that woods, open, wood and soil are too broad of categories, however, and that mites are capable of sensing the environment at minute levels. It may be necessary to ask more specific questions in future studies such as: what trees are present in the woods?, is the open environment urban or rural?, is the wood moist? and what kind of tree does it come from? Most of the

“open” environments were suburban or urban suggesting a confounding factor, disturbance, within this variable. Disturbance may be a more influential factor, leading to the lower richness of mite species observed in open environments.

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The Heterostigmatina showed a highly significant increase in richness in the woods and wood substrates. There were also many more associated species in Lasius nests than non Lasius nests. A possible explanation for this may lie in the symbiotic fungus relationship occurring within the ant genus. Ascomycete fungal associates are cultured by Lasius ants for nest architecture. Ants provision the fungi with honeydew and shredded wood creating carton-like building material (Schlick-Steiner et al., 2008).

Fungal associates show specificity within the Lasius genus, and this specificity may be mirrored by the Heterostigmatina, which may feed upon the fungi or competing fungal species. The Heterostigmatina may have more complex roles within the colony than just as commensals.

As colony size increased, prevalence and # of mite species also increased in all categories. A larger colony holds a greater resource patch for any organisms living within, be it brood, food stores, dead ants, , or other mite species. Mesostigmata generally have large body sizes, and many are predatory. A small resource patch would be less capable of providing for Mesostigmata. Aside from the large mites, one would expect mites to be less often encountered in small colonies. The ant colonies may be governed by rules similar to those of the Theory of Island Biogeography (MacArthur &

Wilson, 1967). For example, a large colony may provide a larger habitat area, more opportunities for variation within the habitat, and less probability of extinction due to random events, while small colonies harbor fewer resources, and may be more prone to chance extinctions.

The Importance of Rearing

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The fact that attempts in rearing Astigmata were generally fruitful with limited equipment and time investment, marks a big step toward better understanding of taxonomy of ant associates. Only a few studies have described more than one instar of the mite species (Ebermann, 1981; Ebermann & Rack, 1982; Wurst, 2001). Further studies should attempt rearing in order to link the heteromorphic deutonymphs of the Astigmata to their feeding instars. Another option for such linking might rely on DNA barcoding.

Caveats

Uneven sampling was an issue anticipated at the start of this project. Ant species make nests of varying sizes and many of the average sizes of colonies have not been documented. A protocol had to be developed for sampling ant colonies of various sizes.

Usually by the time 30 ants had been inspected in the colony, I was beginning to see mostly the same mites, but in cases of more diverse mite assemblages, more ants were sometimes checked. Small ant colonies gave little room for adjustment, and usually all of the ants in the colony were examined. Despite the issues I had with forming a protocol, based on the results of the total ants regression it appears that number of ants inspected explained little of the variance observed in the mite numbers, thus my sampling method within colonies was adequate with at least 20 ants per colony being checked for inclusion in the analysis (10 for extremely small ant colonies).

However, for many ant species the number of colonies collected was not adequate to estimate the overall number of associated mite species. For example, for the 13 ant species without any mites I did not have more than 3 colonies collected. Further collections of those ant species would likely yield mites. Species accumulation curves of

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Lasius and Aphaenogaster species (although the most thoroughly collected) still indicate more species awaiting discovery (Figure 2 and Figure 3). Others, such as Solenopsis molesta, and Ponera pennsylvanica show clear asymptotes indicating thorough sampling

(Figure 4).

Summary of Conclusions

Generally most mites appear to be in a commensal relationship with the ants by feeding on dead ants, bacterial and fungal resources within the ant nests, while a few were found on brood or had clearly parasitic relationships with their ant hosts (such as

Antennophorus). Mite species richness and prevalence were significantly greater in colonies when 1) the ant subfamily was Formicinae, 2) the ant genus was Lasius, 3) the ants were large, 4) the ant species established its nest parasitically, 5) the colony was populous, 6) the colony was in the woods, and 7) the nest substrate was wood.

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GENUS FAMILY MORPHOSPECIES HOST ANT SPECIES # SLIDES # COLONIES Acaridae Armacarus sp1 18 2 Lasius umbratus 3 3 Acaridae Cosmoglyphus sp1 Solenopsis molesta 11 7 Acaridae Cosmoglyphus sp2 Lasius claviger 3 2 Lasius neoniger 9 5 Acaridae Forcellinia sp1 Lasius neoniger 19 6 Acaridae Forcellinia sp2 Lasius neoniger 3 1 Acaridae Forcellinia sp3 cerasi 2 1 Crematogaster lineolata 1 1 Acaridae Forcellinia sp4 Camponotus nearcticus 33 2 Tapinoma sessile 1 1 Acaridae Forcellinia sp5 Camponotus chromaiodes 1 1 Acaridae Forcellinia sp6 Crematogaster lineolata 2 1 Acaridae Forcellinia sp7 Tetramorium caespitum 3 1 Acaridae Forcellinia sp8 Formica nitidiventris 6 1 Acaridae Forcellinia sp9 Myrmica americana 10 1 Acaridae Forcellinia sp10 Lasius claviger 8 1 Lasius interjectus 4 1 Lasius speculiventris 1 1 Lasius umbratus 23 7 Acaridae Forcellinia sp11 Aphaenogaster picea 1 1 complex Acaridae Froriepia sp1 Lasius claviger 5 3 Lasius interjectus 11 1 Acaridae Froriepia sp2 Lasius umbratus 1 1 Acaridae Froriepia sp3 Solenopsis molesta 2 2 Acaridae Sancassania sp1 Formica exsectoides 1 1 Acaridae Sancassania sp3 Lasius nearcticus 1 1 Acaridae Schwiebea sp1 Lasius interjectus 2 1 Lasius umbratus 14 6 Acaridae Schwiebea sp2 Lasius claviger 12 1 Lasius interjectus 1 1 Continued Table 10. Astigmata Mites and Hosts. Mite species are shown with their associated host species, number of representative slides, and number of colonies in which the mite species were found for each host.

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Table 10 continued GENUS FAMILY MORPHOSPECIES HOST ANT SPECIES # SLIDES # COLONIES Acaridae Schwiebea sp3 Lasius claviger 3 2 Lasius interjectus 1 1 Lasius umbratus 3 1 Acaridae Schwiebea sp4 Aphaenogaster picea complex 4 1 Acaridae Schwiebea sp5 Lasius speculiventris 1 1 Lasius umbratus 1 1 Acaridae Schwiebea sp6 1 1 Acaridae Schwiebea sp7 Aphaenogaster picea complex 2 1 Myrmica americana 1 1 Acaridae Schwiebea sp8 Aphaenogaster fulva 6 2 Camponotus nearcticus 2 1 Crematogaster cerasi 5 2 Crematogaster pilosa 2 1 Acaridae Schwiebea sp9 Tetramorium caespitum 1 1 Acaridae Schwiebea sp10 Formica exsectoides 1 1 Acaridae Schwiebea sp11 Lasius umbratus 1 1 Acaridae Schwiebea sp12 Prenolepis imparis 3 1 Astigmata undet undet gen sp3 Lasius umbratus 1 1 Astigmata undet undet gen sp4 Lasius umbratus 1 1 Histiostomatidae Histiostoma sp1 Camponotus chromaiodes 1 1 Camponotus pennsylvanicus 2 1 Histiostomatidae Histiostoma sp2 Lasius claviger 3 1 Lasius umbratus 15 3 Histiostomatidae Histiostoma sp3 Lasius claviger 1 1 Histiostomatidae Histiostoma sp4 Aphaenogaster fulva 1 1 Aphaenogaster picea complex 1 1 Histiostomatidae Histiostoma sp5 Aphaenogaster tennesseensis 3 1 Camponotus chromaiodes 9 1 Myrmecina americana 1 1 Histiostomatidae Histiostoma sp6 Aphaenogaster fulva 27 3 Aphaenogaster rudis complex 1 1 Histiostomatidae Histiostoma sp7 Aphaenogaster fulva 2 1 Aphaenogaster picea complex 1 1 Histiostomatidae Histiostoma sp8 Aphaenogaster fulva 3 1 Histiostomatidae Histiostoma sp9 Aphaenogaster fulva 1 1 Histiostomatidae Histiostoma sp10 Tetramorium caespitum 2 1 Continued 72

Table 10 continued GENUS FAMILY MORPHOSPECIES HOST ANT SPECIES # SLIDES # COLONIES Histiostomatidae Histiostoma sp11 Lasius claviger 3 1 Histiostomatidae Histiostoma sp12 Lasius alienus 78 13 Lasius claviger 26 2 Lasius umbratus 11 5 Histiostomatidae Histiostoma sp13 Lasius alienus 1 1 Lasius claviger 7 1 Lasius interjectus 1 1 Lasius neoniger 2 1 Lasius speculiventris 5 1 Lasius umbratus 43 11 Myrmecina americana 2 1 Histiostomatidae Histiostoma sp14 Camponotus pennsylvanicus 2 1 Lasius alienus 2 1 Lasius neoniger 1 1 Lasius umbratus 1 1 Tetramorium caespitum 12 2 Histiostomatidae Histiostoma sp15 Lasius alienus 1 1 Lasius interjectus 20 1 Lasius neoniger 3 2 Lasius umbratus 18 6 Histiostomatidae Histiostoma sp16 Crematogaster cerasi 2 1 Histiostomatidae Histiostoma sp17 Lasius speculiventris 2 1 Lasius umbratus 17 6 Histiostomatidae Histiostoma sp18 Formica exsectoides 25 4 Formica nitidiventris 2 1 Histiostomatidae Histiostoma sp19 Formica exsectoides 2 1 Formica nitidiventris 7 1 18 4 Histiostomatidae Histiostoma sp20 Formica exsectoides 1 1 Formica subsericea 7 2 Histiostomatidae Histiostoma sp21 Lasius neoniger 2 2 Pyramica ohioensis 1 1 Histiostomatidae Histiostoma sp22 Camponotus pennsylvanicus 2 1 Histiostomatidae Histiostoma sp23 Lasius neoniger 8 3 Histiostomatidae Histiostoma sp24 Camponotus subbarbatus 1 1 Continued

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Table 10 continued GENUS FAMILY MORPHOSPECIES HOST ANT SPECIES # SLIDES # COLONIES Histiostomatidae Histiostoma sp25 Crematogaster cerasi 1 1 Histiostomatidae Histiostoma sp26 Aphaenogaster tennesseensis 2 1 Histiostomatidae Histiostoma sp27 Prenolepis imparis 2 2 Totals: 60 mite species 667

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GENUS FAMILY MORPHOSPECIES HOST ANT SPECIES # SLIDES # COLONIES Microdispidae Caesarodispus sp2 Camponotus chromaiodes 1 1 Microdispidae Caesarodispus sp11 Lasius speculiventris 1 1 Lasius umbratus 4 1 Lasius nearcticus 6 2 Microdispidae Caesarodispus sp15 Formica subsericea 1 1 Microdispidae Caesarodispus sp16 Prenolepis imparis 8 2 Microdispidae Unguidispus sp10 Lasius speculiventris 1 1 Lasius umbratus 11 4 Pygmephoridae Bakerdania sp3 Camponotus chromaiodes 1 1 Pygmephoridae Bakerdania sp5 Formica subsericea 5 2 Pygmephoridae Bakerdania sp7 Lasius alienus 1 1 Pygmephoridae Bakerdania sp9 Lasius claviger 3 2 Pygmephoridae Bakerdania sp12 Lasius claviger 1 1 Lasius umbratus 3 2 Pygmephoridae Bakerdania sp13 Lasius umbratus 3 2 Pygmephoridae Bakerdania sp14 Formica subsericea 1 1 Solenopsis molesta 2 2 Pygmephoridae Petalomium sp6 Lasius alienus 1 1 Pygmephoridae Petalomium sp8 Lasius claviger 5 2 Lasius interjectus 8 1 Lasius neoniger 1 1 Lasius umbratus 4 2 Scutacaridae Diversipes sp1 Lasius alienus 3 1 Scutacaridae Imparipes sp1 Aphaenogaster fulva 16 7 Aphaenogaster picea complex 21 13 Aphaenogaster rudis complex 17 8 Aphaenogaster tennesseensis 9 2 Scutacaridae Imparipes sp2 Aphaenogaster picea complex 6 4 Aphaenogaster rudis complex 2 2 Continued Table 11. Heterostigmatina Mites and Hosts. Mite species are shown with their associated host species, number of representative slides, and number of colonies in which the mite species were found for each host.

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Table 11 continued GENUS FAMILY MORPHOSPECIES HOST ANT SPECIES # SLIDES # COLONIES Scutacaridae Imparipes sp4 Lasius alienus 9 3 Lasius claviger 21 3 Lasius interjectus 3 1 Lasius umbratus 25 5 Scutacaridae Imparipes sp5 Lasius claviger 2 1 Lasius umbratus 9 5 Scutacaridae Imparipes sp6 Lasius umbratus 3 3 Scutacaridae Imparipes sp7 Solenopsis molesta 3 3 Scutacaridae Imparipes sp8 Solenopsis molesta 7 6 Scutacaridae Imparipes sp9 Ponera pennsylvanica 9 4 Scutacaridae Imparipes sp10 Ponera pennsylvanica 3 3 Scutacaridae Imparipes sp11 Tetramorium caespitum 16 4 Scutacaridae Imparipes sp12 Lasius neoniger 1 1 Scutacaridae Imparipes sp13 Lasius neoniger 1 1 Scutacaridae Imparipes sp14 Formica subsericea 1 1 Scutacaridae Pygmodispus sp1 Lasius alienus 2 1 Scutacaridae Scutacarus sp1 Aphaenogaster picea complex 4 3 Aphaenogaster rudis complex 2 1 Scutacaridae Scutacarus sp2 Lasius alienus 12 7 Lasius neoniger 1 1 Scutacaridae Scutacarus sp3 Lasius alienus 5 3 Scutacaridae Scutacarus sp4 Lasius speculiventris 3 1 Lasius umbratus 27 9 Scutacaridae Scutacarus sp5 Lasius umbratus 5 3 Scutacaridae Scutacarus sp6 Lasius speculiventris 4 1 Lasius umbratus 2 2 Scutacaridae Scutacarus sp7 Lasius umbratus 2 2 Scutacaridae Scutacarus sp8 Lasius speculiventris 1 1 Lasius umbratus 1 1 Scutacaridae Scutacarus sp9 Lasius umbratus 3 2 Scutacaridae Scutacarus sp10 Aphaenogaster rudis complex 6 1 Scutacaridae Scutacarus sp11 Aphaenogaster fulva 12 3 Aphaenogaster picea complex 7 2 Aphaenogaster rudis complex 11 2 Aphaenogaster tennesseensis 3 2 Continued

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Table 11 continued GENUS FAMILY MORPHOSPECIES HOST ANT SPECIES # SLIDES # COLONIES Scutacaridae Scutacarus sp12 Aphaenogaster fulva 15 9 Aphaenogaster picea complex 3 2 Aphaenogaster rudis complex 16 7 Aphaenogaster tennesseensis 3 2 Ponera pennsylvanica 1 1 Scutacaridae Scutacarus sp13 Camponotus chromaiodes 10 3 Scutacaridae Scutacarus sp14 Formica subsericea 2 2 Scutacaridae Scutacarus sp15 Prenolepis imparis 2 2 Scutacaridae Scutacarus sp17 Formica exsectoides 2 2 Scutacaridae Scutacarus sp18 Formica nitidiventris 1 1 Scutacaridae Scutacarus sp19 Tapinoma sessile 1 1 Scutacaridae Scutacarus sp20 Lasius claviger 2 1 Lasius umbratus 2 2 Scutacaridae Scutacarus sp21 Lasius umbratus 5 2 Scutacaridae Scutacarus sp22 Lasius neoniger 1 1 Scutacaridae Scutacarus sp23 Lasius neoniger 2 1 Scutacaridae Scutacarus sp24 Lasius interjectus 1 1 Scutacaridae Scutacarus sp25 Lasius interjectus 2 1 Scutacaridae Scutacarus sp26 Lasius interjectus 5 1 Scutacaridae Scutacarus sp27 Lasius neoniger 3 2 Scutacaridae Scutacarus sp28 Lasius umbratus 1 1 Totals: 56 species 452

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GENUS FAMILY MORPHOSPECIES HOST # SLIDES # COLONIES Laelapidae Cosmolaelaps sp3 Lasius alienus 4 1 Laelapidae Cosmolaelaps sp4 Camponotus chromaiodes 1 1 Laelapidae Cosmolaelaps sp8 Lasius umbratus 1 1 Laelapidae Cosmolaelaps sp9 Formica exsectoides 3 1 Laelapidae Cosmolaelaps sp10 Lasius umbratus 2 2 Laelapidae Cosmolaelaps sp13 Aphaenogaster rudis complex 1 1 Laelapidae Cosmolaelaps sp14 Aphaenogaster rudis complex 2 1 Laelapidae Cosmolaelaps sp15 Lasius neoniger 1 1 Laelapidae Cosmolaelaps sp19 Formica exsectoides 4 1 Laelapidae Cosmolaelaps sp21 Lasius claviger 1 1 Laelapidae Gaeolaelaps sp2 Formica exsectoides 11 6 Laelapidae Gaeolaelaps sp3 Aphaenogaster fulva 1 1 Laelapidae Gaeolaelaps sp4 Lasius claviger 1 1 Lasius umbratus 1 1 Laelapidae Gaeolaelaps sp6 Formica integra 6 1 Laelapidae Gaeolaelaps sp7 Formica nitidiventris 12 2 Laelapidae Gaeolaelaps sp8 Formica subsericea 3 2 Laelapidae Laelaspis cf. dubitatus Aphaenogaster picea complex 2 1 Aphaenogaster rudis complex 4 3 Laelapidae Laelaspis cf. vitzthumi Formica exsectoides 2 1 Laelapidae Laelaspis nr. picketti Tetramorium caespitum 4 1 Oplitidae Oplitis aktius Camponotus nearcticus 3 1 Oplitidae Oplitis alienorum Lasius alienus 16 4 Oplitidae Oplitis exsectoidesorum Formica exsectoides 2 2 Oplitidae Oplitis sarcinulus Tetramorium caespitum 7 4 Oplitidae Oplitis sp1 Lasius neoniger 10 6 Oplitidae Oplitis sp2 Camponotus chromaiodes 1 1 Continued Table 12. Mesostigmata Mites and Hosts. Mite species are shown with their associated host species, number of representative slides, and number of colonies in which the mite species were found for each host.

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Table 12 continued GENUS FAMILY MORPHOSPECIES HOST # SLIDES # COLONIES Oplitidae Oplitis sp3 Camponotus chromaiodes 1 1 Oplitidae Oplitis sp8 Formica nitidiventris 2 1 Oplitidae Oplitis sp9 Formica integra 2 1 Oplitidae Oplitis sp10 Formica subsericea 1 1 Oplitidae Oplitis (Uroplitana) sp11 Lasius speculiventris 1 1 Lasius umbratus 1 1 Rhodacaridae undet gen sp1 Formica exsectoides 1 1 Uropodidae Uropolyaspis sp1 Lasius umbratus 6 6 Uropodidae Uropolyaspis sp2 Lasius claviger 1 1 Antennophoridae Antennophorus sp1 Lasius umbratus 12 3 Antennophoridae Antennophorus sp2 Lasius claviger 1 1 Totals 35 species 135

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TOTAL # PHORETIC GENUS SPECIES # COLONIES ANTS SPP Amblyopone pallipes 3 19 0 Aphaenogaster fulva 15 604 10 Aphaenogaster picea complex 22 944 11 Aphaenogaster rudis complex 15 497 10 Aphaenogaster tennessensis 6 232 5 Brachymyrmex depilis 3 69 0 Camponotus chromaiodes 14 196 9 Camponotus nearcticus 3 50 3 Camponotus pennsylvanicus 4 8 3 Camponotus subbarbatus 3 74 1 Crematogaster cerasi 4 225 4 Crematogaster lineolata 2 61 2 Crematogaster pilosa 2 20 1 Formica exectoides 11 360 12 Formica integra 1 89 2 Formica nitidiventris 3 60 6 Formica rubicunda 1 11 0 Formica subsericea 8 192 9 Lasius alienus 19 708 14 Lasius claviger 4 182 21 Lasius interjectus 1 17 12 Lasius nearcticus 4 160 2 Lasius neoniger 15 468 17 Lasius speculiventris 1 56 10 Lasius umbratus 18 611 38 minimum 3 107 0 Myrmecina americana 3 17 2 Myrmica americana 2 23 2 Myrmica latifrons 1 2 0 Myrmica punctiventris 2 24 0 Paratrechina faisonensis 3 94 0 Paratrechina parvula 1 70 0 Pheidole bicarinata 2 32 0 Ponera pennsylvanica 20 126 3 Prenolepis imparis 3 167 4 Proceratium pergandei 1 1 0 Pyramica laevinasis 1 1 0 Pyramica ohioensis 2 53 1 Continued Table 13. Number of Phoretic Mites. Each ant species is shown with number of colonies and individuals inspected and the resulting number of phoretic mites encountered. 80

Table 13 continued TOTAL # PHORETIC GENUS SPECIES # COLONIES ANTS SPP Pyramica reflexa 1 13 0 Pyramica sp. 1 1 0 Solenopsis molesta 16 838 5 Tapinoma sessile 10 409 2 Tetramorium caespitum 19 1202 7

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ANT SPECIES MITE FAMILY MITE SPECIES # SLIDES # COLONIES Aphaenogaster fulva Acaridae Schwiebea sp8 6 2 Histiostomatidae Histiostoma sp4 1 1 Histiostomatidae Histiostoma sp6 27 3 Histiostomatidae Histiostoma sp7 2 1 Histiostomatidae Histiostoma sp8 3 1 Histiostomatidae Histiostoma sp9 1 1 Scutacaridae Imparipes sp1 16 7 Scutacaridae Scutacarus sp11 12 3 Scutacaridae Scutacarus sp12 15 9 Laelapidae Gaeolaelaps sp3 1 1 Acaridae Forcellinia sp11 1 1 Aphaenogaster picea Acaridae Schwiebea sp4 4 1 complex Acaridae Schwiebea sp7 2 1 Histiostomatidae Histiostoma sp4 1 1 Histiostomatidae Histiostoma sp7 1 1 Scutacaridae Imparipes sp1 21 13 Scutacaridae Imparipes sp2 6 4 Scutacaridae Scutacarus sp1 4 3 Scutacaridae Scutacarus sp11 7 2 Scutacaridae Scutacarus sp12 3 2 Laelapidae Laelaspis cf. dubitatus 2 1 Histiostomatidae Histiostoma sp6 1 1 Aphaenogaster rudis Scutacaridae Imparipes sp1 17 8 complex Scutacaridae Imparipes sp2 2 2 Scutacaridae Scutacarus sp1 2 1 Scutacaridae Scutacarus sp10 6 1 Scutacaridae Scutacarus sp11 11 2 Scutacaridae Scutacarus sp12 16 7 Laelapidae Laelaspis cf. dubitatus 4 3 Laelapidae Cosmolaelaps sp13 1 1 Laelapidae Cosmolaelaps sp14 2 1 Continued Table 14. Ants and their Associated Mite Species.

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Table 14 continued ANT SPECIES MITE FAMILY MITE SPECIES # SLIDES # COLONIES Aphaenogaster Histiostomatidae Histiostoma sp5 3 1 tennesseensis Histiostomatidae Histiostoma sp26 2 1 Scutacaridae Imparipes sp1 9 2 Scutacaridae Scutacarus sp11 3 2 Scutacaridae Scutacarus sp12 3 2 Camponotus Acaridae Forcellinia sp5 1 1 chromaiodes Histiostomatidae Histiostoma sp1 1 1 Histiostomatidae Histiostoma sp5 9 1 Microdispidae Caesarodispus sp2 1 1 Pygmephoridae Bakerdania sp3 1 1 Scutacaridae Scutacarus sp13 10 3 Laelapidae Cosmolaelaps sp4 1 1 Oplitidae Oplitis sp2 1 1 Oplitidae Oplitis sp3 1 1 Camponotus nearcticus Acaridae Forcellinia sp4 33 2 Acaridae Schwiebea sp8 2 1 Oplitidae Oplitis aktius 3 1 Camponotus Histiostomatidae Histiostoma sp1 2 1 pennsylvanicus Histiostomatidae Histiostoma sp14 2 1 Histiostomatidae Histiostoma sp22 2 1 Camponotus subbarbatus Histiostomatidae Histiostoma sp24 1 1 Crematogaster cerasi Acaridae Forcellinia sp3 2 1 Acaridae Schwiebea sp8 5 2 Histiostomatidae Histiostoma sp16 2 1 Histiostomatidae Histiostoma sp25 1 1 Crematogaster lineolata Acaridae Forcellinia sp3 1 1 Acaridae Forcellinia sp6 2 1 Crematogaster pilosa Acaridae Schwiebea sp8 2 1 Formica exsectoides Acaridae Schwiebea sp10 1 1 Acaridae Sancassania sp1 1 1 Histiostomatidae Histiostoma sp18 25 4 Histiostomatidae Histiostoma sp19 2 1

Histiostomatidae Histiostoma sp20 1 1

Scutacaridae Scutacarus sp17 2 2

Laelapidae Cosmolaelaps sp9 3 1 Continued

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Table 14 continued ANT SPECIES MITE FAMILY MITE SPECIES # SLIDES # COLONIES Formica exsectoides Laelapidae Cosmolaelaps sp19 4 1 (continued) Laelapidae Gaeolaelaps sp2 11 6 Laelapidae Laelaspis cf. vitzthumi 2 1 Oplitidae Oplitis exsectoidesorum 2 2 Rhodacaridae undet gen sp1 1 1 Formica integra Laelapidae Gaeolaelaps sp6 6 1 Oplitidae Oplitis sp9 2 1 Formica nitidiventris Acaridae Forcellinia sp8 6 1 Histiostomatidae Histiostoma sp18 2 1 Histiostomatidae Histiostoma sp19 7 1 Scutacaridae Scutacarus sp18 1 1 Laelapidae Gaeolaelaps sp7 12 2 Oplitidae Oplitis sp8 2 1 Formica subsericea Histiostomatidae Histiostoma sp19 18 4

Histiostomatidae Histiostoma sp20 7 2 Microdispidae Caesarodispus sp15 1 1 Pygmephoridae Bakerdania sp5 5 2 Pygmephoridae Bakerdania sp14 1 1 Scutacaridae Imparipes sp14 1 1 Scutacaridae Scutacarus sp14 2 2 Laelapidae Gaeolaelaps sp8 3 2 Oplitidae Oplitis sp10 1 1 Lasius alienus Acaridae Schwiebea sp6 1 1

Histiostomatidae Histiostoma sp12 78 13

Histiostomatidae Histiostoma sp13 1 1 Histiostomatidae Histiostoma sp14 2 1 Histiostomatidae Histiostoma sp15 1 1 Pygmephoridae Bakerdania sp7 1 1 Pygmephoridae Petalomium sp6 1 1 Scutacaridae Diversipes sp1 3 1 Scutacaridae Imparipes sp4 9 3 Scutacaridae Pygmodispus sp1 2 1 Scutacaridae Scutacarus sp2 12 7 Scutacaridae Scutacarus sp3 5 3 Continued

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Table 14 continued ANT SPECIES MITE FAMILY MITE SPECIES # SLIDES # COLONIES Lasius alienus Laelapidae Cosmolaelaps sp3 4 1 (continued) Oplitidae Oplitis alienorum 16 4 Lasius claviger Acaridae Armacarus sp1 18 2 Acaridae Cosmoglyphus sp2 3 2 Acaridae Forcellinia sp10 8 1 Acaridae Froriepia sp1 5 3 Acaridae Schwiebea sp2 12 1 Acaridae Schwiebea sp3 3 2 Histiostomatidae Histiostoma sp2 3 1 Histiostomatidae Histiostoma sp3 1 1 Histiostomatidae Histiostoma sp11 3 1 Histiostomatidae Histiostoma sp12 26 2 Histiostomatidae Histiostoma sp13 7 1 Pygmephoridae Bakerdania sp9 3 2 Pygmephoridae Bakerdania sp12 1 1 Pygmephoridae Petalomium sp8 5 2 Scutacaridae Imparipes sp4 21 3 Scutacaridae Imparipes sp5 2 1 Scutacaridae Scutacarus sp20 2 1 Antennophoridae Antennophorus sp2 1 1 Laelapidae Cosmolaelaps sp21 1 1 Laelapidae Gaeolaelaps sp4 1 1 Uropodidae Uropolyaspis sp2 1 1 Lasius interjectus Acaridae Forcellinia sp10 4 1 Acaridae Froriepia sp1 11 1 Acaridae Schwiebea sp1 2 1 Acaridae Schwiebea sp2 1 1 Acaridae Schwiebea sp3 1 1 Histiostomatidae Histiostoma sp13 1 1 Histiostomatidae Histiostoma sp15 20 1 Pygmephoridae Petalomium sp8 8 1 Scutacaridae Imparipes sp4 3 1 Scutacaridae Scutacarus sp24 1 1 Scutacaridae Scutacarus sp25 2 1 Continued

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Table 14 continued ANT SPECIES MITE FAMILY MITE SPECIES # SLIDES # COLONIES Lasius interjectus Scutacaridae Scutacarus sp26 5 1 (continued) Lasius nearcticus Microdispidae Caesarodispus sp11 6 2 Acaridae Sancassania sp3 1 1 Lasius neoniger Acaridae Cosmoglyphus sp2 9 5 Acaridae Forcellinia sp1 19 6 Acaridae Forcellinia sp2 3 1 Histiostomatidae Histiostoma sp13 2 1 Histiostomatidae Histiostoma sp14 1 1 Histiostomatidae Histiostoma sp15 3 2 Histiostomatidae Histiostoma sp21 2 2 Histiostomatidae Histiostoma sp23 8 3 Pygmephoridae Petalomium sp8 1 1 Scutacaridae Imparipes sp12 1 1 Scutacaridae Imparipes sp13 1 1 Scutacaridae Scutacarus sp2 1 1 Scutacaridae Scutacarus sp22 1 1 Scutacaridae Scutacarus sp23 2 1 Scutacaridae Scutacarus sp27 3 2 Laelapidae Cosmolaelaps sp15 1 1 Oplitidae Oplitis sp1 10 6 Lasius speculiventris Acaridae Forcellinia sp10 1 1

Acaridae Schwiebea sp5 1 1

Histiostomatidae Histiostoma sp13 5 1 Histiostomatidae Histiostoma sp17 2 1 Microdispidae Caesarodispus sp11 1 1 Microdispidae Unguidispus sp10 1 1 Scutacaridae Scutacarus sp4 3 1 Scutacaridae Scutacarus sp6 4 1 Scutacaridae Scutacarus sp8 1 1 Oplitidae Oplitis (Uroplitana) sp11 1 1 Lasius umbratus Acaridae Armacarus sp1 3 3 Acaridae Forcellinia sp10 23 7 Acaridae Froriepia sp2 1 1 Acaridae Schwiebea sp1 14 6 Acaridae Schwiebea sp3 3 1 Continued

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Table 14 continued ANT SPECIES MITE FAMILY MITE SPECIES # SLIDES # COLONIES Lasius umbratus Acaridae Schwiebea sp5 1 1 (continued) Acaridae Schwiebea sp11 1 1 Astigmata undet undet gen sp3 1 1 Astigmata undet undet gen sp4 1 1 Histiostomatidae Histiostoma sp2 15 3 Histiostomatidae Histiostoma sp12 11 5 Histiostomatidae Histiostoma sp13 43 11 Histiostomatidae Histiostoma sp14 1 1 Histiostomatidae Histiostoma sp15 18 6 Histiostomatidae Histiostoma sp17 17 6 Microdispidae Caesarodispus sp11 6 2 Microdispidae Unguidispus sp10 11 4 Pygmephoridae Bakerdania sp12 3 2 Pygmephoridae Bakerdania sp13 3 2 Pygmephoridae Petalomium sp8 4 2 Scutacaridae Imparipes sp4 25 5 Scutacaridae Imparipes sp5 9 5 Scutacaridae Imparipes sp6 3 3 Scutacaridae Scutacarus sp4 27 9 Scutacaridae Scutacarus sp5 5 3 Scutacaridae Scutacarus sp6 2 2 Scutacaridae Scutacarus sp7 2 2 Scutacaridae Scutacarus sp8 1 1 Scutacaridae Scutacarus sp9 3 2 Scutacaridae Scutacarus sp20 2 2 Scutacaridae Scutacarus sp21 5 2 Scutacaridae Scutacarus sp28 1 1 Antennophoridae Antennophorus sp1 12 3 Laelapidae Cosmolaelaps sp8 1 1 Laelapidae Cosmolaelaps sp10 2 2 Laelapidae Gaeolaelaps sp4 1 1 Oplitidae Oplitis (Uroplitana) sp11 1 1 Uropodidae Uropolyaspis sp1 6 6 Continued

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Table 14 continued ANT SPECIES MITE FAMILY MITE SPECIES # SLIDES # COLONIES Myrmecina americana Histiostomatidae Histiostoma sp5 1 1 Histiostomatidae Histiostoma sp13 2 1 Myrmica americana Acaridae Forcellinia sp9 10 1 Acaridae Schwiebea sp7 1 1 Ponera pennsylvanica Scutacaridae Imparipes sp9 9 4

Scutacaridae Imparipes sp10 3 3

Scutacaridae Scutacarus sp12 1 1 Prenolepis imparis Acaridae Schwiebea sp12 3 1 Histiostomatidae Histiostoma sp27 2 2 Microdispidae Caesarodispus sp16 8 2 Scutacaridae Scutacarus sp15 2 2 Pyramica ohioensis Histiostomatidae Histiostoma sp21 1 1 Solenopsis molesta Acaridae Cosmoglyphus sp1 11 7

Acaridae Froriepia sp3 2 2

Pygmephoridae Bakerdania sp14 2 2 Scutacaridae Imparipes sp7 3 3 Scutacaridae Imparipes sp8 7 6 Tapinoma sessile Acaridae Forcellinia sp4 1 1 Scutacaridae Scutacarus sp19 1 1 Tetramorium caespitum Acaridae Forcellinia sp7 3 1

Acaridae Schwiebea sp9 1 1

Histiostomatidae Histiostoma sp10 2 1

Histiostomatidae Histiostoma sp14 12 2 Scutacaridae Imparipes sp11 16 4 Laelapidae Laelaspis nr. picketti 4 1 Oplitidae Oplitis sarcinulus 7 4

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ASTIG ASTIG FAMILY ASTIG SPECIES SPECIFICITY Acaridae Armacarus sp1 2 Acaridae Cosmoglyphus sp1 1 Acaridae Cosmoglyphus sp2 2 Acaridae Forcellinia sp1 1 Acaridae Forcellinia sp2 1 Acaridae Forcellinia sp3 2 Acaridae Forcellinia sp4 3 Acaridae Forcellinia sp5 1 Acaridae Forcellinia sp6 1 Acaridae Forcellinia sp7 1 Acaridae Forcellinia sp8 1 Acaridae Forcellinia sp9 1 Acaridae Forcellinia sp10 2 Acaridae Forcellinia sp11 1 Acaridae Froriepia sp1 2 Acaridae Froriepia sp2 1 Acaridae Froriepia sp3 1 Acaridae Sancassania sp1 1 Acaridae Sancassania sp3 1 Acaridae Schwiebea sp1 2 Acaridae Schwiebea sp2 2 Acaridae Schwiebea sp3 2 Acaridae Schwiebea sp4 1 Acaridae Schwiebea sp5 2 Acaridae Schwiebea sp6 1 Acaridae Schwiebea sp7 3 Acaridae Schwiebea sp8 3 Acaridae Schwiebea sp9 1 Acaridae Schwiebea sp10 1 Acaridae Schwiebea sp11 1 Acaridae Schwiebea sp12 1 Astigmata undet undet gen sp3 1 Astigmata undet undet gen sp4 1 Histiostomatidae Histiostoma sp1 2 Histiostomatidae Histiostoma sp2 2 Histiostomatidae Histiostoma sp3 1 Histiostomatidae Histiostoma sp4 2 Histiostomatidae Histiostoma sp5 3 Histiostomatidae Histiostoma sp6 2 Histiostomatidae Histiostoma sp7 2 Histiostomatidae Histiostoma sp8 1 Histiostomatidae Histiostoma sp9 1 Histiostomatidae Histiostoma sp10 1 Histiostomatidae Histiostoma sp11 1 Histiostomatidae Histiostoma sp12 2 Continued Table 15. Astigmata host specificity. All phoretic Astigmatid species are shown with codes designating their specificity levels. 1 = associated with only a single host species, 2= associated with multiple host species in the same genus, 3 = associated with multiple hosts in separate genera.

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Table 15 continued ASTIG ASTIG FAMILY ASTIG SPECIES SPECIFICITY Histiostomatidae Histiostoma sp13 3 Histiostomatidae Histiostoma sp14 3 Histiostomatidae Histiostoma sp15 2 Histiostomatidae Histiostoma sp16 1 Histiostomatidae Histiostoma sp17 2 Histiostomatidae Histiostoma sp18 2 Histiostomatidae Histiostoma sp19 2 Histiostomatidae Histiostoma sp20 2 Histiostomatidae Histiostoma sp21 3 Histiostomatidae Histiostoma sp22 1 Histiostomatidae Histiostoma sp23 1 Histiostomatidae Histiostoma sp24 1 Histiostomatidae Histiostoma sp25 1 Histiostomatidae Histiostoma sp26 1 Histiostomatidae Histiostoma sp27 1

HETERO HETERO FAMILY HETERO SP SPECIFICITY Microdispidae Caesarodispus sp2 1 Microdispidae Caesarodispus sp15 1 Microdispidae Caesarodispus sp16 1 Microdispidae Unguidispus sp10 2 Pygmephoridae Bakerdania sp3 1 Pygmephoridae Bakerdania sp5 1 Pygmephoridae Bakerdania sp7 1 Pygmephoridae Bakerdania sp9 1 Pygmephoridae Bakerdania sp11 2 Pygmephoridae Bakerdania sp12 2 Pygmephoridae Bakerdania sp13 1 Pygmephoridae Bakerdania sp14 3 Pygmephoridae Petalomium sp6 1 Pygmephoridae Petalomium sp8 2 Scutacaridae Diversipes sp1 1 Scutacaridae Imparipes sp1 2 Scutacaridae Imparipes sp2 2 Scutacaridae Imparipes sp4 2 Scutacaridae Imparipes sp5 2 Scutacaridae Imparipes sp6 1 Continued Table 16. Heterostigmatina Host specificity. All phoretic Heterostigmatid species are shown with codes designating their specificity levels. 1 = associated with only a single host species, 2= associated with multiple host species in the same genus, 3 = associated with multiple hosts in separate genera.

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Table 16 continued HETERO HETERO FAMILY HETERO SP SPECIFICITY Scutacaridae Imparipes sp7 1 Scutacaridae Imparipes sp8 1 Scutacaridae Imparipes sp9 1 Scutacaridae Imparipes sp10 1 Scutacaridae Imparipes sp11 1 Scutacaridae Imparipes sp12 1 Scutacaridae Imparipes sp13 1 Scutacaridae Imparipes sp14 1 Scutacaridae Pygmodispus sp1 1 Scutacaridae Scutacarus sp1 2 Scutacaridae Scutacarus sp2 2 Scutacaridae Scutacarus sp3 1 Scutacaridae Scutacarus sp4 2 Scutacaridae Scutacarus sp5 1 Scutacaridae Scutacarus sp6 2 Scutacaridae Scutacarus sp7 1 Scutacaridae Scutacarus sp8 2 Scutacaridae Scutacarus sp9 1 Scutacaridae Scutacarus sp10 1 Scutacaridae Scutacarus sp11 2 Scutacaridae Scutacarus sp12 3 Scutacaridae Scutacarus sp13 1 Scutacaridae Scutacarus sp14 1 Scutacaridae Scutacarus sp15 1 Scutacaridae Scutacarus sp17 1 Scutacaridae Scutacarus sp18 1 Scutacaridae Scutacarus sp19 1 Scutacaridae Scutacarus sp20 2 Scutacaridae Scutacarus sp21 1 Scutacaridae Scutacarus sp22 1 Scutacaridae Scutacarus sp23 1 Scutacaridae Scutacarus sp24 1 Scutacaridae Scutacarus sp25 1 Scutacaridae Scutacarus sp26 1 Scutacaridae Scutacarus sp27 1 Scutacaridae Scutacarus sp28 1

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MESO MESO FAMILY MESO SP SPECIFICITY Antennophoridae Antennophorus sp1 1 Antennophoridae Antennophorus sp2 1 Laelapidae Cosmolaelaps sp3 1 Laelapidae Cosmolaelaps sp4 1 Laelapidae Cosmolaelaps sp8 1 Laelapidae Cosmolaelaps sp9 1 Laelapidae Cosmolaelaps sp10 1 Laelapidae Cosmolaelaps sp13 1 Laelapidae Cosmolaelaps sp14 1 Laelapidae Cosmolaelaps sp15 1 Laelapidae Cosmolaelaps sp19 1 Laelapidae Cosmolaelaps sp21 1 Laelapidae Gaeolaelaps sp2 1 Laelapidae Gaeolaelaps sp3 1 Laelapidae Gaeolaelaps sp4 2 Laelapidae Gaeolaelaps sp6 1 Laelapidae Gaeolaelaps sp7 1 Laelapidae Gaeolaelaps sp8 1 Laelapidae Laelaspis cf. dubitatus 2 Laelapidae Laelaspis cf. vitzthumi 1 Laelapidae Laelaspis nr. picketti 1 Oplitidae Oplitis aktius 1 Oplitidae Oplitis alienorum 1 Oplitidae Oplitis exsectoidesorum 1 Oplitidae Oplitis sarcinulus 1 Oplitidae Oplitis sp1 1 Oplitidae Oplitis sp2 1 Oplitidae Oplitis sp3 1 Oplitidae Oplitis sp8 1 Oplitidae Oplitis sp9 1 Oplitidae Oplitis sp10 1 Oplitidae Oplitis (Uroplitana) sp11 2 Rhodacaridae undet gen sp1 1 Uropodidae Uropolyaspis sp1 1 Uropodidae Uropolyaspis sp2 1 Table 17. Mesostigmata Host Specificity. All phoretic Mesostigmatid species are shown with codes designating their specificity levels. 1 = associated with only a single host species, 2= associated with multiple host species in the same genus.

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PHORETIC ATTACHMENT SITE CODE TOTAL ASTIG FAMILY ASTIG SPECIES 1 2 3 4 5 6 7 8 SLIDES Acaridae Armacarus sp1 19 2 21 Acaridae Cosmoglyphus sp1 9 2 11 Acaridae Cosmoglyphus sp2 9 1 1 1 12 Acaridae Forcellinia sp1 1 5 9 4 19 Acaridae Forcellinia sp10 1 2 1 1 25 2 4 36 Acaridae Forcellinia sp11 1 1 Acaridae Forcellinia sp2 1 1 1 3 Acaridae Forcellinia sp3 3 3 Acaridae Forcellinia sp4 1 6 16 10 1 34 Acaridae Forcellinia sp5 1 1 Acaridae Forcellinia sp6 2 2 Acaridae Forcellinia sp7 1 2 3 Acaridae Forcellinia sp8 1 4 1 6 Acaridae Forcellinia sp9 9 1 10 Acaridae Froriepia sp1 1 8 4 3 16 Acaridae Froriepia sp2 1 1 Acaridae Froriepia sp3 2 2 Acaridae Sancassania sp1 1 1 Acaridae Sancassania sp3 1 1 Acaridae Schwiebea sp1 4 4 5 1 2 16 Acaridae Schwiebea sp2 1 4 8 13 Acaridae Schwiebea sp3 2 3 2 7 Acaridae Schwiebea sp4 2 2 4 Acaridae Schwiebea sp5 1 1 2 Acaridae Schwiebea sp6 1 1 Acaridae Schwiebea sp7 1 1 1 3 Acaridae Schwiebea sp8 1 2 6 6 15 Acaridae Schwiebea sp9 1 1 Acaridae Schwiebea sp10 1 1 Acaridae Schwiebea sp11 1 1 Acaridae Schwiebea sp12 1 1 1 3 Astigmata undet undet gen sp3 1 1 Astigmata undet undet gen sp4 1 1 Histiostomatidae Histiostoma sp1 1 2 3 Histiostomatidae Histiostoma sp2 1 3 14 18 Histiostomatidae Histiostoma sp3 1 1 Histiostomatidae Histiostoma sp4 1 1 2 Histiostomatidae Histiostoma sp5 2 1 1 9 13 Histiostomatidae Histiostoma sp6 1 27 28 Histiostomatidae Histiostoma sp7 1 1 1 3 Continued Table 18. Astigmata Phoretic Attachment Site Specificity. Number of specimens (slides) collected at each of 8 phoretic locations are shown. Phoretic attachment locations are as follows: 1-thorax/petiole, 2-antennal, 3-head, 4-between legs/coxae, 5-on legs, 6-gaster, 7-wing, 8-unknown.

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Table 18 continued PHORETIC ATTACHMENT SITE CODE TOTAL ASTIG FAMILY ASTIG SPECIES 1 2 3 4 5 6 7 8 SLIDES Histiostomatidae Histiostoma sp8 3 3 Histiostomatidae Histiostoma sp9 1 1 Histiostomatidae Histiostoma sp10 2 2 Histiostomatidae Histiostoma sp11 2 1 3 Histiostomatidae Histiostoma sp12 1 2 2 1 54 55 115 Histiostomatidae Histiostoma sp13 1 1 8 1 1 40 9 61 Histiostomatidae Histiostoma sp14 2 1 1 7 7 18 Histiostomatidae Histiostoma sp15 4 1 3 1 2 5 26 42 Histiostomatidae Histiostoma sp16 2 2 Histiostomatidae Histiostoma sp17 2 1 1 1 6 8 19 Histiostomatidae Histiostoma sp18 2 1 11 13 27 Histiostomatidae Histiostoma sp19 16 5 2 4 27 Histiostomatidae Histiostoma sp20 2 1 1 3 1 8 Histiostomatidae Histiostoma sp21 2 1 3 Histiostomatidae Histiostoma sp22 2 2 Histiostomatidae Histiostoma sp23 8 8 Histiostomatidae Histiostoma sp24 1 1 Histiostomatidae Histiostoma sp25 1 1 Histiostomatidae Histiostoma sp26 2 2 Histiostomatidae Histiostoma sp27 1 1 2 TOTAL 16 15 96 21 96 235 2 186 667

PHORETIC ATTACHMENT SITE CODE TOTAL HETERO FAMILY HETERO SPECIES 1 2 3 4 5 6 7 8 SLIDES Microdispidae Caesarodispus sp2 1 1 Microdispidae Caesarodispus sp11 2 3 6 11 Microdispidae Caesarodispus sp15 1 1 Microdispidae Caesarodispus sp16 7 1 8 Microdispidae Unguidispus sp10 2 10 12 Pygmephoridae Bakerdania sp3 1 1 Pygmephoridae Bakerdania sp5 5 5 Pygmephoridae Bakerdania sp7 1 1 Pygmephoridae Bakerdania sp9 1 2 3 Pygmephoridae Bakerdania sp12 1 2 1 4 Pygmephoridae Bakerdania sp13 1 2 3 Pygmephoridae Bakerdania sp14 2 1 3 Pygmephoridae Petalomium sp6 1 1 Continued Table 19. Heterostigmatina Phoretic Attachment Site Specificity. Number of specimens (slides) collected at each of 8 phoretic locations are shown. Phoretic attachment locations are as follows: 1-thorax/petiole, 2-antennal, 3-head, 4-between legs/coxae, 5-on legs, 6- gaster, 7-wing, 8-unknown.

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Table 19 continued PHORETIC ATTACHMENT SITE CODE TOTAL HETERO FAMILY HETERO SPECIES 1 2 3 4 5 6 7 8 SLIDES Pygmephoridae Petalomium sp8 2 3 1 12 18 Scutacaridae Diversipes sp1 1 2 3 Scutacaridae Imparipes sp1 5 42 16 63 Scutacaridae Imparipes sp2 5 3 8 Scutacaridae Imparipes sp4 3 2 37 16 58 Scutacaridae Imparipes sp5 1 1 5 1 3 11 Scutacaridae Imparipes sp6 1 1 1 3 Scutacaridae Imparipes sp7 1 2 3 Scutacaridae Imparipes sp8 6 1 7 Scutacaridae Imparipes sp9 3 6 9 Scutacaridae Imparipes sp10 2 1 3 Scutacaridae Imparipes sp11 15 1 16 Scutacaridae Imparipes sp12 1 1 Scutacaridae Imparipes sp13 1 1 Scutacaridae Imparipes sp14 1 1 Scutacaridae Pygmodispus sp1 2 2 Scutacaridae Scutacarus sp1 5 1 6 Scutacaridae Scutacarus sp2 10 1 1 1 13 Scutacaridae Scutacarus sp3 1 4 5 Scutacaridae Scutacarus sp4 15 5 1 1 1 7 30 Scutacaridae Scutacarus sp5 1 1 1 1 1 5 Scutacaridae Scutacarus sp6 1 5 6 Scutacaridae Scutacarus sp7 1 1 2 Scutacaridae Scutacarus sp8 2 2 Scutacaridae Scutacarus sp9 2 1 3 Scutacaridae Scutacarus sp10 5 1 6 Scutacaridae Scutacarus sp11 28 2 3 33 Scutacaridae Scutacarus sp12 30 2 1 5 38 Scutacaridae Scutacarus sp13 9 1 10 Scutacaridae Scutacarus sp14 2 2 Scutacaridae Scutacarus sp15 1 1 2 Scutacaridae Scutacarus sp17 1 1 2 Scutacaridae Scutacarus sp18 1 1 Scutacaridae Scutacarus sp19 1 1 Scutacaridae Scutacarus sp20 1 2 1 4 Scutacaridae Scutacarus sp21 1 2 1 1 5 Scutacaridae Scutacarus sp22 1 1 Scutacaridae Scutacarus sp23 2 2 Scutacaridae Scutacarus sp24 1 1 Scutacaridae Scutacarus sp25 1 1 2 Scutacaridae Scutacarus sp26 5 5 Scutacaridae Scutacarus sp27 1 2 3 Scutacaridae Scutacarus sp28 1 1 TOTAL 14 105 35 115 23 52 1 107 452

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PHORETIC ATTACHMENT SITE CODE TOTAL MESO FAMILY MESO SPECIES 1 2 3 4 5 6 7 8 SLIDES Antennophoridae Antennophorus sp1 11 1 12 Antennophoridae Antennophorus sp2 1 1 Laelapidae Cosmolaelaps sp3 4 4 Laelapidae Cosmolaelaps sp4 1 1 Laelapidae Cosmolaelaps sp8 1 1 Laelapidae Cosmolaelaps sp9 1 2 3 Laelapidae Cosmolaelaps sp10 1 1 2 Laelapidae Cosmolaelaps sp13 1 1 Laelapidae Cosmolaelaps sp14 2 2 Laelapidae Cosmolaelaps sp15 1 1 Laelapidae Cosmolaelaps sp19 1 3 4 Laelapidae Cosmolaelaps sp21 1 1 Laelapidae Gaeolaelaps sp2 1 1 1 8 11 Laelapidae Gaeolaelaps sp3 1 1 Laelapidae Gaeolaelaps sp4 2 2 Laelapidae Gaeolaelaps sp6 6 6 Laelapidae Gaeolaelaps sp7 8 2 2 12 Laelapidae Gaeolaelaps sp8 2 1 3 Laelapidae Laelaspis cf. dubitatus 1 2 3 6 Laelapidae Laelaspis cf. vitzthumi 2 2 Laelapidae Laelaspis nr. picketti 4 4 Oplitidae Oplitis aktius 3 3 Oplitidae Oplitis alienorum 15 1 16 Oplitidae Oplitis exsectoidesorum 2 2 Oplitidae Oplitis sarcinulus 1 6 7 Oplitidae Oplitis sp1 9 1 10 Oplitidae Oplitis sp2 1 1 Oplitidae Oplitis sp3 1 1 Oplitidae Oplitis sp8 2 2 Oplitidae Oplitis sp9 2 2 Oplitidae Oplitis sp10 1 1 Oplitidae Oplitis (Uroplitana) sp11 1 1 2 Rhodacaridae undet gen sp1 1 1 Uropodidae Uropolyaspis sp1 5 1 6 Uropodidae Uropolyaspis sp2 1 1 TOTAL 25 0 13 2 43 10 0 42 135 Table 20. Mesostigmata Phoretic Attachment Site Specificity. Number of specimens (slides) collected at each of 8 phoretic locations are shown. Phoretic attachment locations are as follows: 1-thorax/petiole, 2-antennal, 3-head, 4-between legs/coxae, 5-on legs, 6- gaster, 7-wing, 8-unknown.

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Figure 2. Aphaenogaster Species Accumulation Curves. Cumulative number of mite species, Sobs (Mao-Tau), per number of colonies collected.

Figure 3. Lasius Species Accumulation Curves. Cumulative number of mite species, Sobs (Mao-Tau), per number of colonies collected.

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Figure 4. Ponera and Solenopsis Species Accumulation Curves. Cumulative number of mite species, Sobs (Mao Tau), per number of colonies collected.

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ASTIG # # ASSOCIATION FAMILY ASTIG SPECIES ANT SPECIES SLIDES COLONIES TYPE Acaridae sp1 Lasius umbratus 25 1 soil in nest Acaridae Acotyledon sp1 Formica integra 3 1 soil in nest Acaridae Forcellinia sp5* Camponotus sp. 1 1 ant carcass Acaridae Forcellinia sp10* Lasius umbratus 1 1 ant carcass 2 1 soil in nest Acaridae Sancassania sp21 Aphaenogaster 35 1 ant food rudis complex Acaridae Sancassania undet Aphaenogaster 18 1 ant food tennesseensis Formica 2 1 ant food exsectoides Paratrechina 12 1 ant food faisonensis Prenolepis imparis 1 1 ant food 2 2 ant carcass Acaridae Schwiebea sp1*1 Lasius umbratus 46 1 ant food Acaridae Schwiebea sp3* Lasius claviger 2 1 ant carcass Acaridae Schwiebea sp8* Aphaenogaster 1 1 on brood fulva Lasius alienus 2 1 ant food Acaridae Schwiebia undet Lasius neoniger 3 2 soil in nest Histiostomatidae Glyphanoetus undet Aphaenogaster 5 1 soil in nest fulva Aphaenogaster 1 1 ant food tennesseensis Histiostomatidae Histiostoma sp2* Lasius umbratus 2 1 ant carcass 2 1 soil in nest Histiostomatidae Histiostoma sp12* Lasius claviger 1 1 ant carcass Histiostomatidae Histiostoma sp14* Aphaenogaster 4 1 soil in nest fulva 1 1 ant food Histiostomatidae Histiostoma sp15* Lasius umbratus 2 1 ant food Histiostomatidae Histiostoma sp17* Lasius umbratus 3 1 ant food 2 1 soil in nest Continued Table 21. Non-phoretic Astigmatid mites, their hosts, and locations where the mites were found in the nest. *designates a deutonymph which was also observed in a phoretic relationship. 1 Indicates a species (Sancassania sp2) which was reared in the lab. Undetermined species (undet) may actually be different species.

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Table 21 continued ASTIG # # ASSOCIATION FAMILY ASTIG SPECIES ANT SPECIES SLIDES COLONIES TYPE Histiostomatidae Histiostoma undet Aphaenogaster 7 1 soil in nest fulva 3 2 ant food Aphaenogaster 5 2 ant food picea complex Aphaenogaster sp. 8 1 ant food Aphaenogaster 6 1 ant food tennesseensis Camponotus 3 1 ant carcass chromaiodes Lasius alienus 1 1 soil in nest 5 1 ant carcass Lasius claviger 5 1 ant carcass 1 1 on brood Lasius umbratus 7 2 ant carcass 15 2 ant food 11 1 soil in nest

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PROSTIG PROSTIG # # ASSOCIATION ANT SPECIES FAMILY SPECIES SLIDES COLONIES TYPE Microdispidae Unguidispus sp10* Lasius umbratus 1 1 soil in nest Pygmephoridae Pygmephorellus sp1 Crematogaster 1 1 ant food lineolata Scutacaridae Imparipes sp1* Aphaenogaster 2 1 on brood fulva Aphaenogaster 2 1 on brood picea complex Scutacaridae Imparipes sp2* Aphaenogaster 2 1 ant food tennesseensis Scutacaridae Imparipes sp4* Lasius alienus 1 1 ant food Lasius claviger 3 1 ant carcass Lasius umbratus 2 2 ant food 1 1 on brood

Scutacaridae Imparipes undet sp Paratrechina 1 1 ant food faisonensis Scutacaridae Pygmodispus sp1* Lasius alienus 1 1 ant food Scutacaridae Scutacarus sp12* Aphaenogaster 1 1 ant carcass rudis complex Scutacaridae Scutacarus sp16 Formica integra 2 1 soil in nest Tarsonemidae undet gen sp1 Lasius alienus 3 2 ant food Tarsonemidae undet gen sp2 Camponotus sp. 1 1 ant carcass Tarsonemidae undet gen sp3 Aphaenogaster 2 1 fungus sp. Tarsonemidae undet gen sp3 Camponotus 2 1 ant food chromaiodes Tarsonemidae undet gen sp4 Aphaenogaster 3 1 ant food rudis complex Aphaenogaster 3 1 fungus sp. Tarsonemidae undet gen sp5 Aphaenogaster 3 2 ant food fulva Tarsonemidae undet gen sp6 Lasius umbratus 1 1 ant food undet gen sp1 Lasius alienus 1 1 fungus undet gen sp1 Formica integra 1 1 soil in nest undet gen sp1 Lasius alienus 1 1 soil in nest undet gen sp1 Formica integra 1 1 soil in nest

Continued Table 22. Non-phoretic Prostigmatid mites, their hosts, and locations where the mites were found in the nest. *designates a mite which was also observed in a phoretic relationship.

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# # ASSOCIATION MESO FAMILY MESO SPECIES ANT SPECIES SLIDES COLONIES TYPE Blattisocidae sp1 Lasius umbratus 1 1 ant food undet gen sp1 Aphaenogaster 9 1 soil in nest fulva Digamasellidae undet gen sp2 Aphaenogaster 1 1 ant food tennesseensis Digamasellidae undet gen sp3 Lasius umbratus 2 1 soil in nest Digamasellidae undet gen sp4 Aphaenogaster 2 1 soil in nest rudis complex Digamasellidae undet gen sp5 Lasius alienus 3 1 ant carcass Digamasellidae undet gen sp5 Lasius alienus 3 1 soil in nest Dinychus sp1 Formica integra 4 1 soil in nest Lasius umbratus 1 1 soil in nest undet gen sp1 Formica integra 1 1 soil in nest Laelapidae Cosmolaelaps imm sp Aphaenogaster 1 1 on pupa picea complex Aphaenogaster 4 1 ant food rudis complex 1 1 ant carcass 3 3 on brood Aphaenogaster 7 2 ant food tennesseensis Camponotus 1 1 on brood chromaiodes Formica integra 1 1 soil in nest Lasius alienus 1 1 soil in nest 1 1 on brood Lasius umbratus 1 1 soil in nest Paratrechina 3 1 ant food faisonensis Prenolepis 2 1 fungus on food imparis Laelapidae Cosmolaelaps sp1 Camponotus 1 1 on brood pennsylvanicus Laelapidae Cosmolaelaps sp2 Paratrechina 8 1 ant food faisonensis Laelapidae Cosmolaelaps sp7 Aphaenogaster 4 2 on pupa picea complex Laelapidae Cosmolaelaps sp11 Lasius umbratus 1 1 ant food 4 1 ant carcass Continued Table 23. Non-phoretic Mesostigmatid mites, their hosts, and locations where the mites were found in the nest. *designates a mite which was also observed in a phoretic relationship.

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Table 23 continued

# # ASSOCIATION MESO FAMILY MESO SPECIES ANT SPECIES SLIDES COLONIES TYPE Laelapidae Cosmolaelaps sp16 Aphaenogaster 1 1 on pupa picea complex Laelapidae Cosmolaelaps sp18 Aphaenogsater 2 1 soil in nest tennesseensis Laelapidae Cosmolaelaps sp20 Lasius alienus 1 1 soil in nest Laelapidae Cosmolaelaps sp21* Lasius claviger 4 1 ant carcass 1 1 on brood Laelapidae Gaeolaelaps sp1 Solenopsis 4 1 on brood molesta Laelapidae Gaeolaelaps sp5 Aphaenogaster 1 1 ant food tennesseensis Laelapidae Gaeolaelaps sp6* Formica integra 3 1 soil in nest Laelapidae Laelaspis n.sp. nr. Aphaenogaster 4 2 ant food vitzthumi tennesseensis Laelapidae sp1 Aphaenogaster 3 1 ant carcass tennesseensis 1 1 ant food Laelapidae undet gen sp1 Brachymrymex 1 1 near brood depilis Macrochelidae Macrocheles sp1 Formica integra 2 1 soil in nest Macrochelidae Macrocheles sp2 Aphaenogaster 1 1 ant food tennesseensis Gamasiphis sp1 Lasius alienus 1 1 soil in nest Oplitidae Oplitis alienorum* Lasius alienus 1 1 ant carcass 1 1 soil in nest 1 1 ant food Holaspina sp1 Aphaenogaster 2 1 ant food tennesseensis Parholaspididae Holaspina sp2 Formica integra 1 1 soil in nest Parholaspididae Holaspina sp3 Lasius alienus 1 1 on brood 3 2 soil in nest Circocyllibanidae Trichocylliba sp1 Lasius umbratus 10 2 on brood Trichouropoda sp1 Aphaenogaster 3 2 ant food tennesseensis 2 2 soil in nest Uropodellidae Uropodella laciniata Crematogaster 1 1 soil in nest lineolata Uropodidae undet gen sp1 Pyramica 1 1 soil in nest ohioensis Uropodidae undet gen sp3 Aphaenogaster 1 1 soil in nest fulva Uropodidae Uropolyaspis sp1* Lasius umbratus 1 1 soil in nest 8 1 ant carcass

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Literature Cited

Bhatkar, A. and W. H. Whitcomb. (1970). Artificial diet for rearing various species of ants. The Florida Entomologist, 53: 229-232.

Colwell, R. K. (2005). EstimateS: Statistical estimation of species richness and shared species from samples. Version 7.5. User's Guide and application published at: http://purl.oclc.org/estimates.

Coovert, G. (2005). The ants of Ohio (Hymenoptera: Formicidae). Columbus, OH: Ohio Biological Survey, Inc., 196 pp.

Ebermann, E. (1981). Imparipes (Imparipes) brevitarsus n.sp. (Acari, Scutacaridae), eine mit ameisen vergesellschaftete neue milbenart aus Österreich. Mitteilungen des Naturwissenschaftlichen Vereines für Steiermark, 111: 183-192.

Ebermann, E. and G. Rack. (1982). Zur biologogie einer neuen myrmecophilen art der gattung Petalomium (Acari, Pygmephoridae). Entomologische Mitteilungen aus dem Zoologiischen Museum Hamburg, 115: 175-191.

Franks, N. R., Healey, K. J., & Byrom, L. (1991). Studies on the relationship between the ant ectoparasite Antennophorus grandis (Acarina: Antennophoridae) and its host Lasius flavus (Hymenoptera: Formicidae). Journal of the Zoology (London), 225: 59-70.

Hirschmann, W. (1993). Gangsystematik der Parasitiformes. Teil 550. Bestimmungstabellen der Uropodiden der Erde, Atlas der Ganggattungen der Atrichopygidiina. Acarologie, Schriftenreihe für vergleichende Milbenkunde, Nürnberg, 40: 292-370.

Hölldobler, B. and E. O. Wilson. (1990). The Ants, Cambridge: Harvard Press.

Hunter, P. E. and M. H. Farrier. (1975). Mites in the genus Oplitis Berlese (Acarina: Uropodidae) associated with ants (Hymenoptera: Formicidae) in the southeastern United States Part I. Acarologia, 17: 595-623.

104

Hunter, P. E. and M. H. Farrier. (1976). Mites in the genus Oplitis Berlese (Acarina: Uropodidae) associated with ants (Hymenoptera: Formicidae) in the southeastern United States Part II. Acarologia, 18: 20-50.

Ivanov, K. (2010). Ohio Ants. Retrieved May, 2010 from http://www.antweb.org.

Janet, C. M. (1897). Sur les rapports de l’Antennophorus uhlmanni Haller avec le Lasius mixtus Nyl. Compte rendus de l’Academie des sciences (Paris), 124: 583-585.

Johnson, N. F. (2007). Hymenoptera name server version 1.5. Retrieved from http://hns.osu.edu/.

Karg, W. (1989). Acari (Acarina), Milben, Unterordnung Parasitiformes (Anactinochaeta), Uropodina Kramer, Schildkrötenmilben (Vol. 67). Die Tierwelt Deutschlands, Jena: Gustav Fischer Verlag, p. 203.

Krantz, G. W. and B. D. Ainscough. (1990). Acarina: Mesostigmata (Gamasida). In D. L. Dindal (Ed.), Soil Biology Guide (pp.583-665), New York: John Wiley and Sons, Inc.

MacArthur, R. H. and E. O. Wilson. (1967). The Theory of Island Biogeography. Princeton, : Princeton University Press.

Margolis, L., G. W. Esch, J. C. Holmes, A. M. Kuris, and G. A. Schad. (1982). The use of ecological terms in parasitology (report of an ad hoc committee of the American society of parasitologists). The Journal of Parasitology, 68: 131-133.

Neece, K. C. and D. P. Bartell. (1982). A Faunistic Survey of the Organisms Associated with Ants of Western Texas, No. 25. Graduate Studies Texas Tech University. Lubbock, TX: Texas Tech University Press, 36p.

OConnor, B. M. Keys to the genera of astigmatid mites. Acarology Summer Program 2008. Unpublished.

OConnor, B. M. (2009) Cohort Astigmatina. In G. W. Krantz & D. E. Walter (Eds.), A Manual of Acarology (3rd ed.), (pp. 565-657). Lubbock, TX: Texas Tech University Press.

Rettenmeyer, C.W. (1962). Arthropods associated with neotropical army ants with a review of the behavior of these ants (Arthropoda; Formicidae: Dorylinae). Ph.D. Dissertation. University of Kansas, Lawrence, KS. 644p.

Rohde, C. J. Jr. (1956). A modification of the plaster-charcoal technique for the rearing of mites and other small arthropods. Ecology, 37: 843-844.

105

Savulkina, M. M. (1981). Systematics, ecology, and distribution of mites of the family Pygmephoridae Cross, 1965 (Acari, Trombidiformes). Entomological Review, 60: 163-180.

Schlick-Steiner, B. C., F. M. Steiner, H. Konrad, B. Seifert, E. Christian, K. Moder, C. Stauffer, and R. H. Crozier. (2008). Specificity and transmission mosaic of ant nest-wall fungi. Proceedings of the National Academy of Sciences, 105: 940-943.

Walter, D. E., E. E. Lindquist, I. M. Smith, D. R. Cook, and G. W. Krantz. (2009). Order Trombidiformes. In G. W. Krantz & D. E. Walter (Eds), A Manual of Acarology (3rd ed.), (pp. 233-420). Lubbock, TX: Texas Tech University Press.

Wurst, E. (2001). The life cycle of Lemanniella minotauri n. sp. and the erection of the new family Lemannillidae (Acari: Astigmata). Stuttgarter Beiträge zur naturkunde, Serie A (Biologie), 621: 1-34.

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Chapter 3: Mites Associated with Messor pergandei

The Sonoran Desert occupies the southwestern portion of Arizona and extends into California and Mexico. It is a harsh environment characterized by little rainfall (<25 cm a year), shrubby and widely spaced vegetation, saguaro cacti (Carnegiea gigantea), and high daily summer temperatures. The soil is dry, sandy, and fine (Wheeler &

Wheeler, 1973). Many plants and animals have adapted to survival in the desert biome, and some are highly dependent on one another. These interspecies co-dependencies create a complex ecosystem, with many components yet to be discovered. To contribute to the current knowledge base, this section centers around a granivorous desert ant,

Messor (=) pergandei, and its acarine associates.

Interdependence in the Desert: Seed Harvester Ants

Seed harvester ants fall into 4 genera (all in subfamily Myrmecinae):

Pogonomyrmex, Pheidole, Aphaenogaster, and Messor (Johnson, 2000; 2001). Seeds are a more reliable food source in the desert than typical ant resources such as insect prey or honeydew from homopterans (Tevis, 1958). Pogonomyrmex rugosus and Messor pergandei are the two most conspicuous and thoroughly studied species of seed harvesters found in the southwestern United States (Johnson, 2000; 2001). Almost all species of Pogonomyrmex and Messor rely heavily on seed harvesting but also take insect

107 prey when available. Most Aphaenogaster species are opportunistic seed harvesters, and only about 30-40 of the approximate 100 Pheidole species in the United States are granivorous (Johnson, 2001).

Many desert plant populations remain in the seed stage for long periods of time awaiting a wet season or other necessary environmental conditions. Seed predation by harvester ants might seem a detriment to the desert plant ecosystem; however, desert plants compensate for the losses with improved seed dispersal, protection, and nutrients that increase seedling survivorship. Seed dispersal primarily occurs when foraging workers sometimes drop their seeds along the foraging trail, do not clip the seed radicle prior to storage in the nest, or deposit viable seeds along with the husks into the fertile chaff (refuse) piles surrounding the nest entrance (O’Dowd & Hay, 1980; Rissing, 1986;

Steinberger et al., 1991; Tevis, 1958). Ants also provide soil aeration for the desert, through the creation of galleries and chambers, and they mix deep and upper layers of soil and incorporate organic refuse into the soil (Beattie & Culver, 1983). Finally, because there are relatively few other insects in the desert, ants make up a large portion of the diets of many reptiles and birds (Wheeler & Wheeler, 1973).

Associates of Seed Harvesters

Generally seed harvester ants have received little attention in studies of myrmecophiles. Mites associated with Messor spp. are shown in Table 27 located at the end of this chapter. These and other seed harvester associates can also be found in the full mite associate list described in Chapter 1: Appendix A (Table 29 to Table 32).

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Messor pergandei

Natural History

Messor pergandei is one of the most frequently encountered ants in the Sonoran

Desert communities of Arizona. It is a small, shiny, black ant species with polymorphic workers (2½ -7 mm in length); like other desert ants it has a psammophore (stiff beard- like hairs on the underside of the head that aid in the transport of sandy soil) (Wheeler &

Wheeler, 1973). M. pergandei form populous, long-lived colonies with an estimated

30,000 to 50,000 workers (Johnson, 2000). It is known to have extensive foraging columns up to 40 m long with several thousand workers in a column (Johnson, 2000;

Wheeler & Rissing, 1975b). This species forms conspicuous crater entrances (usually 2-

3 entrances per nest), each of which is surrounded by a chaff pile (Wheeler & Rissing,

1975a). Nests have been estimated to span about 15.5 m (50 ft) in diameter underground and can be deeper than 4 m (Tevis, 1958). The chamber closest to the surface, known as the vestibule, is the processing chamber for seeds. Below the vestibule are the chambers in which husked seeds are stored. Brood is located in even deeper chambers, while the queen is likely in the lowest chamber, but no study has successfully recovered her (Tevis,

1958; Wheeler & Rissing, 1975a).

Mating flights in this species have been noted mainly in February under clear conditions when temperatures reach approximately 22º C. Alates gather in the vestibule prior to departure, and as in other ant species, they out, mate, and upon landing females begin excavation of a nest (Pollock & Rissing, 1985).

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A geographical transition in colony founding behavior has been reported in this species of ant. After the mating flight, unrelated M. pergandei dealate gynes (wingless mated females) can be frequently observed cooperatively founding colonies

(pleiometrosis) in Arizona and southeastern California, while colonies are almost exclusively founded by single dealates (haplometrosis) in southwestern California (Cahan et al., 1998; Rissing & Johnson, 2000).

The Scope of this Study

The biology, behavior, and ecology of M. pergandei has been studied in detail by many researchers, but no previously published studies have mentioned the mites found in association with this common ant species. S. W. Rissing and W. H. Schaedla deposited

12 mite specimens (4 species) taken from M. pergandei alates, into the Acarology

Collection at The Ohio State University (in 1997 and 2007), an action from which this study stems. Two specimens were Petalomium sp. (Heterostigmatina: Pygmephoridae) collected in 2007. The other ten specimens belonged to the Astigmata. They include representatives of Cosmoglyphus sp. (4 slides, from 1997 and 2007) and Forcellinia sp.

(1 slide from 1997), family Acaridae and Lemanniella sp. (5 slides all from 2007, taken only from female alates), family Lemanniellidae.

To follow up on these initial collections the following study aims to answer the questions: 1) What mite phoretic species are associated with M. pergandei alates?, 2) Do the mites have specific phoretic attachment sites?, 3) Are the phoretic mites preferentially using the female alates rather than the males for dispersal?, 4) Are mite infestations similar for alates and dealates?, 5) Do the phoretic mites appear in the chaff piles?, and 6)

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If phoretic mites are successfully reared in a laboratory setting, what roles do they play in the nests?

Methods

Locality and collection dates

The majority of the mite collections from M. pergandei alates, dealates and nests were made at W McCartney Rd, East of I-10 (32.9398º N, 111.6641º W), Casa Grande,

Pinal County, Arizona. A couple of collections were made at another nearby site: N Cox

Rd at W McCartney Rd, East of I-10 (32.9299º N, 111.6891º W). The dominant vegetation of the study area is the creosote bush, Larrea divaricata, with small numbers of saguaro and fishhook barrel cacti. Collections were made from February 12 to

February 22, 2008. The two collection sites are located very near to each other and encompass the same general location at which Rissing and Schaedla made their collections on March 7, 1997, and February 21, 2007.

Field Collection

In order to collect alates prior to their mating flights, a standard shovel was used to excavate the vestibule chambers of the colonies to a depth of 1 to 2 feet around the entrance craters. If no alates were apparent after a few excavation attempts at each entrance, the colony was assumed to not have any accessible alates. If alates were uncovered, excavation continued until no more alates were found. Alates were deposited individually into empty 2 ml Eppendorf tubes which were then labeled with numbers in chronological order using permanent marker on the lid. Tubes with alates from different colonies were placed into separate plastic bags and given a field number in the format

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“collector initials-08-monthday-colony number”. Care was taken to determine boundaries of the colony, so as not to sample the same colony twice. In one nest, 30 workers were collected.

Dealates were collected by searching for newly excavated nests which appeared as small holes surrounded by half-moon shaped piles of sand. Usually one shovel full of sand approximately 10 inches deep was adequate to gain access to the nest chamber.

Dealates seemed to prefer disturbed soil (edges of dirt roads, dry stream banks, and horse hoof prints), so often simply following horse tracks yielded a queen in every couple of prints. Because this area is in the geographical range of pleiometrosis, multiple queens sometimes were collected from the same hole. When dealates were uncovered, they were deposited individually into 15 ml Falcon tubes which then were labeled chronologically.

Chaff accumulates as a contiguous mass outside of active nest entrances and can be easily peeled up from the sand like carpet. Chaff piles were collected from six nest entrances (including multiple entrances) prior to excavation and placed inside 473 ml (16 oz) clear plastic deli containers with lids for transport. Collapsible nylon Berlese funnels

(using 40 watt bulbs) were hung in a garage (for 2 days) to extract any chaff-inhabiting arthropods into Nasco Whirl-Paks® approximately 50% full of 95% ethanol.

After returning from the field, live alates and dealates were grasped between thumb and forefinger, and all surfaces were carefully examined for phoretic mites under a stereomicroscope. Each ant’s field number, sex, and the number and location of phoretic mites were recorded in a notebook.

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Almost all alates were then returned to their original Eppendorf tube, a field number label written in indelible ink was inserted, and the tube was filled with 95% ethanol. Five female alates with phoretic mites were kept alive (with wings artificially removed) to see if they would lay eggs which might induce mite development. The dealates, however, were to be reared, so each was individually deposited into clean

Falcon tubes which had been half filled with tap water and plugged with a cotton ball. A piece of aluminum foil was wrapped around the tube to block light. This created a dark, humid chamber in which the gyne could begin laying eggs. A total of 23 dealates without mites were paired with some of the alates collected on the last day (male and female, 18 live and 5 dead) to see if mites would transfer between hosts.

Laboratory Observation of Dealates

All collections (ants and chaff) were transported to the Acarology lab at The Ohio

State University. Dealates were transferred to glass culture tubes (16 x150 mm) half full of distilled water and plugged with cotton on February, 27th, 2008. Another cotton ball was used to plug the opening of the test tube, and foil was wrapped around the tube.

These tubes were put into two incubators maintained at 35º C. Daily to weekly checks were made to note and remove any dead dealates and to record colony development. The dead dealates were inspected for mites to salvage any which remained. The first workers

(minims) were noted on April 2nd, 2008.

Colonies were transferred to nest chambers on April 7th. Laboratory nest chambers consisted of two types. Type 1 was a translucent plastic storage container (11 x

20.5 x 34.5cm) partially filled with a substrate comprised of 9 parts Plaster of Paris

113 mixed with 1 part activated carbon. Several glass test tubes (16 x 150 mm) were placed on top of the substrate. These tubes either were half filled with water and plugged with cotton or left empty (as potential storage chambers). The substrate was moistened and the test tube containing the dealate and minims was inserted among the new tubes. The rim of the box was coated with fluon “Insect-a-slip” (BioquipTM, Rancho Dominguez, CA) in order to prevent ants from escaping. All of the nests were placed in incubators. Type 2 nests consisted of two 20.3 x 25.4 mm (8 x 10 in) pieces of glass with 6.35 mm (1/4 in) square wooden dowels holding the glass apart at the perimeter. Dowels were wood glued together at the corners except for the top piece which was removable allowing access into the colony. Binder clips were used to hold the nest together. Sifted sandy soil from the original collection site in Arizona (which had been heated in a microscope slide oven for a week) was poured into the nest chamber to about halfway. It was then moistened by allowing the base of the chamber to sit in a tub of water. Dealates and the contents of their rearing test tubes were gently transferred into individual nest chambers. Dealates quickly excavated a chamber a couple of centimeters below the soil surface to which they transferred their brood. Nests were stored vertically in incubators.

When the minims began to eclose, thistle bird seed and meal worms (Tenebrio molitor) were given to the colony on a piece of foil (Type 1 nests) or placed directly onto the soil surface (Type 2 nests). Workers were seen moving the seeds, husking the seeds, and placing them among the larvae, but they paid little attention to the meal worms. On

April 15th one of the incubators malfunctioned and temperatures rose above 55º C, killing

114 all but one of the dealates. The remaining colonies were stored in the functional incubator and weekly checks continued until all colonies were dead (July 24, 2008).

Recovery, Preservation, and Identification of Mites from Ants and Chaff

Contents of the Eppendorf tubes holding the alates with phoretic mites were examined and mites counted and identified under a stereomicroscope. Mites were rarely observed still clinging to the host, so examination of tube contents was required. The contents were emptied into a small petri dish, and the Eppendorf tube was rinsed with ethanol into the petri dish. Slow circular swirling motions usually brought most of the mites into the center of the dish. Numbers of apparently different species of mites were recorded, and specimens (usually 1-3 mites) of each morphospecies were placed into lactophenol for clearing prior to slide making. Whenever possible, specimens of each species were also retained in 95% ethanol, but in cases of low mite numbers, individuals were placed into lactophenol only, as identification was vital. The host alate was placed with a label into a glass shell vial plugged with cotton, then into a glass (20 ml) scintillation vial (in some cases with the remaining mites in another shell vial). Several attempts were usually required to recover the field-documented number of mites, but sometimes despite numerous rinses and careful inspection, collection of all mites was not successful.

Mite attachment sites on host alates were pieced together by merging lab and field notes; the number of mites of each species recovered from a host was compared to field notes as to the number at each bodily location. For example in the case of KAU-08-0218-

2 #39, field records indicated that there were five mites under the head and four mites

115 between the 3rd coxae. Five Lemanniella sp. and four Petalomium sp. were recovered from the vial, thus was it assumed that the five mites under the head were Lemanniella sp., and the four mites between the coxae were Petalomium sp. Generally this method was successful for documenting the locations of the mites, but in cases of undocumented, lost specimens, or if there were equal numbers of mites of different species in different locations, exact locations could not be reasonably determined and these mites were not included in the analyses.

Dead dealates from laboratory nest chambers or culture tubes were treated in a similar way. Their rearing tubes and brood were carefully inspected and contents rinsed.

Recovered mites were placed in lactophenol or in 95% ethanol, if they were part of a large series of the same species.

Berlese chaff outputs were searched for any arthropods. Morphologically distinct mites were placed in lactophenol and duplicates were stored in 95% ethanol. Insects and miscellaneous larvae were put into a separate vial but not identified.

Cleared mites were mounted on slides as described in “Chapter 2: Preservation and Identification of Ants and Mites”. Mites on slides were identified to genus and morphospecies using several keys including the key to families (OConnor, 2009) and unpublished key to the genera for the Astigmata (OConnor, 2008), Savulkina’s (1981) key for pygmephoroids (Heterostigmatina), and the Walter et al. (2009) key to the families of soil Prostigmata (for non-Heterostigmatids).

A mite species accumulation curve was constructed from the accumulated number of mite species taken from alates (Sobs Mao Tau) per field colony sampled. Sobs Mao Tau

116 was calculated using the diversity statistical program, EstimateS (Version 7.5, Colwell,

2005).

Results

Colony Collections

A total of 330 alates (140 males, 190 females) was collected from a total of 16 ant colonies. Numbers of alates varied from 2 to 61 per colony. Most colonies produced both sexes, but ratios were usually skewed toward one sex or the other; rarely were they found in approximately equal proportions. Only five colonies were collected with only 1 of the sexes). Table 24 summarizes the alate collections. The 30 workers collected did not have mites.

TOTAL TOTAL TOTAL # M # F % M % F COLONY # ALATES M F WITH WITH WITH WITH KAU-08-0214-1 21 15 6 2 6 13.33 100 KAU-08-0216-1 57 9 48 0 11 0 22.92 KAU-08-0216-2 37 7 30 0 6 0 20 KAU-08-0217-1 3 2 1 0 0 0 0 KAU-08-0217-21 4 1 3 0 2 0 66.67 KAU-08-0217-4 10 0 10 0 7 n/a 70 KAU-08-0217-5 1 0 1 0 0 n/a 0 KAU-08-0218-1 6 5 1 0 0 0 0 KAU-08-0218-21 2 0 2 0 2 n/a 100 KAU-08-0218-3 4 0 4 0 0 n/a 0 KAU-08-0218-4 61 56 5 4 2 7.14 40 KAU-08-0219-1 48 2 46 0 46 0 100 KAU-08-0220-1 14 14 0 0 0 0 n/a KAU-08-0220-2 17 4 13 0 0 0 0 KAU-08-0220-3 26 10 16 0 0 0 0 KAU-08-0221-1 19 15 4 0 0 0 0 Totals 330 140 190 6 82 4.29 43.16 Table 24. Alate Summary and Sex Preference of Mites. Total number of male (M) and female (F) alates collected per colony and number with phoretic mites. 1Signifies two colonies collected at the Cox Rd site. All other colonies were collected at the W McCartney Rd site. “n/a” is used when no alates of that sex were found within the colony.

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Mite Association with Female versus Male Alates

Alates from 16 colonies were collected consisting of 140 males and 190 females.

Of these 16 colonies, 8 colonies had mites. A total of 90 males and 150 female alates were collected from the mite infested colonies and 88 total alates had phoretic mites. The percentage infested (using only the infested colonies) was calculated for both alate sexes.

Infestation was 6.7% for males (N=6) and 54.7% for females (N =82). The mean number of mites per alate was 6.8 mites (range 1 to 30, SD=7.0) and the median was 4.5 mites.

The detailed findings are shown in Table 24.

Phoretic Mite Species and Abundance

A total of 593 mite specimens were collected belonging to 6 species. Three species were previously collected by Rissing and Schaedla and deposited at the

Acarology Lab: Lemanniella sp., Petalomium sp., and Forcellinia sp. Their fourth mite species (Cosmoglyphus sp.) was not collected in any of the colonies sampled in 2008, but three additional species were found: Armacarus sp. (Astigmata: Acaridae), Histiostoma sp. (Astigmata: Histiostomatidae), and Unguidispus sp. (Heterostigmatina:

Microdispidae). One mite belonging to the family was found in a vial with a female alate, but it is believed to be a contaminant, since no members in this family are known to be phoretic, but many are found in soil habitats.

Approximately 83% of the mites collected were Armacarus sp. (N =490), 488 of which were found in a single colony. Lemanniella sp. constituted 10% and Petalomium

118 sp.7%. Forcellinia sp., Histiostoma sp. and Unguidispus sp. were collected in very low numbers. Total abundance of the mite species per colony is shown in Table 25.

Armacarus

Forcellinia

Petalomium

Histiostoma

Lemanniella

Unguidispus MITES TOTAL COLONY # M F M F M F F F M KAU-08-0214-1 0 1 2 15 0 2 0 0 1 20 KAU-08-0216-1 0 0 0 0 0 13 0 0 0 13 KAU-08-0216-2 0 0 0 0 0 6 0 0 0 6 KAU-08-0217-2 0 0 0 9 0 0 0 0 0 9 KAU-08-0217-4 0 0 0 20 0 2 0 0 0 22 KAU-08-0218-2 0 0 0 6 0 4 0 0 0 10 KAU-08-0218-4 0 1 0 0 4 1 0 0 0 6 KAU-08-0219-1 0 488 0 4 0 10 2 2 0 507 TOTALS 0 490 2 54 4 38 2 2 1 593 SPECIES 82.63 9.44 7.08 0.34 0.34 0.17 % OF TOTAL Table 25. Host Sex Preference of Mite Species. Number of individuals of the 6 mite species found in each colony on male (M) and female (F) alates and percentages of the total mite fauna that each species comprises (N=593).

Attachment Site Specificity

Observation of living ant specimens prior to placement in alcohol is necessary to gain a clear understanding of the mite attachment sites. Unfortunately, in some cases, the number of field-documented mites did not directly correspond to the number of recovered mites. Eighty-two of the anticipated mites were not recovered from their respective vials; however, 40 previously undocumented mites were recovered in other vials. Specimens that were documented in the field, but not subsequently recovered in the lab were not included in the attachment site specificity summary, and any which were undocumented in the field were categorized as specimens with “unknown” phoretic locations. 119

Table 26 details the number of mites found in phoretic locations on the host for each of the 6 species. Armacarus sp. were found in various sites on the body, but the majority (388 of the 490 mites) were found attached to the gaster, usually anterior- ventrally. Additionally, Armacarus sp. would often arrange themselves in the same direction, with legs I pointing toward the posterior of the ant. Lemanniella sp. were found primarily (48 of 56 mites) under the head, beneath the psammophore. Petalomium sp. were generally (29 of 42 mites) found between all coxae in the ventral position, but especially between the second and third coxae. Forcellinia, Histiostoma, and

Unguidispus species were found in too low of numbers to make generalizations concerning their phoretic location.

LOCATION Armacarus Lemanniella Petalomium Forcellinia Histiostoma Unguidispus Ventral head 7 48 0 0 1 1 Neck 0 0 3 0 0 0 Dorsal thorax 28 1 1 0 0 0 Lateral thorax 11 0 0 0 0 0 Ventral coxae 6 0 29 0 0 0 On leg 11 0 3 2 0 0 Petiole 16 0 0 0 0 0 Gaster 388 0 0 0 0 0 Unknown 23 7 6 0 1 0 Total 490 56 42 2 2 1 Table 26. Phoretic Locations. Number of mites for each species found at corresponding locations on the hosts.

Mites on Dealates

A total of 44 dealates from newly excavated nests was collected, all from the W

McCartney Rd location. Four dealates were found as pairs, founding their nest pleimetrotically; in both cases only one of the two had mites. Twelve dealates had mites 120

(37.5%) each with an average of 6.4 mites (range from 1 to 29, SD = 8.6) per ant. Median number of mites was 3 per dealate, a slightly lower number of mites and infestation rate than seen in alates. Identification of many of the mites was not possible because they were put into the rearing containers with the founding queens, but those that were identified belonged to the same species as found on alates.

Dealates that died in the nest chambers (unknown causes or incubator malfunction) were sometimes found with mites still on them, but often, no mites were recoverable. In one case, a dead dealate had developed white fungal mycelia, and a

Petalomium sp. mite was observed feeding on the fungus. Inspection of colonies prior to and after death of the gyne yielded no other mite activity observations. Lack of apparent mite activity might be attributed to inadequate observation time in an attempt to minimize disturbance to the colony as it grew. Aside from the Petalomium sp. observation, mites such as Armacarus sp. were observed on the host up to a week after death still in their deutonymphal stage suggesting inadequate cues for further development. It was expected that if the appropriate resources were available for the development of the mites, subsequent instars would be detected fairly quickly.

In all instances (N=23) of dealates without mites paired with alates possessing phoretic mites, transfer between hosts was not evident. Mites remained on their original hosts. In cases of alates with wings artificially removed, eggs were laid, but mite development did not occur. It is possible that lack of mite development was due to laboratory conditions.

Mites in Chaff Piles

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Only 3 of the 6 chaff piles had mites and generally in low numbers. The majority of those recovered from chaff piles were small, soft bodied mites in the Prostigmata. Five morphospecies of mites were collected in the chaff piles, but all were represented by singletons except for one, a very common species found in two of the chaff piles,

Tydeidae sp. Other mites include Procaeculus sp. (), Anystidae sp., a non- phoretic female of Bakerdania sp. (Pygmephoridae), and Tetranychidae sp.

Voucher Specimens

Voucher specimens for each morphospecies are deposited at the Acarology

Collection at The Ohio State University under the following names and code numbers

(number of slides of each species are shown in parentheses, but only the OSAL number listed is the voucher specimen): Armacarus Messor sp1- OSAL0007082 (71 total slides),

Lemanniella Messor sp1-OSAL0007039 (28 total slides), Petalomium Messor sp1-

OSAL0007035 (36 total slides), Forcellinia Messor sp1-OSAL0007105 (2 total slides),

Histiostoma Messor sp1-OSAL0092942 (2 total slides), Unguidispus Messor sp1-

OSAL0007060 (1 total slide), Nanorchestidae Messor sp1-OSAL0092938 (1 total slide),

Tydeidae Messor sp1-OSAL0102747 (7 total slides), Procaeculis Messor sp1-

OSAL0102748 (1 total slide), Anystidae Messor sp1-OSAL0102750 (1 total slide),

Bakerdania Messor sp1-OSAL0102740 (1 total slide), Tetranychidae Messor sp1-

OSAL0102749 (1 total slide). A voucher specimen of Messor pergandei is deposited in the Ohio State University Insect Collection under the specimen number OSUC0359951.

Discussion

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Yearly Variation and Mite Species Richness

A total of six mite species were found associated phoretically with M. pergandei reproductives: Armacarus sp. (Acaridae), Lemanniella sp. (Lemanniellidae), Petalomium sp. (Pygmephoridae), Forcellinia sp. (Acaridae), Histiostoma sp. (Histiostomatidae), and

Unguidispus sp. (Microdispidae). The 1997 and 2007 collections by Rissing and

Schaedla included Cosmoglyphus sp., a species not collected in this study; however, they are lacking Armacarus sp., the most abundant mite species (although primarily present in a single colony) found in this study. Also two other undocumented species were found in smaller numbers, Histiostoma sp., and Unguidispus sp. It is likely that mite species exhibit fluctuations in abundance and ubiquity in different years. It is also probable that the mite community may change in composition throughout the year depending on resource availability.

The curve showing the cumulative number of mite species per colony collected in

2008 can be viewed in Figure 5. The expected total number of species is estimated by the point at which the line appears to be approaching an asymptote. These data approach an asymptote at approximately 7 or 8 total species. Further sampling would be required to acquire a definitive list of associated mites and their yearly cycles; however, current data are suggestive of adequate sampling.

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Figure 5. Messor pergandei Species Accumulation Curve. Cumulative number of mite species, Sobs (Mao Tau), per number of colonies collected.

Sex and Attachment Site Specificity

Mites showed a marked preference for female hosts. For all species of mites except for Unguidispus sp., which was represented by a singleton, the majority of individuals were found on the female alates. Male M. pergandei alates die soon after mating, as is the case for most ant species, so the ability to select a female host, would be an advantage for a mite requiring resources found within the nest. That and the desiccating desert environment (allowing little time for survival on a dead host) would select for female host preference. As described in Chapter 2, mites could return to the nest by way of necrophagy by other ants (the limited resources of the desert might, in fact, increase necrophagic activity). Another point worth noting is that the 7 instances of male-riding mites occurred in 2 colonies which were male-skewed. In all cases of female- skewed nests only females had mites, thus, host sex specificity may be strongly

124 influenced by host availability rather than sex of the host for some mite species.

However, it is more likely that males are utilized in situations in which females are rare or more difficult to find. This is supported by the disparity between percentage of each sex with mites and the percentage of the total alates each sex present within the colony

(see Table 24). In both cases of phoresy on males, males were more common than females in the nest, but smaller percentages of males had phoretic mites.

Attachment site specificity is apparent in the well sampled mite species

Armacarus sp., Lemanniella sp., and Petalomium sp. Although most mites were found in several locations on the alates, the data suggest a preference for one or a couple of key locations. In highly infested ants, the mites commonly spill over to locations beyond their primary attachment sites, but when only a few mite individuals are present, they can usually be found in their preferred locations. Interestingly, this attachment site preference is maintained on the different ant sexes despite morphological differences between the sexes. For example, Lemanniella sp. prefers to ride on the underside of the ant’s head, as previously documented by Wurst (2001), in both males and females, but M. pergandei males have much smaller heads and less developed psammophores than the females.

Exactly how the mites select and distribute themselves at attachment locations is still unknown.

Alate versus Dealate Mite Abundance

Dealates appeared to have slightly lower mite loads than alates. While alates are relatively undisturbed while they are residing in the nest, dealates undergo many more obstacles which may cause mites to become lost, such as flying, mating, and digging a

125 nest. It is also possible that the mites may have fallen off during handling, transit to Ohio or during transfer to test tubes or the laboratory nest. Many insect species clean themselves of mites, so loss of mites could also be from hygienic efforts. It appears that mites arrange themselves in more secure locations (under the psammophore, between legs, underside of gaster all facing the same direction) in an effort to be more aerodynamic, protected, and less of a burden for the host, and in so doing, are more likely to reach their destinations, despite the obstacles.

Chaff Piles

Chaff piles appear to hold little mite diversity. They were very dry despite rain in the area during my visit, which is probably the main factor influencing mite numbers in the chaff pile. Chaff piles are primarily made of seeds and small pieces of plants, but seem to be held together by fungal mycelia. The Bakerdania sp. was probably feeding on fungus within the chaff pile. Procaeculus sp. and Anystidae sp. are usually predatory.

Tetranychid mites are plant feeders, and may have been deposited in the chaff by ants gathering plant materials. Mites in the family Tydeidae utilize a wide range of food sources, so no clear role can be determined for the most common of the chaff species.

Caveats

The lack of precise documentation in the cases of unobserved mites was probably due to an inability to thoroughly inspect alates while avoiding injury to them. Loss of mites may be due to accidentally counting the same mite multiple times (when on highly infested ants), difficulty in observing mites on a wriggling host, inability to recover mites from Eppendorf tubes, or loss of mites during handling prior to insertion into the tube.

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This inconsistency led to less robust results regarding attachment site specificity. One solution would require individual collections to be made at the time of the inspection. All mites from each body region would have to be removed in situ and placed in separate vials. This would, of course, be extremely time consuming and fraught with opportunity for loss of mites, but without access to resources for immediate clearing and slide making

(such as those used in Chapter 2), it may be the most reliable alternative. Another possible method might employ a kill jar or freezing of the ants prior to inspection.

Rettenmeyer (1962) used ether kill jars, but noted that many of the mites were caught in condensation and lost. Freezing may work just as well, but any jostling after death would result in the loss of many of the mites in condensation.

Unfortunately only a dozen dealates with mites were collected. These collections are very weather dependent, as mating flights are not easily predictable in this species.

No previous studies had attempted to rear dealates for mite observation, so developing an efficient method was a goal of this study. After the malfunction of the incubator, however, few of the dealates with mites remained, thereby allowing little opportunity for manipulation and observation. The lack of mite disembarkation from the hosts indicates that rearing conditions were not optimal for mite growth and development. With greater collections, it might be possible to determine what conditions elicit a developmental response in the mites.

Conclusion Summary

At least seven mite species are phoretically associated with Messor pergandei:

Armacarus sp., Lemanniella sp., Petalomium sp., Forcellinia sp., Histiostoma sp.,

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Unguidispus sp., and Cosmoglyphus sp. Most of these species appear to have preferred attachment sites for phoresy, and appear to prefer female alates rather than male alates.

Five mite species were found in low numbers inhabiting the chaff piles: Tydeidae sp.,

Procaeculus sp., Anystidae sp., Bakerdania sp., and Tetranychidae sp. The phoretic

Petalomium sp. was observed consuming fungus on a dead dealate, but the roles of the other species are still unclear.

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Messor Species Mite Family Mite Species Association Reference M. barbarus Laelapidae (Hypoaspis) laevis in nest of Berlese, 1904 Messoracaridae Messoracarus mirandus on head Silvestri, 1912 M. capitatus Hunter & Hunter, 1963; Laelapidae Myrmozercon acuminatus in nest of Vitzthum,1930; Berlese, 1904 Hunter & Hunter, 1963; Laelapidae Myrmozercon brachiatus in nest of Vitzthum,1930; Berlese, 1904 M. excursionis Laelapidae Laelaps (Hypoaspis) intermedius in nest of Karawajew, 1909 Scutacaridae Imparipes placidus in nest of Khaustov & Chydyrov, 2004 Circocyllibanidae Cillibano transversalis in nest of Karawajew, 1909 M. meridionalis Oplitidae Oplitis inopina in nest of Hull, 1923

129 129 M. structor Oplitidae Oplitis leonardiana in nest of Berlese, 1904

Oplitidae Oplitis philoctena in nest of Berlese, 1904 Gymnolaelaps myrmophilus in nest of Štorkán , 1940 Trachyuropoda magna in nest of Štorkán, 1940 Messor sp. Scutacaridae Imparipes ignotus in nest of Khaustov & Chydyrov, 2004

Table 27. Mites Associated with Messor Species.

Literature Cited

Beattie, A. J. and D.C. Culver. (1983). The nest chemistry of two seed-dispersing ant species. Oecologia, 56: 99-103.

Berlese, A. (1904). Illustrazione iconografica degli Acari mirmecofili. Redia, 1903 & 1904: 299-474.

Cahan, S., K. R. Helms, and S. W. Rissing. (1998). An abrupt transition in colony founding behavior in the ant Messor pergandei. Animal Behaviour, 55: 1583- 1594.

Colwell, R. K. (2005). EstimateS: Statistical estimation of species richness and shared species from samples. Version 7.5. User's Guide and application published at: http://purl.oclc.org/estimates.

Hull, J. E. (1923). New myrmecophilous Gamasids. Annals and Magazine of the Natural history of London, 12: 610-616.Hunter & Hunter, 1963

Hunter, P. E. and C. A. Hunter (1963). The genus Myrmonyssus with descriptions of two new species (Acarina: Laelaptidae). Acarologia, 5: 335-341.

Johnson, R. A. (2000). Seed-harvester ants (Hymenoptera: Formicidae) of North America: An overview of ecology and biogeography. Sociobiology, 36, 89-122.

Johnson, R. A. (2001). Biogeography and community structure of North American seed- harvester ants. Annual Reviews of Entomology, 46: 1-29.

Karawajew, W. (1909). Myrmecophilen aus Transkapien. Revue russe d’entomologie, Russkoe Entomologicheskoe Obozrenie 3: 227-237.

Khaustov, A. A. and P. R. Chydyrov. (2004). New species of mites of the family Scutacaridae (Acari: Heterostigmata) associated with ants (Hymenoptera, Formicidae) from Turkmenistan. Acarina, 12: 87-103.

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OConnor, B. M. (2009) Cohort Astigmatina. In G. W. Krantz & D. E. Walter (Eds.), A Manual of Acarology (3rd ed.), (pp. 565-657). Lubbock, TX: Texas Tech University Press.

OConnor, B. M. Keys to the genera of astigmatid mites. Acarology Summer Program 2008. Unpublished.

O’Dowd, D. J. and M. E. Hay. (1980). Mutualism between harvester ants and a desert ephemeral: seed escape from rodents. Ecology, 61: 531-540.

Pollock, G. B. and S. W. Rissing. (1985). Mating season and colony foundation of the seed harvester ant, Veromessor pergandei. Psyche, 92, 125-134.

Rettenmeyer, C.W. (1962). Arthropods associated with neotropical army ants with a review of the behavior of these ants (Arthropoda; Formicidae: Dorylinae). Ph.D. Dissertation. University of Kansas, Lawrence, KS. 644p.

Rissing, S. W. and R. A. Johnson. (2000). Colony founding behavior of some desert ants: geographic variation in metrosis. Psyche, 103: 95-101.

Rissing, S. W. (1986). Indirect effects of granivory by harvester ants: plant species composition and reproductive increase near ant nests. Oecologia (Berlin), 68: 231-234.

Savulkina, M. M. (1981). Systematics, ecology, and distribution of mites of the family Pygmephoridae Cross, 1965 (Acari, Trombidiformes). Entomological Review, 60: 163-180.

Silvestri, F. (1912). Contribuzioni alla conoscenza dei mirmecofili. Bollettino del laboratorio di zoologia generale e agrarian della facoltá agraria in portici, 6: 222-245.

Steinberger, Y., H. Leschner, and A. Shmida. (1991). Chaff piles of harvester ant (Messor spp.) nests in a desert ecosystem. Insectes Sociaux, 38: 241-250.

Štorkán, J. (1940). Myrmecophilní Acari z Čech. Vestnik Ceskoslovenske zoologicke spolecnosti v Praze, 8: 166-172.

Tevis, T., Jr. (1958). Interrelations between the harvester ant Veromessor pergandei (Mayr) and some desert ephemerals. Ecology, 39: 695-704.

Vitzthum, H. G. (1930). Ein Ameisengast (Acar.). Deutsche Entomologische Gesellschaft, 1: 89-94.

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Walter, D. E., E. E. Lindquist, I. M. Smith, D. R. Cook, and G. W. Krantz. (2009). Order Trombidiformes. In G. W. Krantz & D. E. Walter (Eds), A Manual of Acarology (3rd ed.), (pp. 233-420). Lubbock, TX: Texas Tech University Press.

Wheeler, J. and S. W. Rissing. (1975a). Natural history of Veromessor pergandei: I. The nest. The Pan-Pacific Entomologist, 51: 205-216.

Wheeler, J. and S. W. Rissing. (1975b). Natural history of Veromessor pergandei: II. Behavior. The Pan-Pacific Entomologist, 51: 302-314.

Wheeler, G. C., Wheeler, J. (1973). Ants of Deep Canyon. University of California, Riverside: Philip L. Boyd Deep Canyon Desert Research Center.

Wurst, E. (2001). The life cycle of Lemanniella minotauri n. sp. and the erection of the new family Lemannillidae (Acari: Astigmata). Stuttgarter Beiträge zur naturkunde Serie A (Biologie), 621: 1-34.

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Chapter 4: Synthesis of the Studies, Implications, and Future Research

This study has brought to the forefront the most abundant, yet most often overlooked of ant nest inhabitants, the mites. In Chapter 1, the reader was introduced to the literature regarding ant mites, and the most complete list of ant associated mites was provided. The compilation of literature surrounding these enigmatic guests will be a great asset to subsequent researchers in acarology and . Mites are a critical component in the ecosystem, and it is my hope that my studes reflect the complexity of the with ecological information regarding the mites found therein.

Distinctive Qualities of this Study

While Rettenmeyer’s (1962) thesis remains a classic examination of mites and other arthropods associated with army ants, conclusions of his study cannot always be applied to non legionary ants and may not adequately portray mites of North America.

Many of the mites discovered by Rettenmeyer are very unusual and rare mites requiring the erection of new families and genera found only with army ants. While they are extremely fascinating mites, many aspects of army ant associated mites cannot be reasonably applied to any system beyond that of army ants. My study differs in that mites of common genera were frequently encountered and documented and roles of many were discerned. The ants in the Ohio study (Chapter 2) span much of the northeastern United

States and many can be found throughout the US. Some such as Camponotus 133 pennsylvanicus, Tapinoma sessile, Tetramorium caespitum, and Lasius neoniger are frequently encountered in human houses as pests. Messor pergandei is the most commonly encountered ant in the study area of Casa Grande, and it also is a common ant throughout much of Arizona and the southwestern United States.

The majority of mites collected the current studies were members of the

Astigmata or Heterostigmatina. This is in clear contrast to the published literature (See

Chapter 1), in which primarily Mesostigmata have been documented. It is likely that the generally smaller size of the Astigmata and Heterostigmatina has led to their subsequent underrepresentation in the literature. Unlike most studies which focused on only a single mite genus or single ant species, this study documented all of the representative mite species in all ant colonies collected and identified them to a level which provided ecological insight.

The studies in Chapter 2 and Chapter 3 not only provided a list of mite associates, but gave possible ecological explanations for trends observed in mite species richness and prevalence. While these were not experimental studies, and therefore cannnot be extrapolated to all ants, all mites, or even all ant mites, they provide a great deal of information to act as a foundation for the explanation of the factors influencing mite species richness.

Major Conclusions of the Projects

The roles of mites within the ant nests range from parasitism (as in

Antennophorus) to apparent commensalism (numerous fungivores and ), but most of their functions remain undocumented or unclear. Laboratory rearing may be the

134 best method to not only link phoretic forms to non-phoretic instars, but also to aid in determining the roles of the mites.

In Ohio (Chapter 2), mite species diversity per colony appeared to be highly influenced by habitat and nest substrate, but habitat categories of woods/open and wood/soil may be too broad to adequately describe the fine-grained level at which mites sense their environment. Overall size of the ant colony, and host size were positively correlated with number of mite species and prevalence within the ant colony.

Data in both Chapter 2 and Chapter 3 showed a marked preference for mite dispersal using female alates rather than male alates, observations that support previous research by Khaustov & Moser (2008) and Fashing (1976). Both of my studies also showed many of the species to have very specific phoretic locations upon their hosts.

Clearly mites have sophisticated means of determining suitable hosts.

In Ohio (and this may also hold true for Arizona, though only M. pergandei was not observed), mites were usually host specific. Perhaps the most intriguing exceptions to this are due to the influence of temporary social parasitism, especially in the genus

Lasius. It appears that mites switch to the parasitic ant species once the original host species is dead. This is in contrast to the species of ants living in close proximity (such as lestobiosis), which elicited no host switching among any of their associated mites.

Implications and Suggestions for Future Research

Studying the ant mites may allow for clearer resolution of ecological questions regarding ants. Mites’ abilities to interact with their environment on a small scale, their specificity to their hosts, and their integral role in nutrient cycling make them valuable

135 organisms for ecological research. Some suggestions for future work in this area include studying: 1) the variation among colonies in mite composition and prevalence and whether these factors can be used to indicate overall health, age, and activity of the ant colony, 2) the role of the mites within the colony and potiential use as biological control agents against invasive and introduced ant species, 3) the transport of mites by way of introduced and invasive ant species and exchange of mites with native ant species, 4) the influence of overall quality of the habitat and disturbance on mite diversity within the ant colonies, 5) parasitic ant colonies and using mites to determine the original host, and 6) seasonal trends in ant mite populations and phoretic activity within a colony.

136

Literature Cited

Fashing, N. J. (1976). The evolutionary modification of dispersal in Naiadacarus arboricola Fashing, a mite restricted to waterfilled treeholes (Acarina: Acaridae). American Midland Naturalist, 95: 337-346.

Khaustov, A. A. and J. C. Moser. (2008). Two new species of mites of the genera Petalomium Cross and Caesarodispus Mahunka (Acari: Heterostigmatina: Neopygmephoridae, Microdispidae) associated with Solenopsis invicta Buren (Hymenoptera: Formicidae) from the U.S.A. International Journal of Acarology, 34: 115-121.

Rettenmeyer, C.W. (1962). Arthropods associated with neotropical army ants with a review of the behavior of these ants (Arthropoda; Formicidae: Dorylinae). Ph.D. Dissertation. University of Kansas, Lawrence, KS. 644p.

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Appendix A: Chapter 1 Supplementary Tables

138

Appendix A: Chapter 1 Supplementary Tables

PARASITIC SPECIES HOST(S) REFERENCE Aphaenogaster tennesseensis A. fulva Coovert, 2005 A. picea Coovert, 2005 A. rudis Coovert, 2005 Formica exsectoides F. fusca group Coovert, 2005 Lasius (Acanthomyops) claviger L. (L.) alienus Raczkowski, 2008 L. (.)umbratus Raczkowski, 2008 L. (A.) interjectus L. (L.) alienus Raczkowski, 2008 L. (A.) claviger Raczkowski, 2008 L. (A.) latipes1 Raczkowski, 2008 L. (Ch.) minutus1 Raczkowski, 2008 L. (A.) murphyi1 Raczkowski, 2008 L. (Ch.) umbratus Raczkowski, 2008 L. (Chthonolasius) umbratus L. (L.) alienus Coovert, 2005 L. (L.) neoniger Coovert, 2005 L. (L.) niger1 Coovert, 2005 L. (L.) pallitarsus1 Coovert, 2005 L. (Ch.) speculiventris L. (Ch.) minutus1 Coovert, 2005 L. (Ch.) umbratus Coovert pers. comm.. L. (A.) latipes1 L. (L.) alienus Raczkowski, 2008 L. (L.) crypticus1 Raczkowski, 2008 L. (A.) interjectus Raczkowski, 2008 L. (L.) neoniger Raczkowski, 2008 L. (A.) murphyi1 L. (L.) neoniger Raczkowski, 2008 L. (Ch.) minutus1 L. (L.) alienus1 Coovert, 2005 L. (L.) pallitarsus1 Coovert, 2005 L. (Ch.) subumbratus1 L. (L.) pallitarsus1 Coovert, 2005 Table 28: Temporary Social Parasitism Interactions among Ohio Ant Species. Data presented are compiled from literature in which mixed colonies were observed or laboratory trials indicated acceptance of the parasitic species. The only social parasite Formica species collected was F. exsectoides. 1Designates a species which was not collected in Chapter 2’s study, but has been documented in Ohio.

139

HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Aenictequidae Aenicteques chapmani Jacot, 1939 Aenictus gracilis in nest of Jacot, 1939 Antennophoridae Antennophorus uhlmanni Haller, 1877 Lasius niger on ants Haller, 1877 Antennophoridae Antennophorus barbatus Wasmann Labidus praedator on ants Wasmann, 1902 Wiśniewski & Hirschmann, Antennophoridae Antennophorus boveni Wiśniewski & Hirschmann, 1992 Lasius flavus phoretic on ants 1992 Antennophorus Antennophoridae donisthorpei Wheeler, 1910 Lasius claviger under head of ant Wheeler, 1910b Antennophorus Antennophoridae donisthorpei Wheeler, 1910 Lasius flavus under head of ant Wheeler, 1910b Antennophorus Antennophoridae donisthorpei Wheeler, 1910 Lasius interjectus under head of ant Wheeler, 1910b Antennophorus Antennophoridae donisthorpei Wheeler, 1910 Lasius latipes under head of ant Wheeler, 1910b Antennophorus Antennophoridae donisthorpei Wheeler, 1910 Lasius nearcticus under head of ant Wheeler, 1910b

140 Antennophorus Pachycondyla Antennophoridae emarginatus Banks, 1916 lutea in nest of Banks, 1916 Antennophoridae Antennophorus foreli Wasm. Lasius alienus in nest of Berlese, 1904 Antennophoridae Antennophorus foreli Wasmann Lasius alienus - Karawajew, 1906 Antennophoridae Antennophorus foreli Wasmann, 1902 Lasius alienus on ants Wasmann, 1902 Antennophoridae Antennophorus foreli Wasmann, 1902 Lasius flavus in nest of Štorkán, 1940 Antennophoridae Antennophorus foreli Wasm. Lasius niger in nest of Berlese, 1904 Antennophoridae Antennophorus foreli Wasm. Lasius niger under head or side gaster Donisthorpe, 1927 Antennophoridae Antennophorus foreli Wasmann Lasius niger - Karawajew, 1906 Antennophoridae Antennophorus foreli Wasmann, 1902 Lasius niger on ants Wasmann, 1902 Wiśniewski & Hirschmann, Antennophoridae Antennophorus goesswaldi Wiśniewski & Hirschmann, 1992 Lasius flavus phoretic on ants 1992 Antennophoridae Antennophorus grandis Berlese Lasius alienus - Karawajew, 1906 Antennophoridae Antennophorus grandis (Berlese, 1903) Lasius flavus cleptoparasite Franks et al., 1991 Antennophoridae Antennophorus grandis Berlese Lasius flavus - Karawajew, 1906 Antennophoridae Antennophorus grandis Berl. Lasius fuliginosus in nest of Berlese, 1904 Table 29. Mesostigmata and their Associated ants from Literature. Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Antennophoridae Antennophorus grandis Berlese Lasius fuliginosus - Karawajew, 1906 Antennophoridae Antennophorus grandis Berlese Lasius fuliginosus under head or side gaster Donisthorpe, 1927 Antennophoridae Antennophorus grandis Berlese, 1904 Lasius niger in nest of Štorkán, 1940 Wiśniewski & Wiśniewski & Antennophoridae Antennophorus pavani Hirschmann, 1992 Lasius sp. phoretic on ants Hirschmann, 1992 Antennophoridae Antennophorus pubescens Wasmann - Karawajew, 1906 Antennophoridae Antennophorus pubescens Hall. Lasius flavus in nest of Wasmann, 1899 Antennophoridae Antennophorus pubescens Wasm. Lasius flavus under head or side gaster Donisthorpe, 1927 Antennophoridae Antennophorus pubescens Wasmann Lasius flavus - Karawajew, 1906 Antennophoridae Antennophorus pubescens Wasmann Lasius flavus in nest of Berlese, 1904 Antennophoridae Antennophorus pubescens Wasmann Lasius flavus on ants Wasmann, 1902 Antennophoridae Antennophorus pubescens Wasmann Lasius fuliginosus - Karawajew, 1906

141 Antennophoridae Antennophorus pubescens Wasmann Lasius mixtus - Karawajew, 1906 Antennophoridae Antennophorus pubescens Wasmann Lasius niger - Karawajew, 1906

Antennophoridae Antennophorus pubescens Wasmann, 1897 Lasius umbratus in nest of Lehtinen, 1987 Antennophoridae Antennophorus raffrayi Raffray & Brauns? Anoplolepis custodiens on ants Wasmann, 1902 Antennophoridae Antennophorus sp. n/a Lasius brunneus under head or side gaster Donisthorpe, 1927 Antennophoridae Antennophorus uhlmanni Hall in nest of Berlese, 1904 Antennophoridae Antennophorus uhlmanni Hall. Lasius alienus on ants Wasmann, 1902 Antennophoridae Antennophorus uhlmanni Hall Lasius fuliginosus - Karawajew, 1906 Antennophoridae Antennophorus uhlmanni Hall. Lasius fuliginosus on ants Wasmann, 1902 Antennophoridae Antennophorus uhlmanni Hall. Lasius mixtus on ants Wasmann, 1902 Antennophoridae Antennophorus uhlmanni Haller Lasius mixtus under head Janet, 1902 Antennophoridae Antennophorus uhlmanni Haller, 1877 Lasius mixtus under head, on gaster Janet, 1897a Antennophoridae Antennophorus uhlmanni Hall Lasius niger in nest of Berlese, 1904 Antennophoridae Antennophorus uhlmanni Hall. Lasius niger in nest of Wasmann, 1899 Antennophoridae Antennophorus uhlmanni Hall. Lasius niger on ants Wasmann, 1902 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Antennophoridae Antennophorus uhlmanni Haller, 1877 Lasius niger under head, on gaster Janet, 1897a Antennophoridae Antennophorus uhlmanni n/a Lasius umbratus under head Mann, 1913 Antennophoridae Antennophorus uhlmanni Hall Lasius umbratus in nest of Berlese, 1904 Antennophoridae Antennophorus uhlmanni Hall. Lasius umbratus on ants Wasmann, 1902 Antennophoridae Antennophorus uhlmanni Hall. Lasius umbratus under head or side gaster Donisthorpe, 1927 Antennophoridae Antennophorus uhlmanni Haller, 1877 Lasius umbratus under head, on gaster Janet, 1897a Antennophoridae Antennophorus wasmanni Wheeler Lasius flavus on ants Park, 1932 Antennophoridae Antennophorus wasmanni Wheeler, 1910 Lasius umbratus in nest of Wheeler, 1910b Antennophoridae Antennophorus wheeleri Wasmann Lasius flavus on ants Wasmann, 1902 Antennophoridae Echinomegistus wheeleri (Wasm.) Lasius flavus in nest of Berlese, 1904 Antennophoridae Paramegistus australis Banks, 1916 Polyrhachis hexacantha in nest of Banks, 1916 Antennophoridae Ptocharus daveyii Silvestri Camponotus aeneopilosus in nest of Banks, 1916

142 Antennophoridae Ptocharus daveyii Silvestri Papyrius nitidus in nest of Banks, 1916 Ascidae Lasioseius ometes (Oudemans, 1903) Lasius fuliginosus in nest of Rack, 1968 Ascidae hypudaei (Oudemans, 1902) Lasius fuliginosus in nest of Rack, 1968 Ascidae Rettenmeyerius agnesae Elzinga, 1998 Labidus praedator in emmigration column Elzinga, 1998 Ascidae Rettenmeyerius agnesae Elzinga, 1998 Nomamyrmex esenbeckii in emmigration column Elzinga, 1998 base of mandibles or Ascidae Rettenmeyerius carli Elzinga, 1998 labrum, feed on larvae Elzinga, 1998 base of mandibles or Ascidae Rettenmeyerius carli Elzinga, 1998 labrum, feed on larvae Elzinga, 1998 Ascidae Rettenmeyerius marianae Elzinga, 1998 Eciton dulcium in emmigration column Elzinga, 1998 Ascidae Rettenmeyerius marianae Elzinga, 1998 in emmigration column Elzinga, 1998 attached to propodial hairs Ascidae Rettenmeyerius plaumanni Elzinga, 1998 Eciton quadriglume of males Elzinga, 1998 Ascidae Rettenmeyerius schneirlai Elzinga, 1998 Labidus praedator in emmigration column Elzinga, 1998 Cercomegistina Seiodes histricinus Berl. Lasius mixtus in nest of Wasmann, 1899 Cercomegistina Seiodes histricinus Berl. Lasius umbratus in nest of Wasmann, 1899 Circocyllibanidae comata Leon. Lasius flavus on larvae Wasmann, 1899 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Circocyllibanidae Cilliba comata Leon Lasius niger on larvae Wasmann, 1899 Circocyllibanidae Cilliba pandata Mich. Lasius alienus on worker Wasmann, 1899 on worker, or Circocyllibanidae Cilliba pandata Mich. Lasius niger near larvae Wasmann, 1899 Circocyllibanidae Cilliba pandata Mich. Myrmica scabrinodis in nest of Wasmann, 1899 Circocyllibanidae Cillibano comata (Leon.) Camponotus aethiops in nest of Berlese, 1904 Circocyllibanidae Cillibano comata (Leon.) Lasius flavus in nest of Berlese, 1904 Circocyllibanidae Cillibano comata (Leon.) Lasius niger in nest of Berlese, 1904 Circocyllibanidae Cillibano comata (Leon.) Lasius mixtus in nest of Berlese, 1904 Circocyllibanidae Cillibano comata (Leon.) Myrmica scabrinodis in nest of Berlese, 1904 Circocyllibanidae Circocylliba brachychaeta Elzinga & Rettenmeyer, 1974 Eciton quadriglume phoretic on ants Elzinga & Rettenmeyer, 1974 Circocyllibanidae Circocylliba camerata Elzinga & Rettenmeyer, 1974 Eciton burchellii phoretic on ants Elzinga & Rettenmeyer, 1974 Circocyllibanidae Circocylliba camerata Sellnick, 1926 Eciton quadriglume phoretic on ants Sellnick, 1926

143 Circocyllibanidae Circocylliba crinita Elzinga & Rettenmeyer, 1974 Eciton dulcium crassinode phoretic on ants Elzinga & Rettenmeyer, 1974

Circocyllibanidae Circocylliba crinita Elzinga & Rettenmeyer, 1974 Eciton vagans phoretic on ants Elzinga & Rettenmeyer, 1974 Neivamyrmex pilosus Circocyllibanidae Circocylliba crinita Elzinga & Rettenmeyer, 1974 mexicanus phoretic on ants Elzinga & Rettenmeyer, 1974 Circocyllibanidae Circocylliba ecitonis Elzinga & Rettenmeyer, 1974 Eciton burchellii foreli phoretic on ants Elzinga & Rettenmeyer, 1974 Circocyllibanidae Circocylliba ecitonis Elzinga & Rettenmeyer, 1974 Eciton dulcium phoretic on ants Elzinga & Rettenmeyer, 1974 Circocyllibanidae Circocylliba ecitonis Elzinga & Rettenmeyer, 1974 Eciton hamatum phoretic on ants Elzinga & Rettenmeyer, 1974 Circocyllibanidae Circocylliba ecitonis Elzinga & Rettenmeyer, 1974 Eciton lucanoides phoretic on ants Elzinga & Rettenmeyer, 1974 Circocyllibanidae Circocylliba ecitonis Elzinga & Rettenmeyer, 1974 Eciton vagans phoretic on ants Elzinga & Rettenmeyer, 1974 Neivamyrmex pilosus Circocyllibanidae Circocylliba ecitonis Elzinga & Rettenmeyer, 1974 mexicanus phoretic on ants Elzinga & Rettenmeyer, 1974 Circocyllibanidae Circocylliba ecuadorensis Elzinga & Rettenmeyer, 1974 Eciton dulcium phoretic on ants Elzinga & Rettenmeyer, 1974 Circocyllibanidae Circocylliba ecuadorensis Elzinga & Rettenmeyer, 1974 Eciton lucanoides phoretic on ants Elzinga & Rettenmeyer, 1974 Circocyllibanidae Circocylliba ecuadorensis Elzinga & Rettenmeyer, 1974 Eciton vagans phoretic on ants Elzinga & Rettenmeyer, 1974 Neivamyrmex pilosus Circocyllibanidae Circocylliba ecuadorensis Elzinga & Rettenmeyer, 1974 mexicanus phoretic on ants Elzinga & Rettenmeyer, 1974 Circocyllibanidae Circocylliba minuta Elzinga & Rettenmeyer, 1974 Labidus praedator phoretic on ants Elzinga & Rettenmeyer, 1974 Circocyllibanidae Circocylliba oligochaeta Elzinga & Rettenmeyer, 1974 Eciton burchellii foreli phoretic on ants Elzinga & Rettenmeyer, 1974 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Elzinga & Circocyllibanidae Circocylliba oligochaeta Elzinga & Rettenmeyer, 1974 Eciton hamatum phoretic on ants Rettenmeyer, 1974 Elzinga & Circocyllibanidae Circocylliba oligochaeta Elzinga & Rettenmeyer, 1974 Eciton lucanoides phoretic on ants Rettenmeyer, 1974 Neivamyrmex pilosus Elzinga & Circocyllibanidae Circocylliba oligochaeta Elzinga & Rettenmeyer, 1974 mexicanus phoretic on ants Rettenmeyer, 1974 Elzinga & Circocyllibanidae Circocylliba weberi Elzinga & Rettenmeyer, 1974 Eciton burchellii cupiens phoretic on ants Rettenmeyer, 1974 Circocyllibanidae Cylliba pandata n/a Lasius niger on ants Wasmann, 1902 Circocyllibanidae Trichocyllaba napoensis Elzinga, 1981 Labidus coecus phoretic on ants Elzinga, 1981 Circocyllibanidae Trichocyllaba suctorpoda Elzinga, 1981 Labidus praedator phoretic on ants Elzinga, 1981 Circocyllibanidae Trichocylliba ablesi Hirschmann, 1973 Lasius interjectus in nest of Hirschmann, 1973 Circocyllibanidae Trichocylliba ablesi Hirschmann, 1973 Lasius umbratus in nest of Hirschmann, 1973 Circocyllibanidae Trichocylliba agnesae Elzinga, 1995 Labidus praedator phoretic on ants Elzinga, 1995

144 Circocyllibanidae Trichocylliba chiapenis Elzinga, 1981 Eciton hamatum phoretic on ants Elzinga, 1981 Circocyllibanidae Trichocylliba galea Elzinga, 1995 Labidus praedator phoretic on ants Elzinga, 1995 Circocyllibanidae Trichocylliba morosa Elzinga, 1995 Cheliomyrmex morosus phoretic on ants Elzinga, 1995 Circocyllibanidae Trichocylliba neili Elzinga, 1995 Labidus coecus phoretic on ants Elzinga, 1995 Circocyllibanidae Trichocylliba praedator Elzinga, 1981 Labidus praedator phoretic on ants Elzinga, 1981 Circocyllibanidae Trichocylliba schneirlai Elzinga, 1981 Labidus coecus phoretic on ants Elzinga, 1981 Circocyllibanidae Trichocylliba tumba Elzinga, 1995 Eciton burchellii refuse deposits Elzinga, 1995 Circocyllibanidae Trichocylliba watkinsi Elzinga, 1981 Neivamyrmex nigrescens phoretic on ants Elzinga, 1981 Circocyllibanidae Trichocylliba watkinsi Elzinga, 1981 Neivamyrmex opacithorax phoretic on ants Elzinga, 1981 Coxequesomidae Coxequesoma collegianorum Sellnick, 1926 Eciton quadriglume phoretic on ants Sellnick, 1926 Coxequesomidae Coxequesoma gignodissidens Elzinga, 1982a Neivamyrmex sp. coxae on ant Elzinga, 1982a Coxequesomidae Coxequesoma hermanni Elzinga, 1982a Eciton hamatum coxae on ant Elzinga, 1982a Coxequesomidae Coxequesoma labidocoxata Elzinga, 1982a Labidus praedator coxae on ant Elzinga, 1982a Coxequesomidae Coxequesoma panamaensis Elzinga, 1982a Labidus praedator coxae on ant Elzinga, 1982a Coxequesomidae Coxequesoma umbocauda Elzinga, 1982a Eciton hamatum coxae on ant Elzinga, 1982a Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Coxequesomidae Coxequesoma umbocauda Elzinga, 1982a Neivamyrmex pilosus mexicanus coxae on ant Elzinga, 1982a Coxequesomidae Habeogula cauda Elzinga, 1989 Labidus praedator ventral ant head Elzinga, 1989 Diplogyniidae Brachysternopsis flechtmanni Hunter, 1993 Atta capiguara on ant Hunter, 1993 Gamasid Myrmeciphis crawleianus Hull, 1923 Bothriomyrmex sp. in nest of Hull, 1923 Gamasidae undetermined sp. n/a Solenopsis richteri attached to worker Collins & Markin, 1971 Klinckowstroemiidae Ptochacarus banksi Womersley, 1958 Camponotus aeneopilosus in nest of Womersley, 1958 Klinckowstroemiidae Ptochacarus daveyi Womersley, 1958 Camponotus aethiops in nest of Womersley, 1958 Klinckowstroemiidae Ptochacarus daveyi Womersley, 1958 Papyrius nitidus in nest of Womersley, 1958 Klinckowstroemiidae Ptochacarus silvestrii Womersley, 1958 unknown (ants) in nest of Womersley, 1958 Laelapidae sp. n/a Solenopsis invicta in nest of Hermann et al., 1970 Laelapidae Cosmolaelaps cuneifer (Michael, 1891) Camponotus aethiops in nest of Štorkán, 1940 Laelapidae Cosmolaelaps cuneifer (Michael, 1891) Camponotus ligniperdus in nest of Štorkán, 1940

145 Laelapidae Cosmolaelaps cuneifer (Michael, 1891) Formica fusca in nest of Štorkán, 1940 Laelapidae Cosmolaelaps cuneifer (Michael, 1891) Formica rufibarbis in nest of Štorkán, 1940 Laelapidae Cosmolaelaps cuneifer (Michael, 1891) in nest of Štorkán, 1940 Laelapidae Cosmolaelaps cuneifer (Michael, 1891) Lasius flavus in nest of Štorkán, 1940 Laelapidae Cosmolaelaps cuneifer (Michael, 1891) Lasius fuliginosus in nest of Štorkán, 1940 Laelapidae Cosmolaelaps cuneifer (Michael, 1891) Lasius umbratus in nest of Štorkán, 1940 Laelapidae Cosmolaelaps cuneifer (Michael, 1891) Tapinoma erraticum in nest of Štorkán, 1940 Laelapidae Cosmolaelaps thysanifer Zeman, 1982 Formica fusca in nest of Zeman, 1982 Laelapidae Cosmolaelaps vacuus (Michael, 1891) Camponotus ligniperdus in nest of Štorkán, 1940 Laelapidae Cosmolaelaps vacuus (Michael, 1891) Formica fusca in nest of Štorkán, 1940 Laelapidae Cosmolaelaps vacuus (Michael, 1891) Formica sanguinea in nest of Štorkán, 1940 Laelapidae Cosmolaelaps vacuus (Michael, 1891) Lasius alienus in nest of Štorkán, 1940 Laelapidae Cosmolaelaps vacuus (Michael, 1891) Lasius flavus in nest of Štorkán, 1940 Laelapidae Cosmolaelaps vacuus (Michael, 1891) Lasius niger in nest of Štorkán, 1940 Laelapidae Cosmolaelaps vacuus (Michael, 1891) Tetramorium caespitum in nest of Štorkán, 1940 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Laelapidae sp. n/a Myrmecocystus mendax with ant larvae Conway, 2003 Laelapidae Gymnolaelaps shealsi Hunter & Costa, 1971 Solenopsis invicta in nest of Hunter & Costa, 1971 Laelapidae Gymnolaelaps sp. n/a Myrmecocystus mendax near larvae Conway, 2003 Laelapidae Holostaspis sp. n/a Solenopsis xyloni in nest of Neece & Bartell, 1982 Laelapidae Holostaspis terreus Canest. Et Fanz in nest of Moniez, 1892 Laelapidae Hypoaspis cardiophorus Berlese, 1916 lundii in nest of Berlese, 1916 Laelapidae Hypoaspis caudicomatus Berlese, 1916 Acromyrmex lundii in nest of Berlese, 1916 Laelapidae Hypoaspis tuberculata Mašán, 1992 Lasius flavus in nest of Mašán, 1992 Laelapidae Hypoaspis australis Hull, 1923 Camponotus chalceus in nest of Hull, 1923 Laelapidae Hypoaspis canestrinii Berl., Costa Messor structor in nest of Hunter, 1967 Laelapidae Hypoaspis deinos Zeman, 1982 Lasius flavus in nest of Zeman, 1982 Laelapidae Hypoaspis phialiger Berlese, 1916 Acromyrmex lundii in nest of Berlese, 1916

146 Laelapidae Hypoaspis sp. n/a Myrmecia dispar in nest of Gray, B., 1974 Laelapidae Hypoaspis tripodiger Berlese, 1916 Acromyrmex lundii in nest of Berlese, 1916 Laelapidae Iphis equitans Mich Aphaenogaster testaceopilosa in nests of Wasmann, 1902 Laelapidae Iphis equitans Mich Myrmica scabrinodis in nests of Wasmann, 1902 Laelapidae Iphis equitans Mich Tetramorium caespitum on larvae Wasmann, 1902 Laelapidae Laelaspis bakeri Hunter & Davis, 1962 Myrmica fracticornis in nest of Hunter & Davis, 1962 Laelapidae Laelaps acutus Mich. Camponotus herculeanus in nest of Berlese, 1904 Laelapidae Laelaps canestrinii Berl. Tetramorium caespitum in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Aphaenogaster testaceopilosa in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Camponotus ligniperdus in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Solenopsis fugax in nest of Berlese, 1904 Laelapidae Laelaps elegantulus Berl. Tapinoma erraticum in nest of Berlese, 1904 Laelapidae Laelaps equitans Mich. Aphaenogaster testaceopilosa in nest of Berlese, 1904 Laelapidae Laelaps equitans Mich. Tetramorium caespitum in nest of Berlese, 1904 Laelaps equitans var. Laelapidae longitarsus Mich. Tetramorium caespitum in nest of Berlese, 1904 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Laelapidae Laelaps finitimus Berl. Tapinoma erraticum in nest of Berlese, 1904 Laelapidae Laelaps humeratus Berlese, 1904 Tetramorium caespitum in nest of Berlese, 1904 Laelapidae Laelaps laevis Mich. Formica subpilosa in nest of Karawajew, 1909 Laelapidae Laelaps myrmecophilus Berl. Lasius flavus in nest of Berlese, 1904 Laelapidae Laelaps myrmophilus Mich. Aphaenogaster testaceopilosa in nest of Berlese, 1904 Laelapidae Laelaps myrmophilus Mich. Lasius niger in nest of Berlese, 1904 Laelapidae Laelaps oophilus n/a Lasius flavus feeds on eggs Mann, 1913 Laelapidae Laelaps ornatus Berl. Solenopsis fugax in nest of Berlese, 1904 Laelapidae Laelaps acuta Mich. Camponotus herculeanus in nest of Moniez, 1892 Laelapidae Laelaps acuta Michael, 1891 Camponotus herculeanus in nest of Michael, 1892 Donisthorpe, Laelapidae Laelaps acutus Mich. Myrmica scabrinodis in nest of 1927 Laelapidae Laelaps ahngeri Karawajew, 1909 pygmaea in nest of Karawajew, 1909

147 Laelapidae Laelaps canstrinii Berl. Tetramorium caespitum in nest of Wasmann, 1899

Laelapidae Laelaps claviger Berl. in nest of Moniez, 1894 Laelapidae Laelaps claviger Berl. Formica exsecta in nest of Moniez, 1894 Laelapidae Laelaps claviger Berlese Lasius mixtus in nest of Janet, 1902 Laelapidae Laelaps comes Moniez, 1894 Formica omnivora in nest of Moniez, 1894 Laelapidae Laelaps complanatus Moniez, 1892 Formica fusca in nest of Moniez, 1892 Laelapidae Laelaps cuneifer Mich. Aphaenogaster picea in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Camponotus aethiops in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Camponotus herculeanus in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Camponotus herculeanus in nest of Moniez, 1892 Laelapidae Laelaps cuneifer Mich. Camponotus herculeanus in nest of Moniez, 1892 Laelapidae Laelaps cuneifer Michael, 1891 Camponotus herculeanus in nest of Michael, 1891 Laelapidae Laelaps cuneifer Mich. Formica exsecta in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Formica exsecta in nest of Wasmann, 1899 Donisthorpe, Laelapidae Laelaps cuneifer Mich. Formica exsecta scavenger on dead ants 1927 Laelapidae Laelaps cuneifer Mich. Formica fusca in nest of Berlese, 1904 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Laelapidae Laelaps cuneifer Mich. Formica fusca in nest of Wasmann, 1899 Laelapidae Laelaps cuneifer Mich. Formica fusca scavenger on dead ants Donisthorpe, 1927 Laelapidae Laelaps cuneifer Mich. in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Formica rufa in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Formica rufa in nest of Wasmann, 1899 Laelapidae Laelaps cuneifer Mich. Formica rufa scavenger on dead ants Donisthorpe, 1927 Laelapidae Laelaps cuneifer Mich. Formica rufibarbis in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Formica rufibarbis in nest of Wasmann, 1899 Laelapidae Laelaps cuneifer Mich. Formica sanguinea in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Formica sanguinea in nest of Wasmann, 1899 Laelapidae Laelaps cuneifer Mich. Formica sanguinea scavenger on dead ants Donisthorpe, 1927 Laelapidae Laelaps cuneifer Mich. Lasius alienus in nest of Berlese, 1904

148 Laelapidae Laelaps cuneifer Mich. Lasius brunneus scavenger on dead ants Donisthorpe, 1927 Laelapidae Laelaps cuneifer Mich. Lasius fuliginosus in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Lasius fuliginosus in nest of Moniez, 1892 Laelapidae Laelaps cuneifer Mich. Lasius fuliginosus in nest of Wasmann, 1899 Laelapidae Laelaps cuneifer Mich. Lasius fuliginosus scavenger on dead ants Donisthorpe, 1927 Laelapidae Laelaps cuneifer Mich. Lasius mixtus scavenger on dead ants Donisthorpe, 1927 Laelapidae Laelaps cuneifer Mich. Lasius niger in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Lasius umbratus in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Lasius umbratus scavenger on dead ants Donisthorpe, 1927 Laelapidae Laelaps cuneifer Mich. Myrmica scabrinodis in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Pheidole pallidula in nest of Berlese, 1904 Laelapidae Laelaps cuneifer Mich. Solenopsis fugax in nest of Wasmann, 1899 Laelapidae Laelaps cuneifer Mich. unknown (ants) dead ants Donisthorpe, 1927 Laelapidae Laelaps dubitatus Hunter, 1964 Aphaenogaster sp in nest of Hunter, 1964 Laelapidae Laelaps equitans Mich. Myrmica scabrinodis in nest of Berlese, 1904 Laelapidae Laelaps equitans Mich. Tetramorium caespitum rides on host Donisthorpe, 1927 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Laelapidae Laelaps equitans Michael, 1891 Tetramorium caespitum in nest of Michael, 1891 Laelapidae Laelaps flexuosa Mich. Camponotus herculeanus in nest of Moniez, 1892 Laelapidae Laelaps flexuosa Mich. Camponotus herculeanus in nest of Moniez, 1892 Laelapidae Laelaps flexuosa Michael, 1891 Camponotus herculeanus in nest of Michael, 1891 Laelapidae Laelaps flexuosa Mich. Formica sanguinea artificial nest Moniez, 1892 Laelapidae Laelaps flexuosus Mich. Camponotus herculeanus in nest of Wasmann, 1899 Laelapidae Laelaps flexuosus Mich. Formica sanguinea in nest of Wasmann, 1899 Laelapidae Laelaps humeratus Berl. Tetramorium caespitum in nest of Donisthorpe, 1927 Laelapidae Laelaps intermedius Karawajew, 1909 Messor excursionis in nest of Karawajew, 1909 Laelapidae Laelaps karawaiewi Berlese, 1904 Formica pratensis in nest of Berlese, 1904 Laelapidae Laelaps laevis Mich. Formica pratensis in nest of Berlese, 1904 Laelapidae Laelaps laevis Mich. Formica pratensis in nest of Wasmann, 1899

149 Laelapidae Laelaps laevis Mich. Formica sanguinea in nest of Berlese, 1904 Laelapidae Laelaps laevis Mich. in nest of Berlese, 1904 Laelapidae Laelaps laevis Mich. Tetramorium caespitum in nest of Donisthorpe, 1927 Laelapidae Laelaps laevis Mich. unknown (ants) in nest of Moniez, 1892 Laelapidae Laelaps laevis Michael, 1891 unknown (ants) in nest of Michael, 1891 Laelapidae Laelaps montanus Berl. Formica fusca in nest of Donisthorpe, 1927 Laelapidae Laelaps montanus Berlese, 1904 Formica fusca in nest of Berlese, 1904 Laelapidae Laelaps montanus Berl. Formica rufa in nest of Donisthorpe, 1927 Laelapidae Laelaps montanus Berl. Lasius niger in nest of Donisthorpe, 1927 Laelapidae Laelaps montanus Berl. Myrmica spp. in nest of Donisthorpe, 1927 Laelapidae Laelaps myrmecophilus Berl. Aphaenogaster barbara in nest of Moniez, 1894 Laelapidae Laelaps myrmecophilus Berl. Formica cunicularia in nest of Moniez, 1894 Laelapidae Laelaps myrmecophilus Berl. Formica cunicularia in nest of Donisthorpe, 1927 Laelapidae Laelaps myrmecophilus Berl. Formica cunicularia predator Donisthorpe, 1927 Laelapidae Laelaps myrmecophilus Berl. Formica fusca in nest of Wasmann, 1899 Laelapidae Laelaps myrmecophilus Berl. Formica rufa in nest of Berlese, 1904 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Laelapidae Laelaps myrmecophilus Berl. Formica rufa in nest of Moniez, 1894 Laelapidae Laelaps myrmecophilus Berl. Formica rufa in nest of Wasmann, 1899 Laelapidae Laelaps myrmecophilus Berl. Formica rufibarbis in nest of Berlese, 1904 Laelapidae Laelaps myrmecophilus Berl. Formica rufibarbis in nest of Wasmann, 1899 Laelapidae Laelaps myrmecophilus n/a Formica sanguinea on ants and Wasmann, 1902 Laelapidae Laelaps myrmecophilus Berl. Formica sanguinea in nest of Wasmann, 1899 Laelapidae Laelaps myrmecophilus Berl. Formica sanguinea in nest of Donisthorpe, 1927 Laelapidae Laelaps myrmecophilus Berlese Formica sanguinea in nest of Wasmann, 1897 Laelapidae Laelaps myrmecophilus Berl. Lasius flavus in nest of Wasmann, 1899 Laelapidae Laelaps myrmecophilus Berl. Lasius niger in nest of Wasmann, 1899 Laelapidae Laelaps myrmecophilus Berl. Lasius niger in nest of Donisthorpe, 1927 Laelapidae Laelaps myrmecophilus Berl. Myrmica rubra in nest of Donisthorpe, 1927

150 Laelapidae Laelaps myrmecophilus Berl. Myrmica ruginodis in nest of Donisthorpe, 1927 Laelapidae Laelaps myrmecophilus Berl. Myrmica scabrinodis in nest of Berlese, 1904 Laelapidae Laelaps myrmecophilus Berl. Myrmica scabrinodis in nest of Wasmann, 1899 Laelapidae Laelaps myrmophila Mich. Aphaenogaster testaceopilosa in nest of Moniez, 1892 Laelapidae Laelaps myrmophila Michael, 1891 Aphaenogaster testaceopilosa in nest of Michael, 1891 Laelapidae Laelaps myrmophilus Mich. Aphaenogaster testaceopilosa in nest of Berlese, 1904 Laelapidae Laelaps myrmophilus Mich. Formica fusca in nest of Berlese, 1904 Laelapidae Laelaps myrmophilus Mich. Lasius fuliginosus in nest of Wasmann, 1899 Laelapidae Laelaps myrmophilus Mich. Myrmica scabrinodis in nest of Berlese, 1904 Laelapidae Laelaps myrmophilus Mich. Pheidole pallidula in nest of Berlese, 1904 Laelapidae Laelaps myrmophylus Mich. Formica fusca in nest of Donisthorpe, 1927 Laelapidae Laelaps myrmophylus Mich. Lasius flavus in nest of Donisthorpe, 1927 Laelapidae Laelaps myrmophylus Mich. Lasius mixtus in nest of Donisthorpe, 1927 Laelapidae Laelaps myrmophylus Mich. Tetramorium caespitum in nest of Donisthorpe, 1927 Laelapidae Laelaps oophilus Wasm. Formica fusca feeds on eggs Donisthorpe, 1927 Laelapidae Laelaps oophilus Wasm. Formica pratensis feeds on eggs Donisthorpe, 1927 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Laelapidae Laelaps oophilus Wasm. Formica pratensis in nest of Berlese, 1904 Laelapidae Laelaps oophilus Wasm. Formica rufa feeds on eggs Donisthorpe, 1927 Laelapidae Laelaps oophilus Wasm. Formica rufa in nest of Berlese, 1904 Laelapidae Laelaps oophilus Wasm. Formica rufa on eggs Wasmann, 1899 Laelapidae Laelaps oophilus Wasm. Formica rufa on larvae Wasmann, 1902 Laelapidae Laelaps oophilus Mon. Formica rufibarbis on eggs Wasmann, 1897 Laelapidae Laelaps oophilus Wasm. Formica rufibarbis in nest of Berlese, 1904 Laelapidae Laelaps oophilus Wasm. Formica rufibarbis on eggs Wasmann, 1899 Laelapidae Laelaps oophilus Wasm. Formica rufibarbis on larvae Wasmann, 1902 Laelapidae Laelaps oophilus Mon. Formica sanguinea on eggs Wasmann, 1897 Laelapidae Laelaps oophilus Wasm. Formica sanguinea feeds on eggs Donisthorpe, 1927 Laelapidae Laelaps oophilus Wasm. Formica sanguinea in nest of Berlese, 1904

151 Laelapidae Laelaps oophilus Wasm. Formica sanguinea on eggs Wasmann, 1899 Laelapidae Laelaps oophilus Wasm. Formica sanguinea on larvae Wasmann, 1902 Laelapidae Laelaps oophilus Wasm. Tetramorium caespitum on larvae Wasmann, 1902 Laelapidae Laelaps ovalis Moniez Formica sanguinea in nest of Wasmann, 1899 Laelapidae Laelaps ovalis Moniez, 1894 Formica sanguinea in nest of Moniez, 1894 Laelapidae Laelaps parvulus Berlese, 1904 Formica fusca in nest of Berlese, 1904 Laelapidae Laelaps similis Moniez, 1894 Formica fusca in nest of Moniez, 1894 Laelapidae Laelaps styliferus Halbert. Lasius flavus in nest of Donisthorpe, 1927 Laelapidae Laelaps vacua Mich. Camponotus herculeanus in nest of Moniez, 1892 Laelapidae Laelaps vacua Michael, 1891 Camponotus herculeanus in nest of Michael, 1891 Laelapidae Laelaps vacuus Mich. Camponotus herculeanus in nest of Berlese, 1904 Laelapidae Laelaps vacuus Mich. Lasius flavus in nest of Donisthorpe, 1927 Laelapidae Laelaps vacuus Mich. Lasius niger in nest of Donisthorpe, 1927 Laelapidae Laelaps vacuus Mich. Myrmica rubra in nest of Berlese, 1904 Laelapidae Laelaps vacuus Mich. Myrmica scabrinodis in nest of Donisthorpe, 1927 Laelapidae Laelaspis brevichelis Hunter, 1964 Crematogaster lineolata in nest of Hunter, 1964 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Laelapidae Laelaspis brevipilis Hunter, 1961 Linepithema humile phoretic on Hunter, 1961 Laelapidae Laelaspis equitans (Michael) Tetramorium caespitum phoretic on Hunter, 1961 Laelapidae Laelaspis equitans var. longitarsa (Berlese) Tetramorium caespitum phoretic on Hunter, 1961 Laelapidae Laelaspis finitimus (Berlese, 1903) Tapinoma erraticum in nest of Hunter, 1961 Laelapidae Laelaspis humeratus (Berlese, 1904) unknown (ants) in nest of Hunter, 1961 Laelapidae Laelaspis ovisugus (Berlese, 1903) unknown (ants) in nest of Hunter, 1961 Laelapidae Laelaspis pauli Hunter & Davis, 1962 Crematogaster sp. in nest of Hunter & Davis, 1962 Laelapidae Laelaspis piloscutuli Hunter, 1961 Eciton burchellii in nest of Hunter, 1961 Laelapidae Laelaspis piloscutuli Hunter, 1961 Neivamyrmex gibbatus in nest of Hunter, 1961 Laelapidae Laelaspis sp. n/a Pogonomyrmex badius soil in nest Porter, 1985 Laelapidae Laelaspis zuluensis Trägårdh, 1906 Pheidole sp. in nest of Hunter, 1961 Laelapidae Myrmoleichus coronatus Berl. Tapinoma erraticum in nest of Berlese, 1904

152 Laelapidae Myrmonyssus acuminatus Berl. Formica rufa in nest of Donisthorpe, 1927 Laelapidae Myrmonyssus acuminatus Berl. in nest of Berlese, 1904 Laelapidae Myrmonyssus acuminatus n/a Messor capitatus in nest of Vitzthum, 1930 Laelapidae Myrmonyssus acuminatus Berlese, 1904 Messor capitatus in nest of Hunter & Hunter, 1963 Laelapidae Myrmonyssus antennophoroides n/a Camponotus aethiops in nest of Vitzthum, 1930 Laelapidae Myrmonyssus antennophoroides Berl. Camponotus aethiops in nest of Berlese, 1904 Laelapidae Myrmonyssus antennophoroides Berlese, 1904 Camponotus aethiops in nest of Hunter & Hunter, 1963 Laelapidae Myrmonyssus brachiatus n/a Messor capitatus in nest of Vitzthum, 1930 Laelapidae Myrmonyssus brachiatus Berl. Messor capitatus in nest of Berlese, 1904 Laelapidae Myrmonyssus brachiatus Berlese, 1904 Messor capitatus in nest of Hunter & Hunter, 1963 Laelapidae Myrmonyssus clarus Hunter & Hunter, 1963 Crematogaster laeviuscula in nest of Hunter & Hunter, 1963 Laelapidae Myrmonyssus diplogenius n/a Camponotus aethiops in nest of Vitzthum, 1930 Laelapidae Myrmonyssus diplogenius Berl. Camponotus aethiops in nest of Berlese, 1904 Laelapidae Myrmonyssus diplogenius Berlese, 1904 Camponotus aethiops in nest of Hunter & Hunter, 1963 Laelapidae Myrmonyssus eidmanni Sellnick, 1940 Crematogaster impressa in nest of Hunter & Hunter, 1963 Laelapidae Myrmonyssus equalis Banks, 1916 Iridomyrmex gracilis in nest of Hunter & Hunter, 1963 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Laelapidae Myrmonyssus flexuosa Michael, 1981 Camponotus herculeanus in nest of Hunter & Hunter, 1963 Laelapidae Myrmonyssus flexuosus (Mich.) Camponotus herculeanus in nest of Berlese, 1904 Laelapidae Myrmonyssus liguricus n/a Crematogaster scutellaris in nest of Vitzthum, 1930 Laelapidae Myrmonyssus liguricus Vitzthum, 1931 Crematogaster scutellaris in nest of Hunter & Hunter, 1963 Laelapidae Myrmonyssus minor Sellnick, 1941 Crematogaster impressa in nest of Hunter & Hunter, 1963 Laelapidae Myrmonyssus scutellatus Hull, 1923 Iridomyrmex gracilis in nest of Hunter & Hunter, 1963 Laelapidae Myrmonyssus scutellatus Hull, 1923 Iridomyrmex gracilis in nest of Hull, 1923 Laelapidae Myrmonyssus spinosus Hunter & Hunter, 1963 Crematogaster sp. in nest of Hunter & Hunter, 1963 Laelapidae Myrmonyssus titan n/a unknown (ants) in nest of Vitzthum, 1930 Laelapidae Myrmonyssus titan Berlese, 1916 unknown (ants) in nest of Berlese, 1916 Laelapidae Myrmozercon beardae Shaw & Seeman, 2009 unknown (ants) in nest of Shaw & Seeman, 2009 Laelapidae Myrmozercon brevipes Berl. Tapinoma erraticum in nest of Berlese, 1904

153 Laelapidae Myrmozercon brevipes Rosario & Hunter, 1988 unknown (ants) in nest of Rosario & Hunter, 1988 Laelapidae Myrmozercon burwelli Shaw & Seeman, 2009 Polyrhachis flavibasis in nest of Shaw & Seeman, 2009 Laelapidae Myrmozercon iainkayi Walter, 2003 Polyrhachis australis in nest of Shaw & Seeman, 2009 Laelapidae Myrmozercon iainkayi Walter, 2003 Polyrhachis sp. on workers and alates Walter, 2003 Laelapidae Myrmozercon ovatum Karawajew, 1909 Cataglyphus emeryi in nest of Karawajew, 1909 Laelapidae Myrmozercon ovatum Karawajew, 1909 Tapinoma erraticum in nest of Karawajew, 1909 Laelapidae Myrmozercon rotundiscutum Rosario & Hunter, 1988 Camponotus sp. in nest of Rosario & Hunter, 1988 Laelapidae Neoberlesia equitans Berl. Tetramorium caespitum in nest of Berlese, 1904 Laelapidae Neoberlesia equitans Mich. unknown (ants) in nest of Moniez, 1892 Laelapidae Neoberlesia sp. n/a unknown (ants) mixtus in nest of Janet, 1902 Laelapidae reticulatus Berl. Tetramorium caespitum phoretic on ants Hölldobler, 1928 Laelapidae Pachylaelaps sp. n/a Solenopsis fugax phoretic on thorax and gaster Hölldobler, 1928 Laelapidae sp. n/a Pogonomyrmex barbatus in nest of Neece & Bartell, 1982 Laelapidae Pseudoparasitus sp. n/a Pogonomyrmex rugosus in nest of Neece & Bartell, 1982 Laelapidae Sphaeroseius ecitonis (Wasm.) Labidus coecus in nest of Berlese, 1904 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Laelapidae Sphaeroseius ecitonis (Wasm.) Neivamyrmex nigrescens in nest of Berlese, 1904 Larvamimidae Larvamima carli Elzinga, 1993 Eciton burchellii with larvae Elzinga, 1993 Larvamimidae Larvamima carli Elzinga, 1993 Eciton hamatum with larvae Elzinga, 1993 Larvamimidae Larvamima cristata Elzinga, 1993 Neivamyrmex cristatus with larvae Elzinga, 1993 Larvamimidae Larvamima marianae Elzinga, 1993 Eciton hamatum with larvae Elzinga, 1993 Larvamimidae Larvamima marianae Elzinga, 1993 Eciton lucanoides with larvae Elzinga, 1993 Larvamimidae Larvamima marianae Elzinga, 1993 Neivamyrmex pilosus mexicanus with larvae Elzinga, 1993 Larvamimidae Larvamima schneirlai Elzinga, 1993 Eciton burchellii with larvae Elzinga, 1993 Larvamimidae Larvamima schneirlai Elzinga, 1993 Eciton hamatum with larvae Elzinga, 1993 Larvamimidae Larvamima schneirlai Elzinga, 1993 Neivamyrmex pilosus mexicanus with larvae Elzinga, 1993 Krantz, 1962; Macrochelidae Aethosoma burchellestes Krantz, 1962 Eciton burchellii urichi with ants Rettenmeyer, 1962 Macrochelidae micrarrhena Berlese, 1916 Acromyrmex lundii in nest of Berlese, 1916

154 Macrochelidae Holostaspella spectabilis Berlese, 1916 Acromyrmex lundii in nest of Berlese, 1916

Krantz, 1962; Macrochelidae Macrocheles dibamos Krantz, 1962 Eciton vagans mutatum on hind pulvilli Rettenmeyer, 1962 Krantz, 1962; Macrochelidae Macrocheles rettenmeyeri Krantz, 1962 Eciton dulcium crassinode on hind pulvilli Rettenmeyer, 1962 Macrochelidae Macrocheles sp. n/a Myrmecina nipponica bodies in refuse sites Masuko, 1994 Messoracaridae Messoracarus mirandus Silvestri, 1912 Messor barbarus on head Silvestri, 1912 Microgynidae Microsejus truncicola Trägårdh, 1942 Formica aquilonia in nest of Lehtinen, 1987 Microgynidae Microsejus truncicola Trägårdh, 1942 Formica fusca in nest of Lehtinen, 1987 Microgynidae Microsejus truncicola Trägårdh, 1942 Formica rufa in nest of Lehtinen, 1987 Microgynidae Microsejus truncicola Trägårdh, 1942 Leptothorax spp. in nest of Lehtinen, 1987 Nanorchestidae Speleorchestes formicorum Trägårdh, 1909 Formica rufa in nest of Donisthorpe, 1927 Nanorchestidae Sphaerolaelaps holothyroides Leon. Lasius mixtus in nest of Donisthorpe, 1927 Nanorchestidae Sphaerolaelaps holothyroides Leon. Lasius umbratus in nest of Donisthorpe, 1927 Oplitidae Marginura adhaerens Sellnick, 1926 Eciton sp. on spur of ant Sellnick, 1926 Oplitidae Oplitis aktius Hunter & Farrier, 1975 Camponotus floridanus worker tibial spur Hunter & Farrier, 1975 Oplitidae Oplitis aktius Hunter & Farrier, 1975 Camponotus nearcticus queen tibial spur Hunter & Farrier, 1975 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Oplitidae Oplitis alienorum Hunter & Farrier, 1975 Lasius alienus worker tibial spur Hunter & Farrier, 1975 Oplitidae Oplitis anisus Hunter & Farrier, 1975 Pogonomyrmex badius in nest of Hunter & Farrier, 1975 Oplitidae Oplitis arboricavi Hunter & Farrier, 1975 Solenopsis geminata in nest of Hunter & Farrier, 1975 Oplitidae Oplitis blufftonensis Hunter & Farrier, 1975 Camponotus floridanus in nest of Hunter & Farrier, 1975 Oplitidae Oplitis carteretensis Hunter & Farrier, 1975 Solenopsis invicta in nest of Hunter & Farrier, 1975 Oplitidae Oplitis cheleuta Hunter & Farrier, 1975 Formica pallidefulva worker tibial spur Hunter & Farrier, 1975 Oplitidae Oplitis communis Hunter & Farrier, 1975 Solenopsis geminata in nest of Hunter & Farrier, 1975 Oplitidae Oplitis communis Hunter & Farrier, 1975 Solenopsis invicta in nest of Hunter & Farrier, 1975 Oplitidae Oplitis exopodi Hunter & Farrier, 1975 Pachycondyla chinensis in nest of Hunter & Farrier, 1975 Oplitidae Oplitis exopodi Hunter & Farrier, 1975 Solenopsis xyloni in nest of Hunter & Farrier, 1975 Oplitidae Oplitis exsectoidesorum Hunter & Farrier, 1976 Formica exsectoides in nest of Hunter & Farrier, 1976 Oplitidae Oplitis garibaldii Hunter & Farrier, 1976 Lasius alienus in nest of Hunter & Farrier, 1976

155 Oplitidae Oplitis literalis Hunter & Farrier, 1976 Brachymyrmex depilis in nest of Hunter & Farrier, 1976 Oplitidae Oplitis literalis Hunter & Farrier, 1976 Crematogaster atkinsoni in nest of Hunter & Farrier, 1976 Oplitidae Oplitis macclellani Hunter & Farrier, 1976 Pogonomyrmex badius in nest of Hunter & Farrier, 1976 Oplitidae Oplitis moseri Hunter & Farrier, 1976 Solenopsis invicta in nest of Hunter & Farrier, 1976 Oplitidae Oplitis nr. Pandata sspA (Michael, 1894) Formica exsecta in nest of Lehtinen, 1987 Oplitidae Oplitis nr. Pandata sspA (Michael, 1894) Formica rufa in nest of Lehtinen, 1987 Oplitidae Oplitis nr. Pandata sspB (Michael, 1894) Camponotus herculeanus in nest of Lehtinen, 1987 Oplitidae Oplitis nr. villosella (Berlese, 1904) Lasius meridionalis in nest of Lehtinen, 1987 Oplitidae Oplitis ovatula (Berlese, 1903) Tetramorium caespitum in nest of Lehtinen, 1987 Oplitidae Oplitis pandata (Michael, 1894) Lasius niger in nest of Lehtinen, 1987 Oplitidae Oplitis sarcinulus Hunter & Farrier, 1976 Tetramorium caespitum in nest of Hunter & Farrier, 1976 Oplitidae Oplitis sp. n/a Pheidole dentata in nest of Neece & Bartell, 1982 Oplitidae Oplitis sp. n/a Pogonomyrmex badius soil in nest Porter, 1985 Oplitidae Oplitis stammeri Greim in Hirschmann, 1957 Lasius fuliginosus in nest of Lehtinen, 1987 Oplitidae Oplitis trachymyrmecon Hunter & Farrier, 1976 Trachymyrmex septentrionalis in nest of Hunter & Farrier, 1976 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Oplitidae Oplitis villosella (Berlese, 1904) Lasius flavus in nest of Lehtinen, 1987 Hunter & Farrier, Hunter & Farrier, Oplitidae Oplitis virgilinus 1976 Crematogaster atkinsoni in nest of 1976 Hunter & Farrier, Hunter & Farrier, Oplitidae Oplitis virgilinus 1976 Formica dolosa in nest of 1976 Hunter & Farrier, Hunter & Farrier, Oplitidae Oplitis virgilinus 1976 Solenopsis invicta in nest of 1976 Oplitidae Urodiscella alophora (Berlese, 1903) Lasius fuliginosus in nest of Lehtinen, 1987 Oplitidae Urodiscella alophora (Berlese, 1903) Lasius mixtus in nest of Lehtinen, 1987 Oplitidae Urodiscella philoctena (Janet, 1897) Lasius mixtus in nest of Lehtinen, 1987 Oplitidae Urodiscella wasmanni (Kneissl, 1916) Lasius flavus in nest of Lehtinen, 1987 Oplitidae Uroplitella brasiliensis Sellnick, 1926 Eciton quadriglume phoretic on ants Sellnick, 1926 Oplitidae Uroplitella comparata Banks, 1916 Myrmecia sp. in nest of Banks, 1916

156 Oplitidae Uroplitella disparata Banks, 1916 Camponotus aeneopilosus in nest of Banks, 1916 Oplitidae Uroplitella disparata Banks, 1916 Camponotus nigriceps in nest of Banks, 1916

Oplitidae Uroplitella disparata Banks, 1916 Myrmecia pyriformis in nest of Banks, 1916 Oplitidae Uroplitella donisthorpii Hull. Lasius flavus in nest of Donisthorpe, 1927 Oplitidae Uroplitella donisthorpii Hull. Myrmica rubra in nest of Donisthorpe, 1927 Oplitidae Uroplitella donisthorpii Hull. Myrmica scabrinodis in nest of Donisthorpe, 1927 Oplitidae Uroplitella inopina Hull, 1923 Messor meridionalis on spurs of Hull, 1923 Oplitidae Uroplitella minutissima Berl. Formica fusca in nest of Donisthorpe, 1927 Oplitidae Uroplitella minutissima Berl. Lasius flavus in nest of Donisthorpe, 1927 Oplitidae Uroplitella minutissima Berl. Lasius fuliginosus in nest of Donisthorpe, 1927 Oplitidae Uroplitella minutissima Berl. Lasius niger galleries of nest Donisthorpe, 1927 Oplitidae Uroplitella minutissima Berl. Lasius umbratus in nest of Donisthorpe, 1927 Oplitidae Uroplitella minutissima Berl. Pheidole pallidula in nest of Berlese, 1904 Oplitidae Uroplitella minutissima var. villosella Berlese, 1904 Aphaenogaster testaceopilosa in nest of Berlese, 1904 Oplitidae Uroplitella sp. n/a Lasius fuliginosus in nest of Sevastianov, 1965 Pachylaelapidae Gymnolaelaps myrmecophilus (Berlese, 1892) Camponotus ligniperdus in nest of Štorkán, 1940 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Pachylaelapidae Gymnolaelaps myrmecophilus (Berlese, 1892) Formica cunicularia in nest of Štorkán, 1940 Pachylaelapidae Gymnolaelaps myrmecophilus (Berlese, 1892) Formica fusca in nest of Štorkán, 1940 Pachylaelapidae Gymnolaelaps myrmecophilus (Berlese, 1892) Formica rufibarbis in nest of Štorkán, 1940 Pachylaelapidae Gymnolaelaps myrmecophilus (Berlese, 1892) Formica sanguinea in nest of Štorkán, 1940 Pachylaelapidae Gymnolaelaps myrmecophilus (Berlese, 1892) Lasius alienus in nest of Štorkán, 1940 Pachylaelapidae Gymnolaelaps myrmecophilus (Berlese, 1892) Myrmica rubra in nest of Štorkán, 1940 Pachylaelapidae Gymnolaelaps myrmecophilus (Berlese, 1892) Myrmica scabrinodis in nest of Štorkán, 1940 Pachylaelapidae Gymnolaelaps myrmecophilus (Berlese, 1892) Tetramorium caespitum in nest of Štorkán, 1940 Pachylaelapidae Gymnolaelaps myrmophilus (Michael, 1892) Formica sanguinea in nest of Štorkán, 1940 Pachylaelapidae Gymnolaelaps myrmophilus (Michael, 1892) Lasius flavus in nest of Štorkán, 1940 Pachylaelapidae Gymnolaelaps myrmophilus (Michael, 1892) Messor structor in nest of Štorkán, 1940 Pachylaelapidae Gymnolaelaps myrmophilus (Michael, 1892) Myrmica rubra in nest of Štorkán, 1940

157 Pachylaelapidae Gymnolaelaps myrmophilus (Michael, 1892) Myrmica ruginodis in nest of Štorkán, 1940 Pachylaelapidae Laelaspis equitans (Michael, 1891) Tetramorium caespitum in nest of Štorkán, 1940 Pachylaelapidae Laelaspis humeratus (Berlese, 1903) Myrmica scabrinodis in nest of Štorkán, 1940 Pachylaelapidae Laelaspis humeratus (Berlese, 1903) Tetramorium caespitum in nest of Štorkán, 1940 Pachylaelapidae Onchodellus reticulatus Berlese, 1904 Lasius niger in nest of Štorkán, 1940 Pachylaelapidae Onchodellus reticulatus Berlese, 1904 Tetramorium caespitum in nest of Štorkán, 1940 Pachylaelapidae V. Lasius brunneus in nest of Moniez, 1894 Pachylaelapidae Pachylaelaps reticulatus Berlese, 1904 Tetramorium caespitum in nest of Berlese, 1904 Pachylaelapidae Sphaerolaelaps calcariger (Berlese, 1902) Formica rufibarbis in nest of Štorkán, 1940 Pachylaelapidae Sphaerolaelaps calcariger Berl. Lasius mixtus in nest of Berlese, 1904 Pachylaelapidae Sphaerolaelaps calcariger (Berlese, 1902) Solenopsis fugax in nest of Štorkán, 1940 Pachylaelapidae Sphaerolaelaps calcariger (Berlese, 1902) Tapinoma erraticum in nest of Štorkán, 1940 Pachylaelapidae Sphaerolaelaps holothyroides (Leon.) Lasius citrinus in nest of Berlese, 1904 Pachylaelapidae Sphaerolaelaps holothyroides (Leon.) Lasius umbratus in nest of Berlese, 1904 Pachylaelapidae Sphaerolaelaps holothyroides (Leonardi, 1896) Lasius umbratus in nest of Štorkán, 1940 Pachylaelapidae Sphaerolaelaps holothyroides (Leon.) Pheidole pallidula in nest of Berlese, 1904 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Celaenopsis breviatus Banks, 1916 Iridomyrmex sp in nest of Banks, 1916 Parasitidae femoralis Banks, 1916 Rhytidoponera metallica in nest of Banks, 1916 Parasitidae Cyrtolaelaps gracilipes Banks, 1916 Camponotus nigriceps in nest of Banks, 1916 Parasitidae Cyrtolaelaps gracilipes Banks, 1916 Pachycondyla lutea in nest of Banks, 1916 Parasitidae Cyrtolaelaps gracilipes Banks, 1916 Polyrhachis hexacantha in nest of Banks, 1916 Parasitidae Cyrtolaelaps gracilipes Banks, 1916 Rhytidoponera metallica in nest of Banks, 1916 Parasitidae Gamasus canestrinii Berl. Formica fusca in nest of Moniez, 1894 Parasitidae Gamasus crassipes L. Formica exsecta in nest of Moniez, 1894 Parasitidae Gamasus crassipes n/a Formica rufa in nest of Wasmann, 1899 Parasitidae Gamasus crassipes L. Formica rufa in nest of Moniez, 1894 Parasitidae Gamasus crassipes L. Formica rufibarbis in nest of Moniez, 1894 Parasitidae Gamasus crassipes L. Lasius brunneus in nest of Moniez, 1894

158 Parasitidae Gamasus crassipes L. Lasius niger in nest of Moniez, 1894 Parasitidae Gamasus crassipes Linnee Lasius niger in nest of Moniez, 1892 Parasitidae Hypoaspis inepilis Banks, 1916 Camponotus sp. in nest of Banks, 1916 Parasitidae Hypoaspis minusculus Banks, 1916 unknown (ants) in nest of Banks, 1916 Parasitidae Myrmonyssus aequalis Banks, 1916 Iridomyrmex gracilis on thorax of ant Banks, 1916 Parasitidae Neoberlesia equitans Berl. Pheidole pallidula on ants Wasmann, 1902 Parasitidae Neoberlesia mexicana Banks, 1915 Pheidole hirtula in nest of Banks, 1915 Parasitidae disparatus Banks, 1916 Polyrhachis hexacantha in nest of Banks, 1916 Parasitidae Parasitus inversus Banks, 1916 Rhytidoponera metallica in nest of Banks, 1916 Parasitidae Parasitus lyratus Banks, 1916 Amblyopone australis in nest of Banks, 1916 Parasitidae Parasitus sp. n/a Aphaenogaster cockerelli in nest of Neece & Bartell, 1982 Parasitidae Parasitus sp. n/a Myrmecina nipponica bodies in refuse sites Masuko, 1994 Parasitidae Parasitus sp. n/a Solenopsis aurea in nest of Neece & Bartell, 1982 Parasitidae sp. n/a Pheidole flavens soil in nest Wilson, 2005 Parholaspididae Holaspis terreus Can. Formica rufa in nest of Wasmann, 1899 Parholaspididae Parholaspis sp. n/a Myrmecina flava bodies in refuse sites Masuko, 1994 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Physalozerconidae Physalozercon raffray (Wasm.) Anoplolepis custodiens in nest of Berlese, 1904 fraterculus Berlese, 1916 Acromyrmex lundii in nest of Berlese, 1916 Phytoseiidae keegani Fox 1947 unknown (ants) on ant eggs Rack, 1968 Phytoseiidae? Garmania sp. n/a Cyphomyrmex rimosus thorax & gaster Weber, 1972 Trachymyrmex Phytoseiidae? Garmania sp. n/a septentrionalis thorax & gaster Weber, 1972 Planodiscidae Planodiscus borgmeieri Elzinga, 1990 Neivamyrmex gradualis in nest of Elzinga, 1990 Planodiscidae Planodiscus borgmeieri Elzinga, 1990 Neivamyrmex legionis in nest of Elzinga, 1990 Planodiscidae Planodiscus burchelli Elzinga & Rettenmeyer, 1966 Eciton quadriglume on ants Elzinga & Rettenmeyer, 1966 Planodiscidae Planodiscus capillilatus Elzinga, 1991 Eciton drepanophorum in nest of Elzinga, 1991 Planodiscidae Planodiscus cupiens Elzinga & Rettenmeyer, 1970 Eciton burchellii cupiens on ants Elzinga & Rettenmeyer, 1970 Planodiscidae Planodiscus elongatus Elzinga & Rettenmeyer, 1970 Eciton dulcium crassinode phoretic on ants Elzinga & Rettenmeyer, 1970 Planodiscidae Planodiscus elongatus Elzinga & Rettenmeyer, 1970 Eciton hamatum phoretic on ants Elzinga & Rettenmeyer, 1970

159 Neivamyrmex pilosus

Planodiscidae Planodiscus elongatus Elzinga & Rettenmeyer, 1970 mexicanus phoretic on ants Elzinga & Rettenmeyer, 1970 Planodiscidae Planodiscus foreli Elzinga & Rettenmeyer, 1970 Eciton burchellii foreli phoretic on ants Elzinga & Rettenmeyer, 1970 Eciton burchellii Planodiscidae Planodiscus foreli Elzinga & Rettenmeyer, 1970 parvispinum phoretic on ants Elzinga & Rettenmeyer, 1970 Planodiscidae Planodiscus hamatus Elzinga & Rettenmeyer, 1970 Eciton hamatum phoretic on ants Elzinga & Rettenmeyer, 1970 Planodiscidae Planodiscus hamatus Elzinga & Rettenmeyer, 1970 Eciton vagans phoretic on ants Elzinga & Rettenmeyer, 1970 Planodiscidae Planodiscus kistneri Elzinga, 1991 Eciton drepanophorum in nest of Elzinga, 1991 Planodiscidae Planodiscus mexicanus Elzinga, 1991 Eciton hamatum in nest of Elzinga, 1990 Planodiscidae Planodiscus setosus Elzinga & Rettenmeyer, 1970 Eciton burchellii cupiens phoretic on ants Elzinga & Rettenmeyer, 1970 Planodiscidae Planodiscus sqamatim Sellnick, 1926 Eciton quadriglume phoretic on ants Sellnick, 1926 Planodiscidae Planodiscus squamatim Elzinga & Rettenmeyer, 1966 Eciton quadriglume on ants Elzinga & Rettenmeyer, 1966 Dipolyaspis testaceus (C.L. Koch, 1836), Wisniewski, criocephali 1979 Camponotus herculeanus in nest of Lehtinen, 1987 Polyaspididae modesta (Leonardi, 1899) Camponotus ligniperdus in nest of Lehtinen, 1987 Polyaspididae Discourella modesta (Leonardi, 1899) Formica fusca in nest of Lehtinen, 1987 Polyaspididae Discourella modesta (Leonardi, 1899) Lasius sp. in nest of Lehtinen, 1987 Polyaspididae platensis Berlese, 1916 Acromyrmex lundii in nest of Berlese, 1916 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Polyaspididae Trachytes aegrota (C.L. Koch, 1841) unknown (many ants) in nests Lehtinen, 1987 Polyaspididae Trachytes minima Trägårdh, 1910 Formica rufa group in nest of Lehtinen, 1987 Polyaspididae Trachytes minima Trägårdh, 1910 Myrmica sp. in nest of Lehtinen, 1987 Prodinychidae Dinychus arcuatus Trägårdh, 1943 in nest of Lehtinen, 1987 Prodinychidae Dinychus carinatus Berlese, 1903 Lasius spp. in nest of Lehtinen, 1987 Prodinychidae Dinychus carinatus Berlese, 1903 Myrmica spp. in nest of Lehtinen, 1987 Prodinychidae Dinychus septentrionalis (Trägårdh, 1943) Formica polyctena in nest of Lehtinen, 1987 Prodinychidae Prodinychus flagelliger (Berlese, 1910) Formica fusca in nest of Lehtinen, 1987 Prodinychidae Trachyxenura pyriformis (Berlese, 1920) Formica aquilonia in nest of Lehtinen, 1987 Prodinychidae Trachyxenura pyriformis (Berlese, 1920) Formica exsecta in nest of Lehtinen, 1987 Rhodacaridae sp. n/a Pogonomyrmex badius soil in nest Porter, 1985 togatus (C.L. Koch, 1836) Formica polyctena in nest of Lehtinen, 1987

160 phoretic on gen. sp. n/a Solenopsis invicta female alates Hermann et al., 1970

Trachyuropodidae Antennequesoma labergei Elzinga, 1982b Neivamyrmex opacithorax on ants Elzinga, 1982b Trachyuropodidae Antennequesoma longissima Elzinga, 1982b Neivamyrmex sumichrasti on ants Elzinga, 1982b Trachyuropodidae Antennequesoma lujai Sellnick, 1926 Eciton burchellii on the feet Sellnick, 1926 Trachyuropodidae Antennequesoma reichenspergeri Sellnick, 1926 Eciton quadriglume phoretic on ants Sellnick, 1926 Trachyuropodidae Antennequesoma rettenmeyeri Elzinga, 1982b Nomamyrmex esenbeckii crassicornis on ants Elzinga, 1982b Trachyuropodidae Antennequesoma tennuatum Elzinga, 1982b Labidus praedator with larvae Elzinga, 1982b Trachyuropodidae Leonardiella riccardiana (Leonardi, 1895) Formica rufibarbis in nest of Štorkán, 1940 Trachyuropodidae Leonardiella riccardiana (Leonardi, 1895) Lasius alienus in nest of Štorkán, 1940 Trachyuropodidae Leonardiella riccardiana (Leonardi, 1895) Tetramorium caespitum in nest of Štorkán, 1940 Trachyuropodidae Trachuropoda canestriniana Berl. Tetramorium caespitum in nest of Berlese, 1904 Trachyuropodidae Trachuropoda excavata (Wasm.) Lasius niger in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda bostoki (Mich.) Lasius flavus in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda bostocki (Michael, 1894) Lasius fuliginosus in nest of Štorkán, 1940 Trachyuropodidae Trachyuropoda bostocki Mich. Lasius mixtus in nest of Donisthorpe, 1927 Trachyuropodidae Trachyuropoda bostocki Mich. Lasius umbratus in nest of Donisthorpe, 1927 Continued

Table 29 continued MITE HIGHER LEVEL MITE SPECIES AUTHOR HOST SPECIES ASSOCIATION REFERENCE Trachyuropodidae Trachyuropoda bostoki (Mich.) Lasius brunneus in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda bostoki (Mich.) Lasius mixtus in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda canestriniana Berl. Tetramorium caespitum in nest of Donisthorpe, 1927 Trachyuropodidae Trachyuropoda canestriniana var. insularis (Berl.) Berl. Tetramorium meridionale in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda canestriniana var. septentrionalis Berl. Tetramorium caespitum in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda coccinea Mich. Camponotus ligniperdus in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda coccinea Mich. Formica fusca in nest of Donisthorpe, 1927 Trachyuropodidae Trachyuropoda coccinea Mich. Formica fusca in nest of Donisthorpe, 1927 Trachyuropodidae Trachyuropoda coccinea Mich. Formica rufa in nest of Donisthorpe, 1927 Trachyuropodidae Trachyuropoda coccinea Mich. Lasius niger in nest of Donisthorpe, 1927 Trachyuropodidae Trachyuropoda coccinea var. sinuata (Mich.) Berl. Aphaenogaster testaceopilosa in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda coccinea var. sinuata (Mich.) Berl. Camponotus aethiops in nest of Berlese, 1904

161 Trachyuropodidae Trachyuropoda coccinea var. sinuata (Mich.) Berl. Formica fusca in nest of Berlese, 1904

Trachyuropodidae Trachyuropoda coccinea var. sinuata (Mich.) Berl. Formica pratensis in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda coccinea var. sinuata (Mich.) Berl. Formica rufibarbis in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda coccinea var. sinuata (Mich.) Berl. Formica sanguinea in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda coccinea var. sinuata (Mich.) Berl. Lasius flavus in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda coccinea var. sinuata (Mich.) Berl. Lasius niger in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda coccinea var. sinuata (Mich.) Berl. Myrmica scabrinodis in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda constricta Banks, 1916 Pachycondyla lutea in nest of Banks, 1916 Trachyuropodidae Trachyuropoda cristiceps (Can.) Camponotus ligniperdus in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda criticeps (Can.) Pheidole pallidula in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda escavata (Wasm.) Tetramorium caespitum in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda excavata Wasm. Lasius fuliginosus in nest of Donisthorpe, 1927 Trachyuropodidae Trachyuropoda excavata Wasm. Lasius niger in nest of Donisthorpe, 1927 Trachyuropodidae Trachyuropoda laminosa (C. et. B.) Formica fusca in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda laminosa C. & B. Formica fusca in nest of Donisthorpe, 1927 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Trachyuropodidae Trachyuropoda laminosa (C. et. B.) Lasius flavus in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda laminosa C. & B. Lasius flavus in nest of Donisthorpe, 1927 Trachyuropodidae Trachyuropoda laminosa (C. et. B.) Lasius niger in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda laminosa C. & B. Lasius niger in nest of Donisthorpe, 1927 Trachyuropodidae Trachyuropoda laminosa (C. et. B.) Tetramorium caespitum in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda laminosa C. & B. Tetramorium caespitum in nest of Donisthorpe, 1927 Trachyuropodidae Trachyuropoda leai Banks, 1916 Aphaenogaster longiceps? in nest of Banks, 1916 Trachyuropodidae Trachyuropoda magna (Leon.) Camponotus aethiops in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda riccardiana (Leon.) Lasius alienus in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda riccardiana (Leon.) Myrmica scabrinodis in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda riccardiana (Leon.) Solenopsis fugax in nest of Berlese, 1904 Trachyuropodidae Trachyuropoda wasmanniana Berl. Lasius mixtus in nest of Berlese, 1904

162 Trachyuropodidae Trachyuropoda wasmanniana Berl. Lasius umbratus in nest of Donisthorpe, 1927 Trachyuropodidae Urojanetia coccinea (Michael, 1891) Camponotus ligniperdus in nest of Štorkán, 1940 Trachyuropodidae Urojanetia coccinea (Michael, 1891) Formica in nest of Lehtinen, 1987 Trachyuropodidae Urojanetia coccinea (Michael, 1891) Formica cunicularia in nest of Štorkán, 1940 Trachyuropodidae Urojanetia coccinea (Michael, 1891) Formica exsecta in nest of Lehtinen, 1987 Trachyuropodidae Urojanetia coccinea (Michael, 1891) Formica rufibarbis in nest of Štorkán, 1940 Trachyuropodidae Urojanetia coccinea (Michael, 1891) Formica sanguinea in nest of Lehtinen, 1987 Trachyuropodidae Urojanetia coccinea (Michael, 1891) Formica sanguinea in nest of Štorkán, 1940 Trachyuropodidae Urojanetia coccinea (Michael, 1891) Lasius niger in nest of Štorkán, 1940 Trachyuropodidae Urojanetia coccinea (Michael, 1891) Myrmica rubra in nest of Štorkán, 1940 Trachyuropodidae Urojanetia coccinea (Michael, 1891) Myrmica scabrinodis in nest of Štorkán, 1940 Trachyuropodidae Urojanetia cristiceps (Canestrini, 1884) Camponotus ligniperdus in nest of Štorkán, 1940 Trachyuropodidae Urojanetia cristiceps (Canestrini, 1884) Formica cunicularia in nest of Štorkán, 1940 Trachyuropodidae Urojanetia cristiceps (Canestrini, 1884) Formica fusca in nest of Štorkán, 1940 Trachyuropodidae Urojanetia cristiceps (Canestrini, 1884) Formica sanguinea in nest of Štorkán, 1940 Trachyuropodidae Urojanetia cristiceps (Canestrini, 1884) Lasius alienus in nest of Štorkán, 1940 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Trachyuropodidae Urojanetia cristiceps (Canestrini, 1884) Myrmica rubra in nest of Štorkán, 1940 Trachyuropodidae Urojanetia cristiceps (Canestrini, 1884) Myrmica scabrinodis in nest of Štorkán, 1940 Trachyuropodidae Urojanetia excavata (Wasmann, 1899) Formica fusca in nest of Štorkán, 1940 Trachyuropodidae Urojanetia excavata (Wasmann, 1899) Tetramorium caespitum in nest of Štorkán, 1940 Trachyuropodidae Urojanetia excavata (Wasmann,1899) Tetramorium caespitum in nest of Lehtinen, 1987 Trachyuropodidae Urojanetia hirschmanni (Pecina, 1980) Lasius meridionalis in nest of Lehtinen, 1987 Trachyuropodidae Urojanetia lamellosa (Canestrini, et Berlese, 1884) Lasius alienus in nest of Štorkán, 1940 Trachyuropodidae Urojanetia magna (Leonardi, 1895) Camponotus ligniperdus in nest of Štorkán, 1940 Trachyuropodidae Urojanetia magna (Leonardi, 1895) Messor structor in nest of Štorkán, 1940 Trachyuropodidae Urojanetia troguloides Gerv. Formica sanguinea in nest of Štorkán, 1940 Trachyuropodidae Urojanetia troguloides Gerv. Lasius alienus in nest of Štorkán, 1940 Trachyuropodidae Urojanetia troguloides Gerv. Lasius niger niger in nest of Štorkán, 1940

163 Trachyuropodidae Urojanetia troguloides Gerv. Myrmica scabrinodis in nest of Štorkán, 1940 Trachyuropodidae Urojanetia wasmanniana (Berlese, 1903) Lasius mixtus in nest of Lehtinen, 1987 Trachyuropodidae Uropolyaspis hamuliferus (Michael, 1894) Lasius niger in nest of Štorkán, 1940 Trachyuropodidae Urotrachytes formicaria (Lubbock, 1881) Lasius flavus in nest of Lehtinen, 1987 Trachyuropodidae Urotrachytes formicarius (Lubbock, 1881) Lasius alienus in nest of Štorkán, 1940 Trachyuropodidae Urotrachytes formicarius (Lubbock, 1881) Lasius flavus in nest of Štorkán, 1940 Trachyuropodidae Urotrachytes formicarius Lubb. Lasius flavus in nest of Donisthorpe, 1927 Trachyuropodidae Urotrachytes formicarius (Lubbock, 1881) Lasius niger in nest of Štorkán, 1940 Trematurinae Dinychopsis catula Hull, 1918 Formica polyctena in nest of Lehtinen, 1987 Trematurinae Ipiduropoda dialveolata Hirschmann & Z. -Nicol, 1961 Lasius flavus in nest of Lehtinen, 1987 Trematurinae Ipiduropoda dialveolata Hirschmann & Z. -Nicol, 1961 Lasius fuliginosus in nest of Lehtinen, 1987 Trematurinae Ipiduropoda dialveolata Hirschmann & Z. -Nicol, 1961 Lasius meridionalis in nest of Lehtinen, 1987 Trematurinae Ipiduropoda interstructura Hirschmann & Z. -Nicol, 1961 Formica pratensis in nest of Lehtinen, 1987 Trematurinae Oodinychus ovalis (C.L. Koch, 1839) Camponotus spp. in nest of Lehtinen, 1987 Trematurinae Oodinychus ovalis (C.L. Koch, 1839) Formica in nest of Lehtinen, 1987 Trematurinae Oodinychus ovalis (C.L. Koch, 1839) Lasius spp. in nest of Lehtinen, 1987 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Trematurinae Oodinychus spatuliferus beckwithi (Moniez, 1892), Wisniewski, 1979 Camponotus herculeanus in nest of Lehtinen, 1987 Trematurinae Oodinychus spatuliferus beckwithi (Moniez, 1892), Wisniewski, 1979 in nest of Lehtinen, 1987 Trematurinae Trematurella elegans (Berlese, 1916) Formica polyctena in nest of Lehtinen, 1987 Trematurinae Trematurella elegans (Berlese, 1916) Lasius niger in nest of Lehtinen, 1987 Trematurinae Urodiaspis tecta (Kramer, 1876) unknown (ants) in nest Lehtinen, 1987 Trematurinae Uroobovella obovata (Canestrini & Berlese, 1884) Formica fusca in nest of Lehtinen, 1987 Trematurinae Uroobovella obovata (Canestrini & Berlese, 1884) Lasius niger in nest of Lehtinen, 1987 Oodinychus janeti (Berlese, 1904) Formica fusca in nest of Štorkán, 1940 Urodinychidae Urobovella cylliboides Hull, 1923 Iridomyrmex sp in nest of Hull, 1923 Urodinychidae Urodinychus janeti Berl. Formica fusca in nest of Donisthorpe, 1927 Urodinychidae Urodinychus janeti Berl. Formica rufa in nest of Donisthorpe, 1927 Urodinychidae Urodiscella ovatula Berl. Lasius flavus in nest of Donisthorpe, 1927

164 Urodinychidae Urodiscella philoctena Trous. Lasius flavus strigil (spur) Donisthorpe, 1927 Urodinychidae Urodiscella philoctena Trous. Lasius fuliginosus strigil (spur) Donisthorpe, 1927 Urodinychidae Urodiscella philoctena Trous. Lasius umbratus strigil (spur) Donisthorpe, 1927 Urodinychidae Urodiscella ricasoliana Berl. Lasius fuliginosus in nest of Donisthorpe, 1927 Urodinychidae Urodiscella ricasoliana Berl. Lasius umbratus in nest of Donisthorpe, 1927 Urodinychidae Urodiscella signata Hull. Lasius flavus in nest of Donisthorpe, 1927 Urodinychidae Uroobovella notabilis Berl. Formica fusca in nest of Donisthorpe, 1927 Urodinychidae Uroobovella obovata C. & B. Formica fusca in nest of Donisthorpe, 1927 Urodinychidae Uroobovella obovata C. & B. Lasius flavus in nest of Donisthorpe, 1927 Uropidina Uroseius myrmecophilus Wiśniewski, 1979 Formica polyctena in nest of Wiśniewski, 1979 Uropodidae Discopoma pulcherrima Berl. Formica fusca in nest of Donisthorpe, 1927 Uropodidae Disparipes nov. sp. n/a Lasius mixtus on dead ants Janet, 1902 Uropodidae Dispocoma comata Berlese Lasius mixtus on larvae Janet, 1902 phoretic on Uropodidae gen. sp. n/a Solenopsis invicta female alates Hermann et al., 1970 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Uropodidae Trichocylliba comata (Leonardi, 1896) Lasius alienus in nest of Štorkán, 1940 Uropodidae Trichocylliba comata (Leonardi, 1896) in nest of Štorkán, 1940 Uropodidae Trichocylliba comata (Leonardi, 1896) Lasius flavus in nest of Štorkán, 1940 Uropodidae Trichocylliba comata (Leonardi, 1896) Lasius niger in nest of Štorkán, 1940 Uropodidae Trichocylliba comata (Leonardi, 1896) Lasius umbratus in nest of Štorkán, 1940 Uropodidae Trichocylliba comata (Leonardi, 1896) Tetramorium caespitum in nest of Štorkán, 1940 Uropodidae undet sp. n/a Pheidole megacephala parasitic on pupae Le Breton, 2006 attached to dorsum of Uropodidae undetermined sp. n/a Solenopsis richteri alate females Collins & Markin, 1971 Uropodidae Urodiscella ricasoliana (Berlese, 1889) Camponotus ligniperdus in nest of Štorkán, 1940 Uropodidae Urodiscella ricasoliana (Berlese, 1889) Formica rufa in nest of Štorkán, 1940 Uropodidae Uroplitella minutissima Berlese, 1903 Lasius alienus in nest of Štorkán, 1940 Uropodidae Uroplitella minutissima Berlese, 1903 Lasius umbratus in nest of Štorkán, 1940

165 Uropodidae Uroplitella paradoxa (Canestrini, et Berlese, 1884) Myrmica scabrinodis in nest of Štorkán, 1940

Uropodidae Uroplitella paradoxa (Canestrini, et Berlese, 1884) Tetramorium caespitum in nest of Štorkán, 1940 Uropodidae Uropolyaspis hamuliferus Mich. Lasius flavus on femur of ant Donisthorpe, 1927 Uropodidae Uropolyaspis hamuliferus Mich. Lasius mixtus middle of femurs Donisthorpe, 1927 Uropodidae Uropolyaspis hamuliferus Mich. Lasius niger on femur of ant Donisthorpe, 1927 Uropodidae Uropolyaspis hamuliferus Mich. Lasius umbratus on femur of ant Donisthorpe, 1927 Uropodina Cilliba comata Leon Lasius fuliginosus in nest of Wasmann, 1902 Uropodina Cilliba pandata Mich. Lasius fuliginosus in nest of Wasmann, 1902 Uropodina Cillibano comata Leon. Lasius flavus gaster of ants Donisthorpe, 1927 Uropodina Cillibano comata Leon. Lasius mixtus gaster and larvae Donisthorpe, 1927 Uropodina Cillibano comata Leon. Lasius umbratus gaster and larvae Donisthorpe, 1927 Uropodina Cillibano hirticoma Berl. Neivamyrmex nigrescens in nest of Berlese, 1904 Uropodina Cillibano transversalis Karawajew, 1909 Messor excursionis in nest of Karawajew, 1909 Wiśniewski & Hirschmann, Wiśniewski & Hirschmann, Uropodina Dinychus camponoti 1983 Camponotus herculeanus in nest of 1983 Uropodina Discopoma comata Berlese Lasius mixtus parasitic on ants Janet, 1897b Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Uropodina Glyphopsis coccinea Mich. Formica sanguinea in nest of Wasmann, 1899 Uropodina Glyphopsis bostocki Mich. Lasius brunneus in nest of Wasmann, 1899 Uropodina Glyphopsis bostocki Michael Lasius flavus in nest of Janet, 1902 Uropodina Glyphopsis bostocki Michael Lasius mixtus in nest of Janet, 1902 Uropodina Glyphopsis coccinea Mich. Formica fusca in nest of Wasmann, 1899 Uropodina Glyphopsis coccinea Mich. Formica rufa in nest of Wasmann, 1899 Uropodina Glyphopsis coccinea Mich. Formica rufibarbis in nest of Wasmann, 1899 Uropodina Glyphopsis coccinea Mich. Lasius flavus in nest of Wasmann, 1899 Uropodina Glyphopsis coccinea Mich. Lasius niger in nest of Wasmann, 1899 Uropodina Glyphopsis coccinea Mich. Myrmica scabrinodis in nest of Wasmann, 1899 Uropodina Glyphopsis coccinea var. excavata n/a Lasius fuliginosus in nest of Wasmann, 1902 Uropodina Glyphopsis coccinea var.excavata Mich. Tetramorium caespitum in nest of Wasmann, 1899

166 Uropodina Glyphopsis formicariae Lubb. Lasius flavus in nest of Wasmann, 1899 Uropodina Glyphopsis lamellosa Canestr. Lasius flavus in nest of Wasmann, 1899 Uropodina Glyphopsis lamellosa Canestr. Lasius fuliginosus in nest of Wasmann, 1899 Uropodina Glyphopsis lamellosa n/a Lasius niger on ants Wasmann, 1902 Uropodina Glyphopsis lamellosa Canestr. Lasius niger in nest of Wasmann, 1899 Uropodina Janetiella auricularis Hull, 1923 Iridomyrmex gracilis on spurs of Hull, 1923 Uropodina Macrodynichus multispinosus Sellnick fulva parasitic on pupae Krantz et al., 2007 Uropodina Macrodynichus sellnicki Hirschmann & Zirngiebl-Nicol Nylanderia fulva parasitic on pupae Krantz et al., 2007 carrying fungus Balazy & Wisniewski, Uropodina Trachyuropoda coccinea (Michael, 1891) Formica in nest 1982 carrying fungus Balazy & Wisniewski, Uropodina Trachyuropoda coccinea (Michael, 1891) Formica polyctena in nest 1982 Mašán, 1999, cites Uropodina Trichocylliba comata (Leonardi, 1895) Camponotus sp. on ants Bernard, 1968 on adults and Uropodina Trichocylliba comata (Leonardi, 1895) Lasius flavus larvae Mašán, 1999 on adults and Mašán, 1999, cites Uropodina Trichocylliba comata (Leonardi, 1895) Lasius niger larvae Bernard, 1968 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Mašán, 1999, cites Uropodina Trichocylliba comata (Leonardi, 1895) Myrmica sp. on ants Bernard, 1968 Uropodina Trichouropoda beckwithi Wiśniewski, 1980 Camponotus herculeanus in nest of Wiśniewski, 1980 Uropodina Urodinychus janeti Berlese, 1904 Formica rufa in nest of Berlese, 1904 Uropodina Urodinychus karawaiewi Berl. Myrmica scabrinodis in nest of Berlese, 1904 Uropodina Urodinychus mancus Berlese, 1916 Acromyrmex lundii in nest of Berlese, 1916 Uropodina Urodiscella alophora Berl. Lasius mixtus in nest of Berlese, 1904 Uropodina Urodiscella philoctena (Trt.) Lasius mixtus in nest of Berlese, 1904 Uropodina Urodiscella philoctena (Trt.) Messor structor in nest of Berlese, 1904 Uropodina Urodiscella ricasoliana Berl. Lasius fuliginosus in nest of Berlese, 1904 Uropodina Urodynichus parallelepipedus Berlese, 1916 Dorymyrmex steigeri in nest of Berlese, 1916 Uropodina Uroobovella notabilis Berl Formica fusca in nest of Berlese, 1904 Uropodina Uroobovella notabilis Berl Lasius niger in nest of Berlese, 1904

167 Uropodina Uroobovella obovata (C. et. B.) Lasius alienus in nest of Berlese, 1904

Uropodina Uroobovella obovata (C. et. B.) Solenopsis fugax in nest of Berlese, 1904 Uropodina Uroobovella wasmanni Kniessl, 1907 Lasius flavus in nest of Kneissl, 1907 Uropodina Urooplitella minutissima Berl. Formica fusca in nest of Berlese, 1904 Uropodina Urooplitella minutissima Berl. Lasius alienus in nest of Berlese, 1904 Uropodina Uroplitella leonardiana Berl. Messor structor in nest of Berlese, 1904 Uropodina Uroplitella minutissima Berl. Lasius niger in nest of Berlese, 1904 Uropodina Uroplitella minutissima Berl. Myrmica scabrinodis in nest of Berlese, 1904 Uropodina Uroplitella minutissima var. villosella Berlese, 1904 Camponotus aethiops in nest of Berlese, 1904 Uropodina Uroplitella minutissima var. villosella Berlese, 1904 Myrmica scabrinodis in nest of Berlese, 1904 Uropodina Uroplitella pennsylvanica Berl. Formica rubicunda in nest of Berlese, 1904 Uropodina Uroplitella pennsylvanica Berl. Formica subsericea in nest of Berlese, 1904 Uropodina amblyoponae Banks, 1916 Amblyopone australis in nest of Banks, 1916 Uropodina Uropoda apicata Banks, 1916 Iridomyrmex rufoniger in nest of Banks, 1916 Uropodina Uropoda apicata Banks, 1916 Pachycondyla lutea in nest of Banks, 1916 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Uropodina Uropoda bifrons Banks, 1916 Myrmecia pyriformis in nest of Banks, 1916 Uropodina Uropoda bilobata Banks, 1916 Aphaenogaster longiceps? in nest of Banks, 1916 Uropodina Uropoda bruckii Berlese, 1916 Acromyrmex lundii in nest of Berlese, 1916 Uropodina Uropoda canestriniana Berlese, 1891? Tetramorium caespitum in nest of Moniez, 1892 Uropodina Uropoda coccinea Mich. Camponotus herculeanus in nest of Moniez, 1892 Uropodina Uropoda coccinea Michael, 1891 Camponotus herculeanus in nest of Michael, 1891 Uropodina Uropoda coccinea Michael, 1891 Formica fusca in nest of Michael, 1891 Uropodina Uropoda convexifrons Banks, 1916 Rhytidoponera metallica in nest of Banks, 1916 Uropodina Uropoda dentifrons Banks, 1916 Rhytidoponera metallica in nest of Banks, 1916 Uropodina Uropoda duplicata Banks, 1916 Polyrhachis ammon in nest of Banks, 1916 Uropodina Uropoda fraterna Banks, 1916 Polyrhachis hexacantha in nest of Banks, 1916 Uropodina Uropoda hamulifera Mich. Lasius fuliginosus in nest of Wasmann, 1902

168 Uropodina Uropoda hamulifera Mich. Lasius niger in nest of Wasmann, 1899 Uropodina Uropoda internata Banks, 1916 Iridomyrmex sp in nest of Banks, 1916 Uropodina Uropoda krameri Canestr. Formica fusca in nest of Wasmann, 1899 Uropodina Uropoda krameri Canestr. Lasius fuliginosus in nest of Wasmann, 1902 Uropodina Uropoda krameri Canestr. Lasius niger in nest of Wasmann, 1899 Uropodina Uropoda lamellosa Canestr. Et Berl. Tetramorium caespitum in nest of Moniez, 1892 Uropodina Uropoda longifrons Banks, 1916 Rhytidoponera metallica in nest of Banks, 1916 Uropodina Uropoda nov. sp n/a Lasius mixtus on leg of Janet, 1902 Uropodina Uropoda obliquifrons Banks, 1916 Amblyopone australis in nest of Banks, 1916 Uropodina Uropoda obscura Berl. Crematogaster schenki in nest of Moniez, 1894 Uropodina Uropoda ovalis Koch Lasius mixtus femur on stalk Janet, 1902 Uropodina Uropoda productior Berlese, 1916 Camponotus ligniperdus in nest of Berlese, 1916 Uropodina Uropoda ricasoliana Berl. Lasius fuliginosus in nest of Wasmann, 1899 Uropodina Uropoda ricasoliana Berl; Wasmann Lasius fuliginosus in nest of Moniez, 1892 Uropodina Uropoda sp. n/a Myrmecina flava bodies in refuse sites Masuko, 1994 Continued

Table 29 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Uropodina Uropoda spatulifera Moniez Formica rufa in nest of Wasmann, 1899 Uropodina Uropoda spatulifera Moniez, 1892 Formica rufa in nest of Moniez, 1892 Uropodina Uropoda submarginata Banks, 1916 Camponotus aeneopilosus in nest of Banks, 1916 Uropodina Uropoda tasmanica Banks, 1916 Myrmecia pyriformis in nest of Banks, 1916 Uropodina Uropoda trilobata Banks, 1916 unknown (ants) in nest of Banks, 1916 Uropodina Uropoda vegetans n/a Formica rufa in nest of Wasmann, 1899 Uropodina Uropolyaspis hamuliferus Mich. Lasius niger in nest of Berlese, 1904 Uropodina Uroseius koehleri Wiśniewski, 1979 Formica rufa group in nest of Wiśniewski, 1979 Uropodina Urotrachytes formicarius (Lubb.) Lasius flavus in nest of Berlese, 1904 Uropodinae Microcylliba minima Kramer, 1882 Lasius mixtus in nest of Lehtinen, 1987 Uropodinae Phaulodinychus hamulifer (Michael, 1894) Lasius niger in nest of Lehtinen, 1987 Uropodinae Phaulodinychus sp. n/a Camponotus herculeanus in nest of Lehtinen, 1987

169 Uropodinae Phaulodinychus spinulosus (Kneissl, 1916) Lasius mixtus in nest of Lehtinen, 1987 Uropodinae Uroseius koehleri Wisniewski, 1979 Formica rufa in nest of Lehtinen, 1987 Parazercon radiatus (Berlese, 1910) unknown (many ants) in nest of Lehtinen, 1987 Zerconidae kochi Sellnick, 1943 unknown (many ants) in nest of Lehtinen, 1987 Zerconidae Prozercon traegardhi (Halbert, 1915) Lasius flavus in nest of Lehtinen, 1987 Zerconidae curiosus Trägårdh, 1910 Camponotus spp. in nest of Lehtinen, 1987 Zerconidae Zercon curiosus Trägårdh, 1910 Formica in nest of Lehtinen, 1987 Zerconidae Zercon curiosus Trägårdh, 1910 Lasius spp. in nest of Lehtinen, 1987 Zerconidae Zercon curiosus Trägårdh, 1910 Leptothorax spp. in nest of Lehtinen, 1987 Zerconidae Zercon curiosus Trägårdh, 1910 Myrmica spp. in nest of Lehtinen, 1987 Zerconidae Zercon lindrothi Lundqvist & Johnston, 1986 Formica fusca in nest of Lehtinen, 1987 Zerconidae Zercon lindrothi Lundqvist & Johnston, 1986 Lasius spp. in nest of Lehtinen, 1987 Zerconidae Zercon nr. spatulatus (C.L. Koch, 1939) Camponotus ligniperdus in nest of Lehtinen, 1987 Zerconidae Zercon nr. spatulatus (C.L. Koch, 1939) Lasius flavus in nest of Lehtinen, 1987 Zerconidae Zercon nr. spatulatus (C.L. Koch, 1939) Lasius niger in nest of Lehtinen, 1987 Zerconidae Zercon solenites Haarløv, 1942 Formica in nest of Lehtinen, 1987 Zerconidae Zercon spatulatus (C.L. Koch, 1939) unknown (many ants) in nest of Lehtinen, 1987

HIGHER MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE LEVEL Microdispidae Archidispus calcarifer Mahunka, 1977 Labidus praedator in nest of Mahunka, 1977b Microdispidae Archidispus repus Mahunka, 1977 Labidus praedator in nest of Mahunka, 1977b Microdispidae Archidispus repus Mahunka, 1977 Nomamyrmex esenbeckii in nest of Mahunka, 1977b Microdispidae Archidispus willmanni Mahunka, 1977 Neivamyrmex pilosus in nest of Mahunka, 1977a Microdispidae Caesarodispus gaius Mahunka, 1977c Myrmica sabuleti in nest of Mahunka, 1977c Khaustov & Microdispidae Caesarodispus klepzigi Khaustov & Moser, 2008 Solenopsis invicta in nest of Moser, 2008 Microdispidae Glyphidomastax rettenmeyeri Cross, 1965 Neivamyrmex nigrescens phoretic on ants Cross, 1965 Microdispidae Glyphidomastax rettenmeyeri Cross, 1965 Neivamyrmex opacithorax phoretic on ants Cross, 1965 Microdispidae Glyphidomastax rettenmeyeri Cross, 1965 Eciton burchellii in nest of Mahunka, 1977b Microdispidae Microdispus bacchaliensis Mahunka, 1977 Eciton hamatum in nest of Mahunka, 1977b Microdispidae Microdispus obtegitus Mahunka, 1977 Nomamyrmex esenbeckii in nest of Mahunka, 1977b 170 Microdispidae Myrmecodispus dorylinis Cross, 1965 Eciton burchellii phoretic on ants Cross, 1965

Microdispidae Myrmecodispus dorylinis Cross, 1965 Eciton vagans phoretic on ants Cross, 1965 Microdispidae Perperipes ornithocephala Cross, 1965 Eciton hamatum with larvae Cross, 1965 Microdispidae Perperipes ornithocephala Cross, 1965 Neivamyrmex gibbatus in nest of Mahunka, 1977a Microdispidae Perperipes ornithocephala Cross, 1965 Eciton burchellii in nest of Mahunka, 1977b Microdispidae Perperipes ornithocephala Cross, 1965 Labidus coecus in nest of Mahunka, 1977b Microdispidae Pygmodispus dorylini Mahunka, 1977 Eciton vagans in nest of Mahunka, 1977a Microdispidae Reductodispus rettenmeyerorum Mahunka, 1977 Labidus praedator in nest of Mahunka, 1977a Microdispidae Rettenmeyerella petropolitana solenifera Mahunka, 1977 Eciton hamatum in nest of Mahunka, 1977a Microdispidae Rettenmeyerella petropolitana solenifera Mahunka, 1977 Neivamyrmex pilosus mexicanus in nest of Mahunka, 1977a Microdispidae Unguidispus stammeri Mahunka, 1970c Formica hamatum in nest of Mahunka, 1970c Pyemotidae Pygmephorus myrmecophilus Mahunka, 1965 unknown (ants) sp. in nest of Mahunka, 1965 Pyemotidae Pygmephorus samsinaki Mahunka, 1967 Formica rufa in nest of Mahunka, 1967 Pyemotidae Siteroptes bohemicus Mahunka, 1967 Formica rufa in nest of Mahunka, 1967 Continued Table 30. Heterostigmatina and their Associated Ants from Literature.

Table 30 continued HIGHER MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE LEVEL Pyemotidae Siteroptes insignis Mahunka, 1968 Formica sp. in nest of Mahunka, 1968 Pygmephoridae Acinogaster (Acinogaster) crassisetosus Ross & Cross, 1979 Eciton burchellii phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster (Acinogaster) crassisetosus Ross & Cross, 1979 Eciton dulcium phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster (Acinogaster) crassisetosus Ross & Cross, 1979 Neivamyrmex pilosus mexicanus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster (Acinogaster) crassisetosus Ross & Cross, 1979 Eciton rapax phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster (Acinogaster) crassisetosus Ross & Cross, 1979 Labidus coecus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster (Acinogaster) crassisetosus Ross & Cross, 1979 Labidus praedator phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster (Acinogaster) crassisetosus Ross & Cross, 1979 Neivamyrmex adnepos phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster (Acinogaster) crassisetosus Ross & Cross, 1979 Neivamyrmex cristatus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster (Acinogaster) crassisetosus Ross & Cross, 1979 Neivamyrmex diana phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster crassisetosus Ross & Cross, 1979 Neivamyrmex gibbatus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster crassisetosus Ross & Cross, 1979 Neivamyrmex gradualis phoretic on ants Ross & Cross, 1979

171 Pygmephoridae Acinogaster crassisetosus Ross & Cross, 1979 Neivamyrmex legionis phoretic on ants Ross & Cross, 1979

Pygmephoridae Acinogaster crassisetosus Ross & Cross, 1979 Neivamyrmex pilosus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster crassisetosus Ross & Cross, 1979 Nomamyrmex esenbeckii phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster crassisetosus Ross & Cross, 1979 Nomamyrmex hartigii phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster gladiatus Ross & Cross, 1979 Labidus coecus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster gladiatus Ross & Cross, 1979 Labidus praedator phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster kansensis Ross & Cross, 1979 Eciton burchellii phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster kansensis Ross & Cross, 1979 Labidus coecus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster kansensis Ross & Cross, 1979 Labidus praedator phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster kansensis Ross & Cross, 1979 Neivamyrmex carolinensis phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster kansensis Ross & Cross, 1979 Neivamyrmex gibbatus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster kansensis Ross & Cross, 1979 Neivamyrmex moseri phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster kansensis Ross & Cross, 1979 Neivamyrmex nigrescens phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster kansensis Ross & Cross, 1979 Neivamyrmex opacithorax phoretic on ants Ross & Cross, 1979 Continued

Table 30 continued HIGHER MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE LEVEL Pygmephoridae Acinogaster kansensis Ross & Cross, 1979 Neivamyrmex pauxillus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster kansensis Ross & Cross, 1979 Neivamyrmex pilosus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster marianae Cross, 1965 Eciton burchellii phoretic on ants Cross, 1965 Pygmephoridae Acinogaster marianae Cross, 1965 Eciton hamatum phoretic on ants Cross, 1965 Pygmephoridae Acinogaster marianae Cross, 1965 Eciton burchellii phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster marianae Cross, 1965 Eciton hamatum phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster marianae Cross, 1965 Eciton lucanoides phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster marianae Cross, 1965 Eciton rapax phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster obovatus Ross & Cross, 1979 Neivamyrmex moseri phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster obovatus Ross & Cross, 1979 Neivamyrmex pauxillus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Eciton burchellii phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Eciton dulcium phoretic on ants Ross & Cross, 1979

172 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Eciton hamatum phoretic on ants Ross & Cross, 1979

Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Eciton lucanoides phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Eciton mexicanum phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Eciton rapax phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Eciton vagans phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Labidus coecus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Labidus praedator phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Neivamyrmex cristatus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Neivamyrmex gradualis phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Neivamyrmex legionis phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Nomamyrmex esenbeckii phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster panamanus Ross & Cross, 1979 Nomamyrmex hartigii phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster parvisetosus Ross & Cross, 1979 Labidus coecus phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster parvisetosus Ross & Cross, 1979 Neivamyrmex legionis phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster quadridens Cross, 1974 (Anomma) sp. phoretic on ants Cross, 1974 Continued

Table 30 continued HIGHER MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE LEVEL Pygmephoridae Acinogaster quadridens Cross, 1974 sp. phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster triangularis Ross & Cross, 1979 Neivamyrmex moseri phoretic on ants Ross & Cross, 1979 Pygmephoridae Acinogaster triangularis Ross & Cross, 1979 Neivamyrmex pauxillus phoretic on ants Ross & Cross, 1979 Pygmephoridae Bakerdania kielczewskii Metwali, 1981 Myrmica rubra in nest of Metwali, 1981 Pygmephoridae Petalomium aleinikovae (Sevastianov, 1967) Myrmica ruginodis phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium aleinikovae (Sevastianov, 1967) Lasius flavus phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium brevisetum Khaustov, 2005 Formica gagates phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium carelitschense (Sevastianov, 1967) Lasius alienus phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium carelitschense (Sevastianov, 1967) Lasius flavus phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium carelitschense (Sevastianov, 1967) Lasius niger phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium carelitschense (Sevastianov, 1967) Myrmica ruginodis phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium carelitschense Mahunka, 1977c Lasius niger in nest of Mahunka, 1977c

173 Pygmephoridae Petalomium carelitschense Mahunka, 1977c Lasius umbratus in nest of Mahunka, 1977c

Pygmephoridae Petalomium chaetosum (Krczal, 1959) Lasius umbratus in nest of Mahunka, 1977c Pygmephoridae Petalomium fimbrisetum Ebermann & Rack, 1982 Lasius flavus fungus in nest Ebermann & Rack, 1982 Pygmephoridae Petalomium fimbrisetum Ebermann, Rack, 1982 Lasius flavus phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium formicarum (Berlese, 1903) Formica cunicularia phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium formicarum (Berlese, 1903) Formica gagates phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium genavensium Mahunka, 1977c Mymica ruginodis in nest of Mahunka, 1977c Pygmephoridae Petalomium gottrauxi Mahunka, 1977 Camponotus aethiops phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium gottrauxi Mahunka, 1977 Myrmica ruginodis phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium gottrauxi Mahunka, 1977c Mymica ruginodis in nest of Mahunka, 1977c Pygmephoridae Petalomium heterotrichus Mahunka, 1970c Camponotus vagus in nest of Mahunka, 1970c Pygmephoridae Petalomium hofstetteri Khaustov & Moser, 2008 Solenopsis invicta in nest of Khaustov & Moser, 2008 Pygmephoridae Petalomium myrmecophilum (Mahunka, 1965) Lasius umbratus in nest of Mahunka, 1977c Pygmephoridae Petalomium nataliae (Sev., 1967) Lasius niger in nest of Mahunka, 1977c Pygmephoridae Petalomium podolicum (Sev., 1967) Lasius niger in nest of Mahunka, 1977c Continued

Table 30 continued HIGHER MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE LEVEL Pygmephoridae Petalomium podolicus Mahunka, 1970 Formica sp. in nest of Mahunka, 1970b Pygmephoridae Petalomium podolicus Mahunka, 1970 Lasius fuliginosus in nest of Mahunka, 1970b Pygmephoridae Petalomium pseudomyrmecophilum Mahunka, 1970 Lasius umbratus in nest of Mahunka, 1977c Pygmephoridae Petalomium pseudomyrmecophylus Mah., 1970 unknown (ants) in ant hill Mahunka, 1970b Pygmephoridae Petalomium rackae Metwali, 1981 Myrmica rubra in nest of Metwali, 1981 Pygmephoridae Petalomium rackae Metwali, 1981 Myrmica ruginodis in nest of Metwali, 1981 Pygmephoridae Petalomium rarum (Sevastianov, 1967) Formica gagates phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium sawtschuki (Sevastianov, 1967) Formica rufa in nest of Mahunka, 1970b Pygmephoridae Petalomium scyphicum (Sevastianov, 1967) Lasius alienus phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium scyphicum (Sevastianov, 1967) Lasius fuliginosus phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium scyphicum (Sevastianov, 1967) Lasius niger phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium scyphicum (Sev., 1967) Lasius niger in nest of Mahunka, 1977c

174 Pygmephoridae Petalomium scyphicus (Sevastianov, 1967) Lasius sp. in nest of Mahunka, 1970b

Pygmephoridae Petalomium tauricum Khaustov, 2005 Formica gagates phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium tauricum Khaustov, 2005 Lasius alienus phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium tothi Mahunka et Zaki, 1984 Lasius flavus phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium tumidisetosum (=genavensium) (Wilmann, 1951) Lasius niger phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium tumidisetosum (=genavensium) (Wilmann, 1951) Tetramorium sp. phoretic on ants Khaustov, 2005 Pygmephoridae Petalomium ucrainicum (Sev., 1967) Formica sanguinea in nest of Mahunka, 1977c Pygmephoridae Petalomium ucrainicus (Sevastianov, 1967) unknown (ants) in nest of Mahunka, 1970b Pygmephoridae Pseudopygmephorus haarloevi Cross, 1965 Lasius flavus in nest of Cross, 1965 Pygmephoridae Pseudopygmephorus willmanni Cross, 1965 Lasius flavus in nest of Cross, 1965 Pygmephoridae Pygmephorus spp. (7) n/a Lasius fuliginosus in nest of Sevastianov, 1965 Scutacaridae Acinogaster panamanus Ross & Cross, 1979 Cheliomyrmex morosus phoretic on ants Berghoff & Franks, 2007 Scutacaridae Disparipes nudus Berl. Lasius fuliginosus in nest of Wasmann, 1899 Continued

Table 30 continued HIGHER MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE LEVEL Scutacaridae Disparipes sp. n/a Formica rufibarbis phoretic on ants Wasmann, 1899 Scutacaridae Disparipes sp. n/a Tapinoma erraticum on eggs Wasmann, 1899 Scutacaridae Imparipes atta Delfinado & Baker, 1976 Atta cephalotes phoretic on ants Delfinado & Baker, 1976 Scutacaridae Imparipes atypicus Karafiat, 1959 Formica rufa phoretic on ants Karafiat, 1959 Scutacaridae Imparipes brevitarsus Ebermann, 1981 Lasius flavus in nest of Ebermann, 1981 Scutacaridae Imparipes cassovaricus Mahunka, 1970 Neivamyrmex pilosus in nest of Mahunka, 1977a Berlese, 1903 in Scutacaridae Imparipes cf. hystricinus Mahunka, 1967 Formica rufa in nest of Mahunka, 1967 Scutacaridae Imparipes comatosimilis Metwali, 1981 Myrmica rubra in nest of Metwali, 1981 Scutacaridae Imparipes comatosimilis Metwali, 1981 Myrmica ruginodis in nest of Metwali, 1981 Scutacaridae Imparipes comatus Mahunka, 1970c Formica sp. in nest of Mahunka, 1970c

175 Scutacaridae Imparipes comatus Mahunka, 1970 Tapinoma erraticum in nest of Mahunka, 1977c Scutacaridae Imparipes compensatus Mahunka, 1977 Labidus praedator in nest of Mahunka, 1977b

Scutacaridae Imparipes compensatus Mahunka, 1977 Neivamyrmex pilosus in nest of Mahunka, 1977b Scutacaridae Imparipes convexus Mahunka, 1977 Labidus praedator in nest of Mahunka, 1977a Scutacaridae Imparipes crudelatus Mahunka, 1977 Eciton burchellii in nest of Mahunka, 1977b Scutacaridae Imparipes crudelatus Mahunka, 1977 Eciton hamatum in nest of Mahunka, 1977b Scutacaridae Imparipes degenerans Berlese, 1903 unknown (ants) phoretic on ants Karafiat, 1959 Scutacaridae Imparipes dimidiatus Mahunka, 1977 Neivamyrmex pilosus in nest of Mahunka, 1977a Scutacaridae Imparipes egisetus Mahunka, 1977 Neivamyrmex pilosus mexicanus in nest of Mahunka, 1977b Scutacaridae Imparipes egisetus Mahunka, 1977 Neivamyrmex gibbatus in nest of Mahunka, 1977b Scutacaridae Imparipes elzingai Mahunka, 1977 Neivamyrmex cristatus in nest of Mahunka, 1977a Continued

Table 30 continued HIGHER MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE LEVEL Scutacaridae Imparipes histricinus Berl. Lasius fuliginosus in nest of Sevastianov, 1965 Scutacaridae Imparipes hystricinus Berlese, 1903 unknown (ants) phoretic on ants Karafiat, 1959 Scutacaridae Imparipes hystricinus Berlese, 1903 Lasius fuliginosus in nest of Štorkán, 1940 Scutacaridae Imparipes hystricinus Berlese, 1903 Lasius umbratus in nest of Štorkán, 1940 Scutacaridae Imparipes hystricinus Berlese, 1903 Tetramorium caespitum in nest of Štorkán, 1940 Scutacaridae Imparipes hystricinus Berlese Lasius fuliginosus in nest of Vitzthum, 1919 Scutacaridae Imparipes ignotus Khaustov & Chydyrov, 2004 Messor sp. in nest of Khaustov & Chydyrov, 2004 Scutacaridae Imparipes imitatus Mahunka, 1977 Eciton rapax in nest of Mahunka, 1977b Scutacaridae Imparipes intermissus Karafiat, 1959 Lasius fuliginosus phoretic on ants Karafiat, 1959

176 Scutacaridae Imparipes kataglyphi Khaustov & Chydyrov, 2004 Cataglyphus emeryi in nest of Khaustov & Chydyrov, 2004

Scutacaridae Imparipes kugitangensis Khaustov & Chydyrov, 2004 Pheidole pallidula in nest of Khaustov & Chydyrov, 2004 Scutacaridae Imparipes lapillatus Mahunka, 1977 Cheliomyrmex morosus phoretic on ants Berghoff & Franks, 2007 Scutacaridae Imparipes lapillatus Mahunka, 1977 Eciton rapax in nest of Mahunka, 1977b Scutacaridae Imparipes latispinus Mahunka, 1970 Pogonomyrmex occidentalis in nest of Mahunka, 1970a Scutacaridae Imparipes louisianae Ebermann & Moser, 2008 Solenopsis invicta phoretic on ants Ebermann & Moser, 2008 Scutacaridae Imparipes marianae Mahunka, 1977 Eciton drepanophorum in nest of Mahunka, 1977a Scutacaridae Imparipes marianae Mahunka, 1977 Eciton burchellii in nest of Mahunka, 1977a Scutacaridae Imparipes mexicanus Mahunka, 1982 apiculatum phoretic on ants Mahunka, 1982 Scutacaridae Imparipes moralesi Mahunka, 1977 Neivamyrmex cristatus in nest of Mahunka, 1977a Scutacaridae Imparipes moricus Mahunka, 1977 Neivamyrmex carolinensis in nest of Mahunka, 1977b Scutacaridae Imparipes nabilatus Mahunka, 1977 Labidus spininodis in nest of Mahunka, 1977b Continued

Table 30 continued HIGHER MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE LEVEL Scutacaridae Imparipes obsoletus Rack, 1966 Lasius flavus in nest of Ebermann, 1979 Scutacaridae Imparipes obsoletus Rack, 1966 Lasius niger in nest of Ebermann, 1979 Scutacaridae Imparipes obsoletus Rack, 1966 Lasius umbratus in nest of Mahunka, 1977c Scutacaridae Imparipes obsoletus Rack, 1966 Lasius umbratus in nest of Mahunka, 1977c Scutacaridae Imparipes odiosus Mahunka, 1977 Neivamyrmex adnepos in nest of Mahunka, 1977b Scutacaridae Imparipes officius Mahunka, 1977 Neivamyrmex pilosus mexicanus in nest of Mahunka, 1977b Scutacaridae Imparipes opusculus Mahunka, 1977 Eciton burchellii in nest of Mahunka, 1977b Scutacaridae Imparipes opusculus Mahunka, 1977 Eciton hamatum in nest of Mahunka, 1977b Scutacaridae Imparipes opusculus Mahunka, 1977 Eciton lucanoides in nest of Mahunka, 1977b

177 Scutacaridae Imparipes opusculus Mahunka, 1977 Labidus coecus in nest of Mahunka, 1977b

Scutacaridae Imparipes parapicolosimilis Metwali, 1981 Tetramorium caespitum in nest of Metwali, 1981 Scutacaridae Imparipes pennatus Karafiat, 1959 Formica rufa phoretic on ants Karafiat, 1959 Scutacaridae Imparipes pennatus Kar. 1959 unknown (ants) in nest of Mahunka, 1970b Scutacaridae Imparipes pennatus Karafiat, 1959 Tapinoma erraticum in nest of Mahunka, 1977c Khaustov & Chydyrov, Scutacaridae Imparipes placidus Khaustov & Chydyrov, 2004 Messor excursionis in nest of 2004 Scutacaridae Imparipes rationis Mahunka, 1977 Eciton vagans in nest of Mahunka, 1977a Scutacaridae Imparipes rationis Mahunka, 1977 Labidus praedator in nest of Mahunka, 1977a Scutacaridae Imparipes rationis Mahunka, 1977 Neivamyrmex cristatus in nest of Mahunka, 1977a Scutacaridae Imparipes recisus Mahunka, 1977 Eciton hamatum in nest of Mahunka, 1977b Scutacaridae Imparipes rectangulatus Mahunka, 1977 Eciton hamatum in nest of Mahunka, 1977a Continued

Table 30 continued HIGHER MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE LEVEL Scutacaridae Imparipes rectangulatus Mahunka, 1977 sp. in nest of Mahunka, 1977a Scutacaridae Imparipes rectangulatus Mahunka, 1977 Neivamyrmex cristatus in nest of Mahunka, 1977a Scutacaridae Imparipes retrosus Mahunka, 1977 Neivamyrmex cristatus in nest of Mahunka, 1977a Scutacaridae Imparipes rettenmeyeri Mahunka, 1977 Labidus praedator in nest of Mahunka, 1977a Scutacaridae Imparipes robustus Karafiat, 1959 Lasius fuliginosus phoretic on ants Karafiat, 1959 Scutacaridae Imparipes robustus Karafiat, 1959 Formica rufa in nest of Mahunka, 1967 Scutacaridae Imparipes saevus Mahunka, 1977 Neivamyrmex pilosus in nest of Mahunka, 1977b Scutacaridae Imparipes suboletus Mahunka, 1977 Eciton burchellii in nest of Mahunka, 1977b Mahunka & Mahunka, Scutacaridae Imparipes vasarhelyii Mahunka & Mahunka, 1980 unknown (ants) in nest of 1980 178 Scutacaridae Lophodispus latus Kurosa, 1972 Lasius niger in nest of Ebermann, 1979

Scutacaridae Scutacaropsis problematicus Mahunka, 1977 Eciton burchellii in nest of Mahunka, 1977a Scutacaridae Scutacaropsis problematicus Mahunka, 1977 Neivamyrmex pilosus mexicanus in nest of Mahunka, 1977a Scutacaridae Scutacaropsis scutacaroides Mahunka, 1977 Eciton lucanoides? in nest of Mahunka, 1977a Scutacaridae Scutacarus andrassy Mahunka Solenopsis invicta phoretic on ants Ebermann & Moser, 2008 Scutacaridae Scutacarus athiashenriotae Mahunka, 1982 Liometopum apiculatum phoretic on ants Mahunka, 1982 Scutacaridae Scutacarus bursula (Berlese, 1903) unknown (ants) phoretic on ants Karafiat, 1959 Mahunka & Mahunka, Scutacaridae Scutacarus carsticus Mahunka & Mahunka, 1980 unknown (ants) in nest of 1980 Scutacaridae Scutacarus cf. deserticolus Mahunka Solenopsis invicta phoretic on ants Ebermann & Moser, 2008 Scutacaridae Scutacarus chadabi Mahunka, 1977 Nomamyrmes esenbeckii in nest of Mahunka, 1977a Scutacaridae Scutacarus concinnus Mah., 1964 unknown (ants) in ant hill Mahunka, 1970b Continued

Table 30 continued HIGHER MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE LEVEL Scutacaridae Scutacarus ellipticus Karafiat, 1959 Myrmica rubra phoretic on ants Karafiat, 1959 Scutacaridae Scutacarus exspectatus Karafiat, 1959 Lasius niger in nest of Karafiat, 1959 Scutacaridae Scutacarus flexisetosus Karafiat, 1959 Lasius fuliginosus phoretic on ants Karafiat, 1959 Scutacaridae Scutacarus flexisetosus Karaf. Lasius fuliginosus in nest of Sevastianov, 1965 Scutacaridae Scutacarus gratus Karafiat, 1959 Lasius fuliginosus phoretic on ants Karafiat, 1959 Scutacaridae Scutacarus gratus Karaf. Lasius fuliginosus in nest of Sevastianov, 1965 Scutacaridae Scutacarus hauseri Mahunka, 1977 Lasius niger in nest of Ebermann, 1979 Scutacaridae Scutacarus hauseri Mahunka, 1977c Myrmica sabuleti in nest of Mahunka, 1977c Scutacaridae Scutacarus kassaii Mahunka, 1965 Camponotus vagus in nest of Mahunka, 1967

179 Scutacaridae Scutacarus latifrons Mah., 1964 unknown (ants) in nest of Mahunka, 1970b

Scutacaridae Scutacarus longisetus (Berlese, 1904) Lasius niger in nest of Ebermann, 1979 Scutacaridae Scutacarus longisetus (Berlese, 1903) Lasius sp. phoretic on ants Karafiat, 1959 Scutacaridae Scutacarus longisetus (Berl., 1903) Lasius umbratus in nest of Mahunka, 1977c Scutacaridae Scutacarus myrmecophilus Metwali, 1981 Myrmica rubra in nest of Metwali, 1981 Scutacaridae Scutacarus myrmecophilus Metwali, 1981 Myrmica ruginodis in nest of Metwali, 1981 Scutacaridae Scutacarus nanus Ebermann & Moser, 2008 Solenopsis invicta phoretic on ants Ebermann & Moser, 2008 Scutacaridae Scutacarus omittatus Mahunka, 1977 Eciton hamatum in nest of Mahunka, 1977a Scutacaridae Scutacarus ovoideosimilis Metwali, 1981 Myrmica rubra in nest of Metwali, 1981 Scutacaridae Scutacarus ovoideosimilis Metwali, 1981 Myrmica ruginodis in nest of Metwali, 1981 Continued

Table 30 continued HIGHER MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE LEVEL Scutacaridae Scutacarus ovoideus Karafiat, 1959 Lasius niger in nest of Ebermann, 1979 Scutacaridae Scutacarus ovoideus Karafiat, 1959 Myrmica ruginodis in nest of Ebermann, 1979 Scutacaridae Scutacarus ovoideus Karafiat, 1959 unknown (ants) in nest of Karafiat, 1959 Scutacaridae Scutacarus ovoideus Karafiat, 1959 Lasius niger in nest of Mahunka, 1977c Scutacaridae Scutacarus ovoideus Karafiat, 1959 Myrmica ruginodis in nest of Mahunka, 1977c Scutacaridae Scutacarus palustris Mahunka, 1968 Formica in nest of Mahunka, 1968 Scutacaridae Scutacarus pseudocomus Mahunka, 1977 Eciton burchellii in nest of Mahunka, 1977b Scutacaridae Scutacarus rettenmeyeri Mahunka, 1977 Myrmicinae sp. in nest of Mahunka, 1977a Scutacaridae Scutacarus rotundatus (Berlese, 1903) Formica rufa in nest of Štorkán, 1940 Scutacaridae Scutacarus rotundatus (Berlese, 1903) Lasius niger in nest of Štorkán, 1940 Scutacaridae Scutacarus rotundulus Khaustov & Chydyrov, 2004 Camponotus fedtschenkoi in nest of Khaustov & Chydyrov, 2004 180 Scutacaridae Scutacarus rotundus (Berlese, 1903) Formica fusca phoretic on ants Karafiat, 1959

Scutacaridae Scutacarus rotundus (Berlese, 1903) Lasius niger phoretic on ants Karafiat, 1959 Scutacaridae Scutacarus rotundus Berl, 1903 Formica rufa in nest of Mahunka, 1970b Scutacaridae Scutacarus sabinaesimilis Khaustov & Chydyrov, 2004 Tapinoma simrothi in nest of Khaustov & Chydyrov, 2004 Scutacaridae Scutacarus setarus Mahunka, 1977 Neivamyrmex pilosus mexicanus in nest of Mahunka, 1977b Scutacaridae Scutacarus sp. n/a Lasius fuliginosus in nest of Sevastianov, 1965 Scutacaridae Scutacarus stammeri Karafiat, 1959 Formica fusca in nest of Karafiat, 1959 Scutacaridae Scutacarus subcomosus Mahunka, 1970 Lasius umbratus in nest of Mahunka, 1977c Scutacaridae Scutacarus subellipticus Delfinado & Baker, 1976 unknown (ants) in ant hill Delfinado & Baker, 1976 Scutacaridae scutacarus subquadratus Khaustov & Chydyrov, 2004 Tetramorium schneideri in nest of Khaustov & Chydyrov, 2004 Scutacaridae Scutacarus subterraneus (Oudemans, 1913) Lasius fuliginosus in nest of Ebermann, 1979 Continued

Table 30 continued HIGHER MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE LEVEL Scutacaridae Scutacarus subterraneus (Oudemans, 1913) Lasius niger in nest of Ebermann, 1979 Scutacaridae Scutacarus subterraneus n/a Lasius fuliginosus in nest of Sevastianov, 1965 Scutacaridae Scutacarus subterraneus (Oudemans, 1913) Lasius fuliginosus in nest of Štorkán, 1940 Scutacaridae Scutacarus subterraneus (Oudemans, 1913) Lasius niger in nest of Štorkán, 1940 Oudemans, 1913, Štorkán, Scutacaridae Scutacarus subterraneus spinosus 1936 unknown (ants) in nest of Karafiat, 1959 Scutacaridae Scutacarus tackei ellipticus Karafiat, 1959 Lasius niger in nest of Mahunka, 1977c Scutacaridae Scutacarus tackei ellipticus Karafiat, 1959 Myrmica ruginodis in nest of Mahunka, 1977c Scutacaridae Scutacarus takei Willman, 1942 Myrmica rubra in nest of Rack, 1964 Scutacaridae Scutacarus terrenus Delfinado & Baker, 1976 unknown (ants) in ant hill Delfinado & Baker, 1976 Scutacaridae Scutacarus tertius Ebermann & Moser, 2008 Solenopsis invicta phoretic on ants Ebermann & Moser, 2008 Scutacaridae Scutacarus transfusionis Mahunka & Mahunka, 1980 unknown (ants) in nest of Mahunka & Mahunka, 1980 181 Scutacaridae Scutacarus wranoskyi Mahunka, 1970 Pogonomyrmex occidentalis in nest of Mahunka, 1970a

Scutacaridae Thaumatopelvis minutissimus Mahunka, 1977 Labidus praedator in nest of Mahunka, 1977a Scutacaridae Thaumatopelvis minutissimus Mahunka, 1977 Neivamyrmex cristatus in nest of Mahunka, 1977a Scutacaridae Thaumatopelvis rugosus Mahunka, 1977 Labidus praedator in nest of Mahunka, 1977a Scutacaridae undetermined sp. n/a Solenopsis richteri in lab colony Collins & Markin, 1971 fed from ant Tarsonemidae Disparipedidae sp. n/a Solenopsis geminata mouth and anus Travis, 1941 Tarsonemidae Tarsonemoides sp. n/a Lasius fuliginosus in nest of Sevastianov, 1965 Tarsonemidae gladifer Mahunka, 1970c unknown (ants) in nest of Mahunka, 1970c Tarsonemidae Tarsonemus sp. n/a Lasius fuliginosus in nest of Sevastianov, 1965 Tarsonemina Parapygmephorus folliger Mahunka & Mahunka, 1980 unknown (ants) in nest of Mahunka & Mahunka, 1980

HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Acaridae Acotyledon mystax Mahunka, 1978 Eciton burchellii in nest of Mahunka, 1978 Acaridae Acotyledon mystax Mahunka, 1978 Eciton mexicanum in nest of Mahunka, 1978 Acaridae Acotyledon solenopsidis Samsinak, 1960 Solenopsis fugax in nest of Samsinak, 1960 Acaridae Armacarus anniversarius Mahunka, 1979 Labidus praedator in nest of Mahunka, 1979 Acaridae Caloglyphus armatus Mahunka, 1979 Eciton drepanophorum in nest of Mahunka, 1979 Acaridae Caloglyphus armatus Mahunka, 1979 Labidus coecus in nest of Mahunka, 1979 Acaridae Caloglyphus capitatus Mahunka, 1979 Neivamyrmex carolinensis in nest of Mahunka, 1979 Acaridae Caloglyphus dorylini Mahunka, 1979 Labidus coecus in nest of Mahunka, 1979 Acaridae Caloglyphus vitzthumi Mahunka, 1979 Neivamyrmex rugulosus in nest of Mahunka, 1979 Acaridae Cosmoglyphus elzingai Mahunka, 1979 Labidus coecus in nest of Mahunka, 1979

182 Acaridae Cosmoglyphus longisetus Mahunka, 1979 Neivamyrmex carolinensis in nest of Mahunka, 1979

Acaridae Dorylacarus rugosus Mahunka, 1979 Labidus praedator in nest of Mahunka, 1979 Acaridae Dorylacarus rugosus Mahunka, 1979 Nomamyrmex esenbeckii in nest of Mahunka, 1979 Acaridae Forcellinia bipunctata Mahunka, 1978 Eciton lucanoides in nest of Fain, 1987 Acaridae Forcellinia bipunctata Mahunka, 1978 Eciton rapax in nest of Fain, 1987 Acaridae Forcellinia bipunctata Mahunka, 1978 Eciton lucanoides in nest of Mahunka, 1978 Acaridae Forcellinia bipunctata Mahunka, 1978 Eciton rapax in nest of Mahunka, 1978 Estebanes-Gonzalez, Acaridae Forcellinia faini Delfinado-Baker & Baker, 1989 Liometopum luctuosum artificial nest 2000 Acaridae Forcellinia fuliginosi Türk & Türk, 1957 Lasius fuliginosus in nest of Fain, 1987 Acaridae Forcellinia fuliginosi Türk. Lasius fuliginosus in nest of Sevastianov, 1965 Continued Table 31. Astigmata and their Associated Ants from Literature.

Table 31 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Acaridae Forcellinia galleriella Womersley, 1963 Linepithema humile in nest of Fain, 1987 Acaridae Forcellinia laesionis Mahunka, 1979 Labidus coecus in nest of Fain, 1987 Acaridae Forcellinia laesionis Mahunka, 1979 Labidus coecus in nest of Mahunka, 1979 Acaridae Forcellinia mystax (Mahunka, 1978) Eciton burchellii in nest of Fain, 1987 Acaridae Forcellinia mystax (Mahunka, 1978) Neivamyrmex pilosus mexicanus in nest of Fain, 1987 Acaridae Forcellinia rugosa (Mahunka, 1979) Labidus praedator in nest of Fain, 1987 Acaridae Forcellinia rugosa (Mahunka, 1979) Nomamyrmex esenbeckii in nest of Fain, 1987 Acaridae Forcellinia spp. (2) n/a Lasius fuliginosus in nest of Sevastianov, 1965 Acaridae Forcellinia vandergooti (Oudemans, 1917) thoracicus in nest of Fain, 1987

183 Acaridae Forcellinia vandergooti Oudeman, 1917 Anoplolepis gracilipes in nest of Fain, 1987 Acaridae Forcellinia wasmanni (Moniez, 1892) Camponotus ligniperdus in nest of Fain, 1987 Acaridae Forcellinia wasmanni (Moniez, 1892) Formica spp. in nest of Fain, 1987 Acaridae Forcellinia wasmanni (Moniez, 1892) Lasius fuliginosus in nest of Fain, 1987 Acaridae Forcellinia wasmanni Moniez Formica fusca in nest of Hermann et al., 1970 Acaridae Forcellinia wasmanni Moniez Lasius umbratus in nest of Hermann et al., 1970 Acaridae Forcellinia wasmanni (Moniez, 1892) Camponotus ligniperdus phoretic on ants Mahunka, 1961 Acaridae Forcellinia wasmanni (Mon.) Lasius fuliginosus in nest of Sevastianov, 1965 Acaridae Forcellinia wasmanni (Moniez, 1892) Formica fusca in nest of Štorkán, 1940 Acaridae Forcellinia wasmanni (Moniez, 1892) Formica rufa in nest of Štorkán, 1940 Acaridae Forcellinia wasmanni (Moniez, 1892) Formica sanguinea in nest of Štorkán, 1940 Continued

Table 31 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Acaridae Forcellinia wasmanni (Moniez, 1892) Lasius alienus in nest of Štorkán, 1940 Acaridae Forcellinia wasmanni (Moniez, 1892) Lasius fuliginosus in nest of Štorkán, 1940 Acaridae Forcellinia wasmanni (Moniez, 1892) Lasius niger in nest of Štorkán, 1940 Acaridae Forcellinia wasmanni (Moniez, 1892) Myrmica ruginodis in nest of Štorkán, 1940 Acaridae Forcellinia wasmanni (Moniez, 1892) Tetramorium caespitum in nest of Štorkán, 1940 Acaridae Froriepia heterotricha Mahunka, 1978 Labidus coecus in nest of Mahunka, 1978 Acaridae Froriepia vimariensis Vitzthum, 1919 Lasius fuliginosus in nest of Vitzthum, 1919 phoretic on thorax and Acaridae Garsaultia gigantonympha (Vitzt., 1920) Camponotus ligniperdus abdomen Mahunka, 1961 phoretic on thorax and Acaridae Garsaultia gigantonympha (Vitzt., 1920) Lasius fuliginosus abdomen Mahunka, 1961 184 Acaridae Garsaultia tetramorii (Türk & Türk, 1957) Tetramorium caespitum in nest of Mahunka, 1961

Fain & Chmielewski, Acaridae Lasioacarus nidicolus Kadzhaja & Sevastianov, 1967 Lasius niger in nest of 1987 Acaridae Monieziella corticalis Mich. Lasius fuliginosus in nest of Sevastianov, 1965 Acaridae Myrmolichus greimae Türk & Türk, 1957 Lasius fuliginosus feeding on fungus, nest walls Wurst, 2001 Acaridae Ocellacarus congretatus Mahunka, 1979 Labidus praedator in nest of Mahunka, 1979 Acaridae Ocellacarus Mahunka, 1979 Labidus praedator in nest of Mahunka, 1979 Acaridae Pinoglyphus cetus Mahunka, 1979 Labidus praedator in nest of Mahunka, 1979 Acaridae Pinoglyphus pilosus Mahunka, 1979 Labidus praedator in nest of Mahunka, 1979 Rettacarus Acaridae rettenmeyerorum Mahunka, 1979 Labidus coecus in nest of Mahunka, 1979 Acaridae Schwiebea schmuttereri Türk. Lasius fuliginosus in nest of Sevastianov, 1965 Acaridae Troupeauia sp. Zach. Lasius fuliginosus in nest of Sevastianov, 1965 Acaridae Tyrophagus brauni n/a Apterostigma sp. phoretic on ants Weber, 1972 Continued

Table 31 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Acaridae Tyrophagus brauni n/a Cyphomyrmex sp. phoretic on ants Weber, 1972 Acaridae Tyrophagus brauni n/a Mycocepurus sp. phoretic on ants Weber, 1972 Acaridae undetermined sp. n/a Solenopsis richteri in lab colony Collins & Markin, 1971 Conglanoetus Anoetidae ornatofoveolatus Mahunka, 1978 Labidus praedator in nest of Mahunka, 1978 Anoetidae Pelzneria afluctuosa Mahunka, 1978 Eciton burchellii in nest of Mahunka, 1978 Anoetidae Pteranoetus incertus Mahunka, 1978 Eciton burchellii in nest of Mahunka, 1978 Anoetidae Pteranoetus latus Mahunka, 1978 Neivamyrmex cristatus in nest of Mahunka, 1978 Anoetidae Pteranoetus latus Mahunka, 1978 unknown (ants) in nest of Mahunka, 1978 Anoetidae Pteranoetus rettenmeyeri Mahunka, 1978 Neivamyrmex cristatus in nest of Mahunka, 1978

185 Anoetidae Pteranoetus rettenmeyeri Mahunka, 1978 Neivamyrmex gradualis in nest of Mahunka, 1978

Anoetidae Pteranoetus rettenmeyeri Mahunka, 1978 Neivamyrmex gradualis in nest of Mahunka, 1978 Histiostomatidae Anoetus dolifer Mahunka, 1977c Myrmica ruginodis in nest of Mahunka, 1977c Histiostomatidae Anoetus dolifer Mahunka, 1977c Myrmica sabuleti in nest of Mahunka, 1977c Histiosoma = Histiostoma Histiostomatidae rostroserratum Megn. Formica rufa phoretic on ants Wasmann, 1899 Histiostomatidae Histiostoma adarcuale Mahunka, 1979 Neivamyrmex rugulosus in nest of Mahunka, 1979 Histiostomatidae Histiostoma ardarcuale Mahunka, 1979 Labidus coecus in nest of Mahunka, 1979 Histiostomatidae Histiostoma capillata Mahunka, 1978 Labidus praedator in nest of Mahunka, 1978 Histiostomatidae Histiostoma clavifer Mahunka, 1978 Eciton hamatum in nest of Mahunka, 1978 Histiostomatidae Histiostoma continuata Mahunka, 1978 Labidus sp. in nest of Mahunka, 1978 Histiostomatidae Histiostoma coronatum Mahunka, 1979 Neivamyrmex rugulosus in nest of Mahunka, 1979 Histiostomatidae Histiostoma diademata Mahunka, 1978 Neivamyrmex gibbatus in nest of Mahunka, 1978 Continued

Table 31 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Histiostomatidae Histiostoma duplex Mahunka, 1979 Eciton burchellii in nest of Mahunka, 1979 Histiostomatidae Histiostoma elzingai Mahunka, 1978 Eciton lucanoides in nest of Mahunka, 1978 Histiostomatidae Histiostoma feroniarum Duf. Lasius fuliginosus in nest of Sevastianov, 1965 Histiostomatidae Histiostoma fuliginosi Scheuch. Lasius fuliginosus in nest of Sevastianov, 1965 Histiostomatidae Histiostoma incurvata Mahunka, 1978 Eciton burchellii in nest of Mahunka, 1978 Histiostomatidae Histiostoma marianae Mahunka, 1979 Eciton burchellii in nest of Mahunka, 1979 Histiostomatidae Histiostoma moralesi Mahunka, 1978 Eciton hamatum in nest of Mahunka, 1978 Histiostomatidae Histiostoma perruptum Mahunka, 1979 Labidus praedator in nest of Mahunka, 1979 Histiostoma Histiostomatidae rettenmeyerorum Mahunka, 1978 Labidus praedator in nest of Mahunka, 1978

186 Histiostomatidae Histiostoma rimosa Mahunka, 1978 Labidus sp. in nest of Mahunka, 1978

Histiostomatidae Histiostoma rostrata Mahunka, 1978 Eciton hamatum in nest of Mahunka, 1978 Histiostoma rostro- Histiostomatidae serratum Mégn. Formica rufa in artificial nest Wasmann, 1897 Histiostoma rostro- Histiostomatidae serratum Mégn. Formica sanguinea in artificial nest Wasmann, 1897 Histiostomatidae Histiostoma scindita Mahunka, 1978 Eciton lucanoides in nest of Mahunka, 1978 Histiostomatidae Histiostoma sp. n/a Solenopsis invicta in nest of Sokolov, et al. 2003 Histiostomatidae Semianoetus banksi Mahunka, 1979 Neivamyrmex sumichrasti in nest of Mahunka, 1979 Lemanniellidae Lemanniella minotauri Wurst, E, 2001 Lasius brunneus fungus in nest Wurst, 2001 Lemanniellidae Lemanniella reducta Mahunka, 1977c Myrmica sabuleti in nest of Mahunka, 1977c Pinoglyphidae Pinoglyphus rettenmeyeri Mahunka, 1978 Labidus sp. in nest of Mahunka, 1978 Continued

Table 31 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Dermatophagoides arnoldii Sevastianov, 1965 Lasius fuliginosus in nest of Sevastianov, 1965 Tyroglyphidae Tyroglyphus ineaequalis Banks, 1916 Aphaenogaster longiceps in nest of Banks, 1916 Tyroglyphidae Tyroglyphus krameri Berl. Camponotus ligniperdus in nest of Moniez, 1892 Tyroglyphidae Tyroglyphus krameri Berl. Formica ? in nest of Moniez, 1892 Tyroglyphidae Tyroglyphus krameri Berl. Formica sanguinea in nest of Moniez, 1892 Tyroglyphidae Tyroglyphus krameri Berl. Lasius fuliginosus in nest of Moniez, 1892 Tyroglyphidae Tyroglyphus krameri Michael Camponotus ligniperdus in natural nest Wasmann, 1897 Tyroglyphidae Tyroglyphus krameri Michael Formica sanguinea in natural nest Wasmann, 1897 Tyroglyphidae Tyroglyphus krameri Michael Lasius fuliginosus in natural nest Wasmann, 1897

187 Tyroglyphidae Tyroglyphus megnini Haller Lasius mixtus in nest of Janet, C. M., 1902

Tyroglyphidae Tyroglyphus siro/infestans L., Berlese Formica sanguinea in artificial nest Wasmann, 1897 Tyroglyphidae Tyroglyphus wasmanni Mon. Formica fusca in nest of Donisthorpe, 1927 Tyroglyphidae Tyroglyphus wasmanni Mon. Lasius umbratus in nest of Donisthorpe, 1927 Tyroglyphidae Tyroglyphus wasmanni Mon. Myrmica rubra in nest of Donisthorpe, 1927 Tyroglyphidae Tyroglyphus wasmanni Mon. Myrmica ruginodis in nest of Donisthorpe, 1927 Tyroglyphidae Tyroglyphus wasmanni Moniez, 1892 Camponotus ligniperdus artificial nest Moniez, 1892 Tyroglyphidae Tyroglyphus wasmanni Moniez, 1892 Formica fusca artificial nest Moniez, 1892 Tyroglyphidae Tyroglyphus wasmanni Moniez, 1892 Formica sanguinea artificial nest Moniez, 1892 Tyroglyphidae Tyroglyphus wasmanni Moniez, 1892 Lasius fuliginosus in nest of Moniez, 1892 Tyroglyphidae Tyroglyphus wasmanni Moniez Camponotus ligniperdus in artificial nest Wasmann, 1897 Tyroglyphidae Tyroglyphus wasmanni Moniez Formica sanguinea in nest of Wasmann, 1897 Tyroglyphidae Tyroglyphus wasmanni Moniez Lasius fuliginosus in artificial nest Wasmann, 1897 Continued

Table 31 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Tyroglyphidae Tyroglyphus wasmanni Moniez Formica rufa in nest of Wasmann, 1899 Tyroglyphidae Tyroglyphus wasmanni Moniez Formica sanguinea in nest of Wasmann, 1899 Tyroglyphidae Tyroglyphus wasmanni Moniez Lasius fuliginosus in nest of Wasmann, 1899 Fain & Chmielewski, Tyroglyphidae Tyrophagus formicetorum Volgin, 1948 Formica rufa in nest of 1987 Tyroglyphidae Tyrophagus formicetorum Volgin, 1948 Formica rufa in nest of Fain, 1987 Tyroglyphidae Tyrophagus formicetorum Mahunka, 1978 Formica rufa in nest of Mahunka, 1978 Calvolia sp. n/a Lasius fuliginosus in nest of Sevastianov, 1965

188

HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Anystidae Erythraeacarus sp. n/a Pogonomyrmex rugosus in nest of Neece & Bartell, 1982 Bdella hospita Banks, 1916 Iridomyrmex sp. in nest of ants Banks, 1916 Bdellidae Bdella hospita Banks, 1916 Polyrhachis hexacantha in nest of ants Banks, 1916 Belbidae undet sp. n/a Pogonomyrmex badius in nest of Porter, 1985 Damaeidae Damaeus verticillipes n/a Pheidole flavens ants eat mites Wilson, 2005 Damaeidae gen. sp. n/a Myrmecina nipponica bodies in refuse sites Masuko, 1994 Erythraeidae Achorolophus trucidatus Hull, 1923 Crematogaster rufotestacea in jaws of Hull, 1923 Erythraeidae sp. n/a Pogonomyrmex barbatus in nest of Neece & Bartell, 1982 Erythraeidae sp. n/a Aphaenogaster cockerelli in nest of Neece & Bartell, 1982 189 Erythraeidae prominens Banks, 1916 Papyrius nitidus in nest of ants Banks, 1916

Southcott, 1989 cites Andre, 1929; Erythraeidae Forania mentonensis (André, 1929) Plagiolepis pygmaea parasitic on ants 1930 Southcott, 1989 cites Andre, 1929; Erythraeidae Forania mentonensis (André, 1929) Pheidole pallidula parasitic on ants 1931 Erythraeidae Leptus clarki Southcott, 1989 Pogonomyrmex salinus parasitic on ants Southcott, 1989 Erythraeidae Leptus debeauforti Oudemans, 1905 unknown (ants) parasitic on ants Southcott, 1989 Erythraeidae Makolia crimeaensis Saboori et al., 2009 Lasius sp. ectoparasite of ants Saboori et al., 2009 Erythraeidae nr. Abrolophus sp. n/a Cyphomyrmex wheeleri in nest of Neece & Bartell, 1982 Erythraeidae nr. Abrolophus sp. n/a Pogonomyrmex imberbiculus in nest of Neece & Bartell, 1982 Erythraeidae Rhyncholophus attolus Banks, 1916 Pachycondyla lutea in nest of ants Banks, 1916 Erythraeidae Rhyncholophus retentus Banks, 1916 Papyrius nitidus in nest of ants Banks, 1916 Continued Table 32. Non Mesostigmatid, Heterostigmatid, and Astigmatid mites and their Associated Ants from Literature.

Table 32 continued MITE HIGHER LEVEL MITE SPECIES AUTHOR HOST SPECIES ASSOCIATION REFERENCE Erythraeidae Rhyncholophus retentus Banks, 1916 Polyrhachis hexacantha in nest of ants Banks, 1916 Galumnidae acutifrons Banks, 1915 Pheidole hirtula in nest of Banks, 1915 Galumnidae Galumna sp. n/a Pheidole flavens ants eat mites Wilson, 2005 Galumnidae Pergalumna nervosa n/a Pheidole flavens ants eat mites Wilson, 2005 Haplozetidae Xylobates lophotrichus n/a Pheidole flavens ants eat mites Wilson, 2005 Nothridae Nothrus sp. n/a Pheidole flavens ants eat mites Wilson, 2005 Oribatida Caradodes sp. n/a Myrmecina nipponica bodies in refuse sites Masuko, 1994 Oribatida Ceratozetella imperatoria (Aoki) Myrmecina nipponica bodies in refuse sites Masuko, 1994 Oribatida Ceratozetella imperatoria (Aoki) Myrmecina nipponica fed on in nest Masuko, 1994

190 Oribatida Ceratozetes mediocris Berlese Myrmecina nipponica bodies in refuse sites Masuko, 1994

Oribatida Eremoeus cymba Nic. Formica rufa in nest of Wasmann, 1899 Oribatida Eremoeus cymba Nic. Formica rufa in nest of Wasmann, 1899 Oribatida Gustavia microcephala (Nicolett) Myrmecina flava fed on in lab Masuko, 1994 Oribatida Gustavia microcephala (Nicolett) Myrmecina nipponica fed on in lab Masuko, 1994 Oribatida Leiosoma longipilis Moniez, 1894 Monomorium antarcticum in nest of Moniez, 1894 Oribatida Leiosoma longipilis Moniez, 1894 Monomorium smithii in nest of Moniez, 1894 Oribatida Leiosoma longipilis Moniez, 1894 Monomorium antarcticum in nest of Moniez, 1894 Oribatida Leiosoma similis n/a Formica rufa in nest of Wasmann, 1899 Oribatida Oribata alata Herm. Formica rufa in nest of Moniez, 1892 Oribatida Oribata alata Herm. Formica rufa in nest of Moniez, 1894 Continued

Table 32 continued

HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Oribatida Oribata alata n/a Formica rufa in nest of Wasmann, 1899 Oribatida Oribata globula Nic. parvulus in nest of Moniez, 1894 Oribatida Planaphacarus kugohi (Aoki) Myrmecina nipponica bodies in refuse sites Masuko, 1994 Oribatida Protoribates myrmecophilus Aoki & Ito, 1997 Myrmecina sp. feeding on wood, groomed by ants Ito & Aoki, 2003 Oribatida Protoribates myrmecophilus Aoki & Ito, 1997 Myrmecina spp. cared for by ants Ito & Aoki, 2003 Oribatida ardua (Koch.) Myrmecina flava bodies in refuse sites Masuko, 1994 Oribatida Rhysotritia ardua (Koch.) Myrmecina nipponica bodies in refuse sites Masuko, 1994 Oribatida Rhysotritia ardua (Koch.) Myrmecina nipponica fed on in nest Masuko, 1994 Oribatida Scutovertes sculptus n/a Formica rufa in nest of Wasmann, 1899

191 Oribatida Tegeocranus labyrinthicus n/a Formica rufa in nest of Wasmann, 1899

Oribatida Xylobates magnus Aoki Myrmecina flava fed on in lab Masuko, 1994 Oribatida Xylobates magnus Aoki Myrmecina nipponica fed on in lab Masuko, 1994 Oribatida Xylobates sp. n/a Myrmecina nipponica bodies in refuse sites Masuko, 1994 Oribatida, Aribatidae Aribates javensis Aoki, et al. 1994 Myrmecina sp. obligate myrmecophile Aoki, et al. 1994 Oribatida, Cepheidae Cepheus tegeocranus Herm. Formica rufa in nest of Moniez, 1892 Oribatida, Cepheidae Eremaeus cymba Nic. Formica rufa in nest of Moniez, 1892 Oribatidae Galumna antalata Banks, 1916 Amblyopone australis in nest of ants Banks, 1916 Oribatidae Galumna antalata Banks, 1916 Rhytidoponera metallica in nest of ants Banks, 1916 Continued

Table 32 continued HIGHER LEVEL MITE SPECIES MITE AUTHOR HOST SPECIES ASSOCIATION REFERENCE Oribatidae Notaspis spinulosa Michael Iridomyrmex sp. in nest of ants Banks, 1916 pacificum n/a Pheidole flavens soil in nest Wilson, 2005 Scheloribates sp. n/a Pheidole flavens ants eat mites Wilson, 2005 undet sp. n/a Pheidole flavens soil in nest Wilson, 2005 Tectocepheidae Tectocepheus sp. n/a Pheidole flavens ants eat mites Wilson, 2005 Neece & Bartell, Eutrombidium sp. n/a Aphaenogaster cockerelli in nest of 1982 Trombidiidae Rhyncholophus phalangloides n/a Formica exsecta in nest of Moniez, 1894 Trombidiidae Rhyncholophus regalis Koch. Formica fusca in nest of Moniez, 1892 Trombidiidae Rhyncholophus regalis Koch. Formica rufa in nest of Moniez, 1892

192 Trombidiidae Rhyncolophus regalis Koch. Formica rufa in nest of Wasmann, 1899 Trombidiidae aequalis Banks, 1916 Pachycondyla lutea in nest of ants Banks, 1916

Trombidiidae Trombidium erythrellum Koch. Tetramorium caespitum in nest of Moniez, 1892

Appendix B: Chapter 2 Supplementary Tables

193

Appendix B: Chapter 2 Supplementary Tables

LOCALITY FULL LOCALITY NAME LAT/LONG DATE OF FIELD NUMBERS # OF TOTAL CODE COLLECTIONS COLL. ANTS KLOC0029 nr. North Adams Public (38.963 N, 83.569 W) 6/11/2008 KAU-08-0611-1 1 118 Library, 17806 SR 247, Seaman, Adams Co. OH, U.S.A. LOC006720 Primitive Family (39.4354 N, 82.5277 5/31/2008 KAU-08-0531-3; KAU-08-0531-4 2 70 Campground, Hocking W) Hills State Forest, Logan, Hocking Co., OH, U.S.A. KLOC0031 Wooster, Wayne Co., OH, (40.785 N, 81.931 W) 5/27/2008 KAU-08-0527-1 1 1 U.S.A. L01191 1315 Kinnear Rd. (39.9976 N, 83.0441 6/20/2008; KAU-08-0620-1; KAU-08-0625-1; KAU-08-0707- 10 362 Columbus, Franklin Co., W) 6/25/2008; 2; KAU-09-0824-10; KAU-09-0824-11; KAU-09- OH, U.S.A. 7/7/2008; 0824-12; KAU-09-0824-13; KAU-09-0824-14a; 194 8/24/2009 KAU-09-0824-14b; KAU-09-0824-14c

LOC006696 Jackson Lake, Hunting and (39.1022 N, 82.7923 8/29/2008; KAU-08-0829-3; KAU-08-0829-4; KAU-08-0829- 38 1185 Fishing Area, nr. Big Rock, W) 6/13/2009; 5; KAU-08-0829-6; KAU-08-0829-7; KAU-08- Jackson Co., OH, U.S.A. 9/9/2009 0829-8; KAU-08-0829-9; KAU-08-0829-10; KAU-08-0829-11; KAU-08-0829-13; KAU-09- 0613-27; KAU-09-0613-28; KAU-09-0613-29; KAU-09-0613-30; KAU-09-0613-31; KAU-09- 0613-32; KAU-09-0613-33; KAU-09-0909-1; KAU-09-0909-2; KAU-09-0909-5; KAU-09-0909- 6; KAU-09-0909-7; KAU-09-0909-9a; KAU-09- 0909-9b; KAU-09-0909-10a; KAU-09-0909-10b; KAU-09-0909-11; KAU-09-0909-12; KAU-09- 0909-13; KAU-09-0909-15; KAU-09-0909-16; KAU-09-0909-17; KAU-09-0909-18;KAU-09- 0909-19; KAU-09-0909-20; KAU-09-0909-21a; KAU-09-0909-21b; KAU-09-0909-22 Continued Table 33. Localities and Colony Collections. Locality codes, full locality names, coordinates, collection dates, colonies collected at each locality (field numbers), and number of colonies and ants are shown.

Table 33 continued LOCALITY FULL LOCALITY NAME LAT/LONG DATE OF FIELD NUMBERS # OF TOTAL CODE COLLECTIONS COLL. ANTS LOC006697 West campus Woods nr (40.0161 N, 83.0452 W) 4/24/2008; KAU-08-0424-3; KAU-08-0529-2; KAU-08- 35 829 Turf Research Building, 5/29/2008; 0529-3; KAU-08-0529-4; KAU-08-0529-5; Ohio State University, 5/30/2008; KAU-08-0706-1; KAU-08-0706-7; KAU-08- Columbus, Franklin Co., 7/6/2008; 0706-8; KAU-08-0706-10; KAU-08-0706-11; OH, U.S.A. 9/3/2008; KAU-08-0706-12; KAU-08-0706-13; KAU-08- 7/7/2009 0706-15; KAU-08-0706-16; KAU-08-0706-20; KAU-08-0706-21; KAU-08-0706-30; KAU-08- 0706-31; KAU-08-0903-3; KAU-08-0903-4; KAU-08-0903-5; KAU-08-0903-6; KAU-08- 0903-7; KAU-08-0903-8; KAU-08-0903-9; KAU-08-0903-10; KAU-08-0903-11; KAU-08- 0903-12; KAU-09-0707-1; KAU-09-0707-2; KAU-09-0707-5; KAU-09-0707-6; KAU-09- 195 0707-8; KAU-09-0707-10; KAU-09-0707-11

LOC006698 Public Hunting and Fishing (39.61 N, 82.00 W) 6/10/2008 KAU-08-0610-1; KAU-08-0610-10; KAU-08- 7 215 Area, OH-555, btwn 0610-4; KAU-08-0610-5; KAU-08-0610-6; Chesterhill and Alexandria, KAU-08-0610-7; KAU-08-0610-8 Morgan Co., OH, U.S.A. LOC006701 Western Campus, Miami (39.5044 N, 84.7241 W) 8/15/2008; KAU-08-0815-1; KAU-09-0424-1a; KAU-09- 30 916 University, Oxford, Butler 4/24/2009; 0424-1b; KAU-09-0424-2; KAU-09-0424-3; Co., OH, U.S.A. 7/21/2009 KAU-09-0424-4; KAU-09-0424-5; KAU-09- 0424-6; KAU-09-0424-7; KAU-09-0424-8; KAU-09-0424-10; KAU-09-0424-12a; KAU- 09-0424-12b; KAU-09-0424-13; KAU-09- 0424-14; KAU-09-0424-15; KAU-09-0424-17; KAU-09-0424-18; KAU-09-0424-19; KAU-09- 0721-2; KAU-09-0721-3; KAU-09-0721-5; KAU-09-0721-7; KAU-09-0721-8; KAU-09- 0721-9; KAU-09-0721-10; KAU-09-0721-11; KAU-09-0721-12; KAU-09-0721-14; KAU-09- 0721-15 LOC006702 55 Kelly Dr., Oxford, (39.5199 N, 84.7423 W) 5/25/2009 KAU-09-0525-1; KAU-09-0525-2 2 6 Butler Co., OH, U.S.A. Continued

Table 33 continued LOCALITY FULL LOCALITY NAME LAT/LONG DATE OF FIELD NUMBERS # OF TOTAL CODE COLLECTIONS COLL. ANTS LOC006703 West campus Woods nr (40.0013 N, 83.0427 9/25/2008; KAU-08-0925-3; KAU-09-0824-15a; KAU-09- 12 483 parking lots, Ohio State W) 8/24/2009 0824-15b; KAU-09-0824-16; KAU-09-0824- University, Columbus, 17a; KAU-09-0824-17b; KAU-09-0824-18; Franklin Co., OH, U.S.A. KAU-09-0824-19; KAU-09-0824-20; KAU-09- 0824-21; KAU-09-0824-23a; KAU-09-0824-23b LOC006704 Kenney Park, 5201 (40.0661 N, 83.0307 8/12/2008; KAU-08-0812-1; KAU-08-0812-2; KAU-10- 4 120 Delawanda Ave, Columbus, W) 3/10/2010 0310-1 Franklin Co., OH, U.S.A. LOC006705 613 Smokey Hollow Rd., (39.0540 N, 83.0999 6/11/2008; KAU-08-0611-2; KAU-08-0611-3; KAU-09- 42 1261 Piketon, Pike Co., OH, U.S.A. W) 6/13/2009; 0613-1; KAU-09-0613-2; KAU-09-0613-3; 9/10/2009 KAU-09-0613-4; KAU-09-0613-5; KAU-09- 0613-6; KAU-09-0613-7; KAU-09-0613-8; KAU-09-0613-9; KAU-09-0613-10; KAU-09- 0613-11; KAU-09-0613-12; KAU-09-0613-13;

196 KAU-09-0613-14; KAU-09-0613-15; KAU-09- 0613-16; KAU-09-0613-17; KAU-09-0613-18; KAU-09-0613-19; KAU-09-0613-20; KAU-09- 0613-21a; KAU-09-0613-21b; KAU-09-0613- 22; KAU-09-0613-24; KAU-09-0613-25; KAU- 09-0613-26; KAU-09-0613-45; KAU-09-0910-2; KAU-09-0910-3; KAU-09-0910-4; KAU-09- 0910-5; KAU-09-0910-7; KAU-09-0910-8; KAU-09-0910-9; KAU-09-0910-10a; KAU-09- 0910-10b; KAU-09-0910-11; KAU-09-0910-12; KAU-09-0910-13; KAU-09-0910-14 LOC006706 Wickline Rd., Beaver, Pike (39.1164 N, 82.8094 7/18/2008; KAU-08-0718-1; KAU-08-0718-2; KAU-09- 11 352 Co., OH, U.S.A. W) 6/13/2009; 0613-34; KAU-09-0613-35; KAU-09-0613-36a; 9/9/2009 KAU-09-0613-36b; KAU-09-0613-37; KAU-09- 0613-38; KAU-09-0909-27; KAU-09-0909-28; KAU-09-0909-29 Continued

Table 33 continued LOCALITY FULL LOCALITY NAME LAT/LONG DATE OF FIELD NUMBERS # OF TOTAL CODE COLLECTIONS COLL. ANTS LOC006707 4307 Longfork Rd., Piketon, (39.0472 N, 83.0764 W) 6/12/2009; KAU-09-0612-1; KAU-09-0612-2; KAU-09- 6 139 Pike Co., OH, U.S.A. 6/14/2009 0612-3; KAU-09-0612-4; KAU-09-0612-5; KAU-09-0614-2 LOC006708 268 Olentangy St. Columbus, (40.0206 N, 83.0054 W) 5/11/2009; KAU-09-0511-1; KAU-09-0518-1 2 127 Franklin Co., OH, U.S.A. 5/18/2009 LOC006709 1098 Afton Rd, Columbus, (40.0284 N, 83.0439 W) 6/2/2008; KAU-08-0602-1; KAU-08-0615-1; KAU-08- 6 593 Franklin Co., OH, U.S.A. 6/15/2008; 0625-2; KAU-08-0707-1a; KAU-08-0707-1b; 6/25/2008; KAU-08-0707-3a 7/7/2008 LOC006710 Glen Echo Ravine, Indianola (40.0189 N, 83.0029 W ) 8/28/2008; KAU-08-0828-1a; KAU-08-0828-1b; KAU-08- 17 442 Ave at Olentangy St., 8/24/2009 0828-2; KAU-08-0828-3; KAU-08-0828-4; Columbus, Franklin Co., OH KAU-09-0824-1; KAU-09-0824-2; KAU-09- U.S.A. 0824-3a; KAU-09-0824-3b; KAU-09-0824-3c; KAU-09-0824-3d; KAU-09-0824-4; KAU-09- 197 0824-5; KAU-09-0824-6; KAU-09-0824-7; KAU-09-0824-8; KAU-09-0824-9 LOC006711 Rocky Fork Rd, Otway, (38.7807 N, 83.2591 W) 6/11/2008; KAU-08-0611-4; KAU-08-0611-5 2 57 Jefferson Twp, Adams Co., 6/12/2008 OH, U.S.A. LOC006712 138 Kensington St, (39.5078 N, 84.3709 W) 7/22/2008; KAU-08-0722-1; KAU-09-0522-1; KAU-09- 14 483 Middletown, Butler Co., OH, 5/22/2009; 0522-2; KAU-09-0522-3a; KAU-09-0522-3b; U.S.A. 7/20/2009; KAU-09-0522-4; KAU-09-0522-6; KAU-09- 3/27/2010 0522-8; KAU-09-0522-9; KAU-09-0522-10; KAU-09-0522-11; KAU-09-0720-1; KAU-10- 0316-1; KAU-10-0327-1 LOC006713 Main Campus near Aronoff (39.9959 N, 83.0157 W 6/8/2009 KAU-09-0608-1 1 124 Lab, Ohio State University, Columbus, Franklin Co., OH, U.S.A. LOC006714 348 E 15th Ave, Columbus, (39.9998 N, 82.9985 W) 6/7/2008; KAU-08-0607-1; KAU-08-0609-1 2 25 Franklin Co., OH, U.S.A. 6/9/2008 LOC006715 Grant park, Indiana Ave and (41.8676 N, 87.6226 W) 9/16/2008 KAU-08-0916-1 1 9 Roosevelt Rd, Chicago, Cook Co., IL, U.S.A. Continued

Table 33 continued LOCALITY FULL LOCALITY NAME LAT/LONG DATE OF FIELD NUMBERS # OF TOTAL CODE COLLECTIONS COLL. ANTS LOC006716 Crane Hollow Nature (39.4885 N, 82.5585 W) 7/25/2009 KAU-09-0725-1; KAU-09-0725-2; KAU-09- 15 744 Preserve, Cream Ridge Rd. at 0725-3; KAU-09-0725-4; KAU-09-0725-5; Extine Rd., Laurel Township, KAU-09-0725-6; KAU-09-0725-7; KAU-09- Hocking Co., OH 0725-8; KAU-09-0725-9; KAU-09-0725-10; KAU-09-0725-11; KAU-09-0725-12; KAU-09- 0725-13; KAU-09-0725-14; KAU-09-0725-15 LOC006717 Research Foundation (37.1815 N, 86.0421 W) 5/16/2009 KAU-09-0516-1; KAU-09-0516-2; KAU-09- 12 432 Research Station, Hamilton 0516-3; KAU-09-0516-4; KAU-09-0516-5; Valley Rd. nr Cave City, KAU-09-0516-6a; KAU-09-0516-6b; KAU-09- Hart Co., KY, U.S.A. 0516-7; KAU-09-0516-8b; KAU-09-0516-9; KAU-09-0516-11; KAU-09-0516-12 TOTAL 273 9093

198

MITE FAMILY MITE SPECIES SPECIMEN NUMBER (OSAL) Acaridae Acarus sp1 OSAL0092833 Acaridae Acotyledon sp1 OSAL0092683 Acaridae Armacarus sp1 OSAL0092871 Acaridae Cosmoglyphus sp1 OSAL0101145 Acaridae Cosmoglyphus sp2 OSAL0101046 Acaridae Forcellinia sp1 OSAL0101152 Acaridae Forcellinia sp2 OSAL0101049 Acaridae Forcellinia sp3 OSAL0102850 Acaridae Forcellinia sp4 OSAL0101554 Acaridae Forcellinia sp5 OSAL0101740 Acaridae Forcellinia sp6 OSAL0092865 Acaridae Forcellinia sp7 OSAL0101203 Acaridae Forcellinia sp8 OSAL0101038 Acaridae Forcellinia sp9 OSAL0101026 Acaridae Forcellinia sp10 OSAL0101298 Acaridae Forcellinia sp11 OSAL0092857 Acaridae Froriepia sp1 OSAL0101021 Acaridae Froriepia sp2 OSAL0102858 Acaridae Froriepia sp3 OSAL0101141 Acaridae Sancassania sp1 OSAL0101556 Acaridae Sancassania sp2 OSAL0092207 Acaridae Sancassania sp3 OSAL0102739 Acaridae Schwiebea sp1 OSAL0101106 Acaridae Schwiebea sp2 OSAL0092817 Acaridae Schwiebea sp3 OSAL0101292 Acaridae Schwiebea sp4 OSAL0101451 Acaridae Schwiebea sp5 OSAL0101246 Acaridae Schwiebea sp6 OSAL0102962 Acaridae Schwiebea sp7 OSAL0101433 Acaridae Schwiebea sp8 OSAL0101731 Acaridae Schwiebea sp9 OSAL0101207 Acaridae Schwiebea sp10 OSAL0101561 Acaridae Schwiebea sp11 OSAL0102825 Acaridae Schwiebea sp12 OSAL0102738 Astigmata undet undet gen sp3 OSAL0101128 Astigmata undet undet gen sp4 OSAL0102885 Histiostomatidae Histiostoma sp1 OSAL0101753 Histiostomatidae Histiostoma sp2 OSAL0092601 Histiostomatidae Histiostoma sp3 OSAL0101349 Histiostomatidae Histiostoma sp4 OSAL0101551 Histiostomatidae Histiostoma sp5 OSAL0101751 Histiostomatidae Histiostoma sp6 OSAL0101579 Histiostomatidae Histiostoma sp7 OSAL0101553 Histiostomatidae Histiostoma sp8 OSAL0101378 Histiostomatidae Histiostoma sp9 OSAL0101656 Histiostomatidae Histiostoma sp10 OSAL0092998 Histiostomatidae Histiostoma sp11 OSAL0092984 Continued Table 34. Mite voucher specimens deposited in the Ohio State University Acarology Collection as slides and their corresponding OSAL barcode numbers.

199

Table 34 continued MITE FAMILY MITE SPECIES SPECIMEN NUMBER (OSAL) Histiostomatidae Histiostoma sp12 OSAL0101237 Histiostomatidae Histiostoma sp13 OSAL0102830 Histiostomatidae Histiostoma sp14 OSAL0101617 Histiostomatidae Histiostoma sp15 OSAL0101255 Histiostomatidae Histiostoma sp16 OSAL0101729 Histiostomatidae Histiostoma sp17 OSAL0101797 Histiostomatidae Histiostoma sp18 OSAL0101693 Histiostomatidae Histiostoma sp19 OSAL0101468 Histiostomatidae Histiostoma sp20 OSAL0101154 Histiostomatidae Histiostoma sp21 OSAL0092230 Histiostomatidae Histiostoma sp22 OSAL0092644 Histiostomatidae Histiostoma sp23 OSAL0101626 Histiostomatidae Histiostoma sp24 OSAL0102839 Histiostomatidae Histiostoma sp25 OSAL0102849 Histiostomatidae Histiostoma sp26 OSAL0102844 Histiostomatidae Histiostoma sp27 OSAL0102735 Antennophoridae Antennophorus sp1 OSAL0101785 Antennophoridae Antennophorus sp2 OSAL0101440 Blattisocidae Lasioseius sp1 OSAL0101326 Circocyllibanidae Trichocylliba sp1 OSAL0101848 Digamasellidae undet gen sp1 OSAL0092717 Digamasellidae undet gen sp2 OSAL0092733 Digamasellidae undet gen sp3 OSAL0092730 Digamasellidae undet gen sp4 OSAL0092734 Digamasellidae undet gen sp5 OSAL0092729 Dinychidae Dinychus sp1 OSAL0092739 Halolaelapidae undet gen sp1 OSAL0092404 Laelapidae Cosmolaelaps sp1 OSAL0092710 Laelapidae Cosmolaelaps sp2 OSAL0102889 Laelapidae Cosmolaelaps sp3 OSAL0101768 Laelapidae Cosmolaelaps sp4 OSAL0101741 Laelapidae Cosmolaelaps sp7 OSAL0101411 Laelapidae Cosmolaelaps sp8 OSAL0101222 Laelapidae Cosmolaelaps sp9 OSAL0101690 Laelapidae Cosmolaelaps sp10 OSAL0102831 Laelapidae Cosmolaelaps sp11 OSAL0101915 Laelapidae Cosmolaelaps sp13 OSAL0101585 Laelapidae Cosmolaelaps sp14 OSAL0101725 Laelapidae Cosmolaelaps sp15 OSAL0092709 Laelapidae Cosmolaelaps sp16 OSAL0101709 Laelapidae Cosmolaelaps sp18 OSAL0092698 Laelapidae Cosmolaelaps sp19 OSAL0101544 Laelapidae Cosmolaelaps sp20 OSAL0092696 Laelapidae Cosmolaelaps sp21 OSAL0092689 Laelapidae Gaeolaelaps sp1 OSAL0101158 Laelapidae Gaeolaelaps sp2 OSAL0101421 Laelapidae Gaeolaelaps sp3 OSAL0101376 Laelapidae Gaeolaelaps sp4 OSAL0101273 Laelapidae Gaeolaelaps sp5 OSAL0092407 Laelapidae Gaeolaelaps sp6 OSAL0101883 Continued 200

Table 34 continued MITE FAMILY MITE SPECIES SPECIMEN NUMBER (OSAL) Laelapidae Gaeolaelaps sp7 OSAL0102944 Laelapidae Gaeolaelaps sp8 OSAL0101164 Laelapidae Laelaspis cf. dubitatus OSAL0101408 Laelapidae Laelaspis cf. vitzthumi OSAL0101697 Laelapidae Laelaspis n.sp. nr. vitzthumi OSAL0092700 Laelapidae Laelaspis nr. picketti OSAL0092761 Laelapidae Stratiolaelaps sp1 OSAL0092692 Laelapidae undet gen sp1 OSAL0101383 Macrochelidae Macrocheles sp1 OSAL0092403 Macrochelidae Macrocheles sp2 OSAL0092417 Ologamasidae Gamasiphis sp1 OSAL0092411 Oplitidae Oplitis (Uroplitana) sp11 OSAL0101442 Oplitidae Oplitis aktius OSAL0101852 Oplitidae Oplitis alienorum OSAL0092569 Oplitidae Oplitis exsectoidesorum OSAL0101120 Oplitidae Oplitis sarcinulus OSAL0101115 Oplitidae Oplitis sp1 OSAL0101118 Oplitidae Oplitis sp2 OSAL0101213 Oplitidae Oplitis sp3 OSAL0101212 Oplitidae Oplitis sp8 OSAL0102942 Oplitidae Oplitis sp9 OSAL0101836 Oplitidae Oplitis sp10 OSAL0101111 Parholaspididae Holaspina sp1 OSAL0092767 Parholaspididae Holaspina sp2 OSAL0092758 Parholaspididae Holaspina sp3 OSAL0092765 Rhodacaridae undet gen sp1 OSAL0101672 Trematuridae Trichouropoda sp1 OSAL0092753 Uropodellidae Uropodella laciniata OSAL0092419 Uropodidae undet gen sp1 OSAL0092567 Uropodidae undet gen sp3 OSAL0092742 Uropodidae Uropolyaspis sp1 OSAL0101258 Uropodidae Uropolyaspis sp2 OSAL0101110 Cunaxidae undet gen sp1 OSAL0102887 Ereynetidae undet gen sp1 OSAL0092401 Eupodidae undet gen sp1 OSAL0092414 Microdispidae Caesarodispus sp2 OSAL0101122 Microdispidae Caesarodispus sp11 OSAL0092887 Microdispidae Caesarodispus sp15 OSAL0102855 Microdispidae Caesarodispus sp16 OSAL0102723 Microdispidae Unguidispus sp10 OSAL0101260 Pygmephoridae Bakerdania sp3 OSAL0101123 Pygmephoridae Bakerdania sp5 OSAL0101200 Pygmephoridae Bakerdania sp7 OSAL0101761 Pygmephoridae Bakerdania sp9 OSAL0102868 Pygmephoridae Bakerdania sp12 OSAL0101234 Pygmephoridae Bakerdania sp13 OSAL0101630 Pygmephoridae Bakerdania sp14 OSAL0101168 Pygmephoridae Petalomium sp6 OSAL0092892 Pygmephoridae Petalomium sp8 OSAL0092896 Pygmephoridae Pygmephorellus sp1 OSAL0092915 Continued

201

Table 34 continued MITE FAMILY MITE SPECIES SPECIMEN NUMBER (OSAL) Rhagidiidae undet gen sp1 OSAL0092400 Scutacaridae Diversipes sp1 OSAL0092617 Scutacaridae Imparipes sp1 OSAL0101560 Scutacaridae Imparipes sp2 OSAL0101907 Scutacaridae Imparipes sp4 OSAL0101985 Scutacaridae Imparipes sp5 OSAL0101308 Scutacaridae Imparipes sp6 OSAL0101217 Scutacaridae Imparipes sp7 OSAL0101172 Scutacaridae Imparipes sp8 OSAL0101171 Scutacaridae Imparipes sp9 OSAL0101756 Scutacaridae Imparipes sp10 OSAL0101755 Scutacaridae Imparipes sp11 OSAL0101623 Scutacaridae Imparipes sp12 OSAL0102938 Scutacaridae Imparipes sp13 OSAL0101318 Scutacaridae Imparipes sp14 OSAL0102853 Scutacaridae Pygmodispus sp1 OSAL0092275 Scutacaridae Scutacarus sp1 OSAL0101583 Scutacaridae Scutacarus sp2 OSAL0101466 Scutacaridae Scutacarus sp3 OSAL0092616 Scutacaridae Scutacarus sp4 OSAL0101227 Scutacaridae Scutacarus sp5 OSAL0101133 Scutacaridae Scutacarus sp6 OSAL0101235 Scutacaridae Scutacarus sp7 OSAL0101342 Scutacaridae Scutacarus sp8 OSAL0101517 Scutacaridae Scutacarus sp9 OSAL0101513 Scutacaridae Scutacarus sp10 OSAL0101902 Scutacaridae Scutacarus sp11 OSAL0101967 Scutacaridae Scutacarus sp12 OSAL0101663 Scutacaridae Scutacarus sp13 OSAL0092281 Scutacaridae Scutacarus sp14 OSAL0101199 Scutacaridae Scutacarus sp15 OSAL0102931 Scutacaridae Scutacarus sp16 OSAL0092272 Scutacaridae Scutacarus sp17 OSAL0101429 Scutacaridae Scutacarus sp18 OSAL0102951 Scutacaridae Scutacarus sp19 OSAL0101555 Scutacaridae Scutacarus sp20 OSAL0101285 Scutacaridae Scutacarus sp21 OSAL0101132 Scutacaridae Scutacarus sp22 OSAL0102937 Scutacaridae Scutacarus sp23 OSAL0101480 Scutacaridae Scutacarus sp24 OSAL0102904 Scutacaridae Scutacarus sp25 OSAL0102909 Scutacaridae Scutacarus sp26 OSAL0102901 Scutacaridae Scutacarus sp27 OSAL0102934 Scutacaridae Scutacarus sp28 OSAL0102856 Tarsonemidae undet gen sp1 OSAL0101060 Tarsonemidae undet gen sp2 OSAL0092380 Tarsonemidae undet gen sp3 OSAL0092907 Tarsonemidae undet gen sp4 OSAL0101065 Tarsonemidae undet gen sp5 OSAL0092908 Tarsonemidae undet gen sp6 OSAL0101058

202

ANT SPECIES SPECIMEN NUMBER (OSUC) Amblyopone pallipes OSUC0359992 Aphaenogaster fulva OSUC0359991 Aphaenogaster picea OSUC0359990 Aphaenogaster rudis OSUC0359989 Aphaenogaster tennesseensis OSUC0359988 Brachymyrmex depilis OSUC0359987 Camponotus chromaiodes OSUC0359986 Camponotus nearcticus OSUC0359985 Camponotus pennsylvanicus OSUC0359984 Camponotus subbarbatus OSUC0359957 Crematogaster cerasi OSUC0359983 Crematogaster lineolata OSUC0359982 Crematogaster pilosa OSUC0359981 Formica exectoides OSUC0359980 Formica integra OSUC0359979 Formica nitidiventris OSUC0359978 Formica rubicunda OSUC0359956 Formica subsericea OSUC0359977 Lasius alienus OSUC0359976 Lasius claviger OSUC0359971 Lasius interjectus OSUC0359970 Lasius nearcticus OSUC0359975 Lasius neoniger OSUC0359974 Lasius speculiventris OSUC0359973 Lasius umbratus OSUC0359972 Messor pergandei OSUC0359951 Monomorium minimum OSUC0359969 Myrmecina americana OSUC0359955 Myrmica americana OSUC0359968 Myrmica latifrons OSUC0359967 Myrmica punctiventris OSUC0359954 Paratrechina faisonensis OSUC0359966 Paratrechina parvula OSUC0359952 Pheidole bicarinata OSUC0359965 Ponera pennsylvanica OSUC0359964 Prenolepis imparis OSUC0359953 Proceratium pergandei OSUC0359963 Pyramica ohioensis OSUC0359962 Pyramica reflexa OSUC0359961 Solenopsis molesta OSUC0359960 Tapinoma sessile OSUC0359959 Tetramorium caespitum OSUC0359958 Table 35. Ant voucher specimens deposited in the Ohio State University Insect Collection as pinned specimens and their corresponding OSUC barcode numbers.

203

# COL. # COL. # COL. TOTAL TOTAL TOTAL F TOTAL M TOTAL TOTAL GENUS SPECIES 2008 2009 2010 COL. WORKERS ALATES ALATES DEALATES ANTS Amblyopone pallipes 1 2 0 3 13 2 3 1 19 Aphaenogaster fulva 4 11 0 15 499 43 62 0 604 Aphaenogaster picea complex 5 17 0 22 846 32 66 0 944 Aphaenogaster rudis complex 3 12 0 15 462 10 24 1 497 Aphaenogaster tennessensis 4 2 0 6 164 33 34 1 232 Brachymyrmex depilis 0 3 0 3 67 0 1 1 69 Camponotus chromaiodes 11 3 0 14 102 73 16 5 196 Camponotus nearcticus 1 2 0 3 12 18 20 0 50 Camponotus pennsylvanicus 4 0 0 4 4 1 1 2 8 Camponotus subbarbatus 0 3 0 3 73 0 0 1 74

204 204 Crematogaster cerasi 0 4 0 4 225 0 0 0 225 Crematogaster lineolata 2 0 0 2 61 0 0 0 61

Crematogaster pilosa 0 2 0 2 20 0 0 0 20 Formica exectoides 2 9 0 11 360 0 0 0 360 Formica integra *1 1 0 1 89 0 0 0 89 Formica nitidiventris 0 3 0 3 60 0 0 0 60 Formica rubicunda 0 1 0 1 11 0 0 0 11 Formica subsericea 0 8 0 8 192 0 0 0 192 Lasius alienus 9 10 0 19 698 0 10 0 708 Lasius claviger 1 3 0 4 155 0 27 0 182 Lasius interjectus 0 1 0 1 17 0 0 0 17 Continued Table 36. Ohio Ant Collection Summary: Ant species, number of colonies collected (# COL.), and number of each caste (F=female, M=male) collected are shown in the table. *Designates a collection made by putting the entire collection on the funnel rather than by the usual method.

Table 36 continued # COL. # COL. # COL. TOTAL TOTAL TOTAL F TOTAL M TOTAL TOTAL GENUS SPECIES 2008 2009 2010 COL. WORKERS ALATES ALATES DEALATES ANTS Lasius nearcticus 1 2 1 4 66 45 49 0 160 Lasius neoniger 4 11 0 15 368 54 44 2 468 Lasius speculiventris 0 1 0 1 55 0 1 0 56 Lasius umbratus 5 12 0 17 435 94 79 3 611 Monomorium minimum 0 3 0 3 107 0 0 0 107 Myrmecina americana 0 3 0 3 15 1 1 0 17 Myrmica americana 0 2 0 2 22 0 0 1 23 Myrmica latifrons 1 0 0 1 2 0 0 0 2 Myrmica punctiventris 0 2 0 2 24 0 0 0 24 Paratrechina faisonensis 1 2 0 3 94 0 0 0 94 Paratrechina parvula 0 1 0 1 67 3 0 0 70

205

205 Pheidole bicarinata 0 2 0 2 32 0 0 0 32

Ponera pennsylvanica 4 16 0 20 121 2 0 3 126 Prenolepis imparis 0 1 2 3 109 4 54 0 167 Proceratium pergandei 1 0 0 1 1 0 0 0 1 Pyramica laevinasis 0 1 0 1 1 0 0 0 1 Pyramica ohioensis 2 0 0 2 40 11 0 2 53 Pyramica reflexa 0 1 0 1 13 0 0 0 13 Pyramica sp. 0 1 0 1 1 0 0 0 1 Solenopsis molesta 1 16 0 17 813 25 0 0 838 Tapinoma sessile 0 10 0 10 387 0 9 13 409 Tetramorium caespitum 11 8 0 19 776 127 192 107 1202 TOTALS: 78 192 3 273 7679 578 693 143 9093

Appendix C: Chapter 2 Raw Data

206

SUBFAMILY GENUS SPECIES FIELD # # PHORETIC # MESO # HETERO # ASTIG PREV WITH Amblyopone pallipes KAU-08-0828-2 0 0 0 0 0.00 Myrmecinae Aphaenogaster fulva KAU-08-0610-10 0 0 0 0 0.61 Myrmecinae Aphaenogaster fulva KAU-08-0706-16 1 0 0 1 0.35 Myrmecinae Aphaenogaster fulva KAU-08-0815-1 3 0 3 0 0.27 Myrmecinae Aphaenogaster fulva KAU-09-0613-17 3 0 2 1 0.15 Myrmecinae Aphaenogaster fulva KAU-09-0613-21a 1 0 0 1 0.46 Myrmecinae Aphaenogaster fulva KAU-09-0707-5 2 0 1 1 0.09 Myrmecinae Aphaenogaster fulva KAU-09-0707-6 1 0 0 1 0.14 Myrmecinae Aphaenogaster fulva KAU-09-0721-11 4 0 3 1 0.48 Myrmecinae Aphaenogaster fulva KAU-09-0721-12 2 0 2 0 0.07 Myrmecinae Aphaenogaster fulva KAU-09-0721-14 5 1 3 1 0.35 Myrmecinae Aphaenogaster fulva KAU-09-0721-15 3 1 2 0 0.43 Myrmecinae Aphaenogaster fulva KAU-09-0721-9 2 0 2 0 0.20

207 Myrmecinae Aphaenogaster picea complex KAU-08-0531-4 0 0 0 0 0.09 Myrmecinae Aphaenogaster picea complex KAU-08-0610-7 0 0 0 0 0.10 Myrmecinae Aphaenogaster picea complex KAU-08-0829-5 0 0 0 0 0.00 Myrmecinae Aphaenogaster picea complex KAU-08-0829-9 2 0 1 1 0.33 Myrmecinae Aphaenogaster picea complex KAU-09-0424-17 1 0 1 0 0.29 Myrmecinae Aphaenogaster picea complex KAU-09-0613-18 2 0 2 0 0.32 Myrmecinae Aphaenogaster picea complex KAU-09-0613-20 0 0 0 0 0.36 Myrmecinae Aphaenogaster picea complex KAU-09-0613-32 2 0 2 0 0.30 Myrmecinae Aphaenogaster picea complex KAU-09-0725-10 2 0 2 0 0.37 Myrmecinae Aphaenogaster picea complex KAU-09-0725-11 3 1 1 1 0.19 Myrmecinae Aphaenogaster picea complex KAU-09-0725-13 1 0 1 0 0.15 Continued Table 37. Raw data for Response variables used in Statisical analyses. # Phoretic indicates total number of phoretic mites found in the colony. “# Meso”, “# Hetero”, and “# Astig” designate number of Mesostigmata, number of Heterostigmatina, and number of Astigmata respectively associated with the colony. “Prev with” is the prevalence of mite infestations in the colony.

Table 37 continued SUBFAMILY GENUS SPECIES FIELD # # PHORETIC # MESO # HETERO # ASTIG PREV WITH Myrmecinae Aphaenogaster picea complex KAU-09-0725-2 1 0 1 0 0.09 Myrmecinae Aphaenogaster picea complex KAU-09-0725-3 1 0 1 0 0.05 Myrmecinae Aphaenogaster picea complex KAU-09-0725-4 1 0 1 0 0.03 Myrmecinae Aphaenogaster picea complex KAU-09-0725-5 2 0 2 0 0.13 Myrmecinae Aphaenogaster picea complex KAU-09-0725-7 1 0 1 0 0.02 Myrmecinae Aphaenogaster picea complex KAU-09-0725-8 3 1 2 0 0.18 Myrmecinae Aphaenogaster picea complex KAU-09-0725-9 2 1 1 0 0.11 Myrmecinae Aphaenogaster picea complex KAU-09-0824-6 3 0 2 1 0.16 Myrmecinae Aphaenogaster picea complex KAU-09-0910-11 3 0 3 0 0.11 Myrmecinae Aphaenogaster picea complex KAU-09-0910-12 1 0 0 1 0.10 Myrmecinae Aphaenogaster rudis complex KAU-08-0706-20 0 0 0 0 0.44 Myrmecinae Aphaenogaster rudis complex KAU-08-0706-31 1 0 1 0 0.11

208 Myrmecinae Aphaenogaster rudis complex KAU-08-0706-8 0 0 0 0 0.13 Myrmecinae Aphaenogaster rudis complex KAU-09-0424-19 2 0 2 0 0.81 Myrmecinae Aphaenogaster rudis complex KAU-09-0424-6 0 0 0 0 0.00 Myrmecinae Aphaenogaster rudis complex KAU-09-0613-11 0 0 0 0 0.05 Myrmecinae Aphaenogaster rudis complex KAU-09-0613-15 2 0 1 1 0.30 Myrmecinae Aphaenogaster rudis complex KAU-09-0613-24 1 0 1 0 0.44 Myrmecinae Aphaenogaster rudis complex KAU-09-0613-25 2 0 2 0 0.45 Myrmecinae Aphaenogaster rudis complex KAU-09-0613-31 3 1 2 0 0.76 Myrmecinae Aphaenogaster rudis complex KAU-09-0613-4 3 1 2 0 0.39 Myrmecinae Aphaenogaster rudis complex KAU-09-0613-45 3 1 2 0 0.32 Myrmecinae Aphaenogaster rudis complex KAU-09-0721-8 3 1 2 0 0.20 Myrmecinae Aphaenogaster tennessensis KAU-08-0610-1 0 0 0 0 0.63 Myrmecinae Aphaenogaster tennessensis KAU-08-0706-11 2 0 2 0 0.49 Continued

Table 37 continued SUBFAMILY GENUS SPECIES FIELD # # PHORETIC # MESO # HETERO # ASTIG PREV WITH Myrmecinae Aphaenogaster tennessensis KAU-08-0812-3 0 0 0 0 0.70 Myrmecinae Aphaenogaster tennessensis KAU-09-0707-8 4 1 3 0 0.10 Myrmecinae Aphaenogaster tennessensis KAU-09-0910-13 2 0 0 2 0.07 Formicinae Brachymyrmex depilis KAU-09-0725-6 0 0 0 0 0.00 Formicinae Brachymyrmex depilis KAU-09-0824-4 0 0 0 0 0.00 Formicinae Camponotus chromaiodes KAU-08-0424-3 1 0 1 0 0.43 Formicinae Camponotus chromaiodes KAU-09-0613-10 0 0 0 0 1.00 Formicinae Camponotus chromaiodes KAU-09-0910-8 5 2 3 0 0.61 Formicinae Camponotus nearcticus KAU-08-0812-1 0 0 0 0 0.05 Formicinae Camponotus subbarbatus KAU-09-0720-1 0 0 0 0 0.00 Formicinae Camponotus subbarbatus KAU-09-0910-10a 1 0 0 1 0.05 Myrmecinae Crematogaster cerasi KAU-09-0522-1 1 0 0 1 0.07

209 Myrmecinae Crematogaster cerasi KAU-09-0522-11 0 0 0 0 0.00 Myrmecinae Crematogaster cerasi KAU-09-0721-5 2 0 0 2 0.53 Myrmecinae Crematogaster cerasi KAU-09-0910-14 2 0 0 2 0.05 Myrmecinae Crematogaster lineolata KAU-08-0829-6 2 0 0 2 0.06 Myrmecinae Crematogaster lineolata KAU-08-0829-7 0 0 0 0 0.03 Formicinae Formica exectoides KAU-08-0718-2 0 0 0 0 0.00 Formicinae Formica exectoides KAU-09-0613-35 7 5 0 2 1.00 Formicinae Formica exectoides KAU-09-0613-36a 2 1 0 1 0.97 Formicinae Formica exectoides KAU-09-0613-37 4 1 0 3 0.90 Formicinae Formica exectoides KAU-09-0613-38 6 3 1 2 1.00 Formicinae Formica exectoides KAU-09-0725-14 2 1 0 1 0.67 Formicinae Formica exectoides KAU-09-0725-15 3 2 1 0 0.44 Formicinae Formica exectoides KAU-09-0909-27 0 0 0 0 0.00 Formicinae Formica exectoides KAU-09-0909-28 0 0 0 0 0.00 Continued

Table 37 continued SUBFAMILY GENUS SPECIES FIELD # # PHORETIC # MESO # HETERO # ASTIG PREV WITH Formicinae Formica exectoides KAU-09-0909-29 0 0 0 0 0.00 Formicinae Formica integra KAU-09-0516-6a 2 2 0 0 0.56 Formicinae Formica nitidiventris KAU-09-0824-17a 2 1 0 1 0.59 Formicinae Formica subsericea KAU-09-0824-14a 1 0 1 0 0.27 Formicinae Formica subsericea KAU-09-0824-18 3 1 1 1 0.88 Formicinae Formica subsericea KAU-09-0909-1 4 1 3 0 0.07 Formicinae Lasius alienus KAU-08-0610-5 0 0 0 0 0.03 Formicinae Lasius alienus KAU-08-0611-3 0 0 0 0 0.59 Formicinae Lasius alienus KAU-08-0611-4 2 1 0 1 0.88 Formicinae Lasius alienus KAU-08-0706-15 4 1 2 1 0.27 Formicinae Lasius alienus KAU-08-0829-11 2 0 1 1 1.00 Formicinae Lasius alienus KAU-08-0903-10 2 0 1 1 0.31

210 Formicinae Lasius alienus KAU-08-0903-12 0 0 0 0 0.01 Formicinae Lasius alienus KAU-09-0424-15 6 2 3 1 0.92 Formicinae Lasius alienus KAU-09-0424-18 5 0 3 2 0.71 Formicinae Lasius alienus KAU-09-0612-2 3 0 1 2 0.65 Formicinae Lasius alienus KAU-09-0613-8 3 0 2 1 0.28 Formicinae Lasius alienus KAU-09-0707-2 2 0 1 1 0.19 Formicinae Lasius alienus KAU-09-0824-2 2 0 1 1 0.50 Formicinae Lasius alienus KAU-09-0910-9 3 0 1 2 0.41 Formicinae Lasius nearcticus KAU-08-0828-3 1 0 1 0 0.17 Formicinae Lasius neoniger KAU-08-0829-13 2 0 0 2 0.36 Formicinae Lasius neoniger KAU-08-0916-1 3 1 0 2 0.29 Formicinae Lasius neoniger KAU-09-0424-1a 7 1 3 3 0.17 Formicinae Lasius neoniger KAU-09-0525-2 2 1 0 1 0.13 Formicinae Lasius neoniger KAU-09-0613-29 1 0 0 1 0.13 Continued

Table 37 continued SUBFAMILY GENUS SPECIES FIELD # # PHORETIC # MESO # HETERO # ASTIG PREV WITH Formicinae Lasius neoniger KAU-09-0613-30 2 0 1 1 0.50 Formicinae Lasius neoniger KAU-09-0824-13 3 1 1 1 0.10 Formicinae Lasius neoniger KAU-09-0824-23a 4 1 0 3 0.08 Formicinae Lasius neoniger KAU-09-0909-15 2 0 0 2 0.33 Formicinae Lasius speculiventris KAU-09-0725-1 10 1 5 4 0.57 Formicinae Lasius umbratus KAU-08-0611-2 4 0 1 3 1.00 Formicinae Lasius umbratus KAU-08-0829-8a &b 8 1 3 4 0.63 Formicinae Lasius umbratus KAU-09-0613-2 8 1 4 3 0.97 Formicinae Lasius umbratus KAU-09-0613-22 8 2 4 2 0.89 Formicinae Lasius umbratus KAU-09-0613-5 11 1 7 3 1.00 Formicinae Lasius umbratus KAU-09-0909-11 7 1 3 3 0.55 Formicinae Lasius umbratus KAU-09-0909-9a 6 2 2 2 0.63

211 Formicinae Lasius umbratus KAU-09-0910-2 19 2 9 8 0.91 Formicinae Lasius umbratus KAU-09-0910-4 12 1 5 6 0.46 Formicinae Lasius umbratus KAU-09-0910-5 8 1 3 4 0.70 Formicinae Lasius umbratus KAU-09-0910-7 8 2 3 3 0.53 Formicinae Lasius (A.) claviger KAU-08-0529-3 10 0 2 8 0.74 Formicinae Lasius (A.) claviger KAU-09-0721-10 11 1 4 6 0.71 Formicinae Lasius (A.) claviger KAU-09-0909-12 7 3 3 1 0.29 Myrmecinae Monomorium minimum KAU-09-0516-5 0 0 0 0 0.00 Myrmecinae Monomorium minimum KAU-09-0909-22 0 0 0 0 0.00 Myrmecinae Myrmecina americana KAU-09-0909-9b 1 0 0 1 0.40 Myrmecinae Myrmica americana KAU-09-0424-4 1 0 0 1 0.41 Myrmecinae Myrmica punctiventris KAU-09-0707-10 0 0 0 0 0.00 Myrmecinae Myrmica punctiventris KAU-09-0707-11 0 0 0 0 0.00 Formicinae Paratrechina faisonensis KAU-08-0722-1 0 0 0 0 0.00 Continued

Table 37 continued SUBFAMILY GENUS SPECIES FIELD # # PHORETIC # MESO # HETERO # ASTIG PREV WITH Formicinae Paratrechina faisonensis KAU-09-0522-2 0 0 0 0 0.00 Formicinae Paratrechina faisonensis KAU-09-0522-3a 0 0 0 0 0.00 Formicinae Paratrechina parvula KAU-09-0909-21a 0 0 0 0 0.00 Myrmecinae Pheidole bicarinata KAU-09-0909-19 0 0 0 0 0.00 Ponerinae Ponera pennsylvanica KAU-08-0706-10 0 0 0 0 0.00 Ponerinae Ponera pennsylvanica KAU-08-0829-10 0 0 0 0 0.00 Ponerinae Ponera pennsylvanica KAU-09-0522-9 1 0 1 0 0.00 Ponerinae Ponera pennsylvanica KAU-09-0613-13 2 0 2 0 0.00 Ponerinae Ponera pennsylvanica KAU-09-0613-19 0 0 0 0 0.00 Ponerinae Ponera pennsylvanica KAU-09-0725-12 0 0 0 0 0.17 Formicinae Prenolepis imparis KAU-10-0316-1 2 0 1 1 0.26 Formicinae Prenolepis imparis KAU-10-0327-1 4 0 2 2 0.11

212 Myrmecinae Pyramica ohioensis KAU-08-0706-7 0 0 0 0 0.00 Myrmecinae Pyramica ohioensis KAU-08-0903-3 1 0 0 1 0.02 Myrmecinae Pyramica reflexa KAU-09-0522-6 0 0 0 0 0.00 Myrmecinae Solenopsis molesta KAU-09-0516-7 1 0 1 0 0.02 Myrmecinae Solenopsis molesta KAU-09-0516-8b 0 0 0 0 0.00 Myrmecinae Solenopsis molesta KAU-09-0516-9 0 0 0 0 0.00 Myrmecinae Solenopsis molesta KAU-09-0612-3 1 0 0 1 0.02 Myrmecinae Solenopsis molesta KAU-09-0613-12 1 0 1 0 0.09 Myrmecinae Solenopsis molesta KAU-09-0613-36b 3 0 2 1 0.10 Myrmecinae Solenopsis molesta KAU-09-0721-2 1 0 1 0 0.03 Myrmecinae Solenopsis molesta KAU-09-0824-14b 0 0 0 0 0.00 Myrmecinae Solenopsis molesta KAU-09-0824-15b 1 0 0 1 0.05 Myrmecinae Solenopsis molesta KAU-09-0824-17b 2 0 2 0 0.02 Myrmecinae Solenopsis molesta KAU-09-0824-19 0 0 0 0 0.05 Continued

Table 37 continued SUBFAMILY GENUS SPECIES FIELD # # PHORETIC # MESO # HETERO # ASTIG PREV WITH Myrmecinae Solenopsis molesta KAU-09-0824-23b 3 0 1 2 0.25 Myrmecinae Solenopsis molesta KAU-09-0909-21b 1 0 1 0 0.02 Dolichoderinae Tapinoma sessile KAU-09-0511-1 0 0 0 0 0.00 Dolichoderinae Tapinoma sessile KAU-09-0518-1 0 0 0 0 0.00 Dolichoderinae Tapinoma sessile KAU-09-0612-5 0 0 0 0 0.00 Dolichoderinae Tapinoma sessile KAU-09-0613-1 2 0 1 1 0.06 Dolichoderinae Tapinoma sessile KAU-09-0613-9 0 0 0 0 0.00 Dolichoderinae Tapinoma sessile KAU-09-0909-16 0 0 0 0 0.00 Dolichoderinae Tapinoma sessile KAU-09-0909-17 0 0 0 0 0.00 Dolichoderinae Tapinoma sessile KAU-09-0909-18 0 0 0 0 0.00 Dolichoderinae Tapinoma sessile KAU-09-0909-2 0 0 0 0 0.00 Myrmecinae Tetramorium caespitum KAU-08-0609-1 0 0 0 0 0.00

213 Myrmecinae Tetramorium caespitum KAU-08-0611-1 0 0 0 0 0.00 Myrmecinae Tetramorium caespitum KAU-08-0620-1 0 0 0 0 0.04 Myrmecinae Tetramorium caespitum KAU-08-0625-1 1 0 0 1 0.07 Myrmecinae Tetramorium caespitum KAU-08-0625-2 0 0 0 0 0.57 Myrmecinae Tetramorium caespitum KAU-08-0707-1a 1 1 0 0 0.08 Myrmecinae Tetramorium caespitum KAU-08-0707-3a 1 1 0 0 0.13 Myrmecinae Tetramorium caespitum KAU-09-0608-1 2 0 1 1 0.39 Myrmecinae Tetramorium caespitum KAU-09-0824-15a 1 1 0 0 0.04 Myrmecinae Tetramorium caespitum KAU-09-0824-8 4 1 1 2 0.06 Myrmecinae Tetramorium caespitum KAU-09-0824-9 1 0 1 0 0.05

SUB PARA COLONY HOST NEST TOTAL SUB FAMILY FAM GENUS SPECIES FIELD # SITIC LASIUS SIZE SIZE HABITAT SUBSTRATE ANTS Amblyoponinae 5 Amblyopone pallipes KAU-08-0828-2 2 2 1 5.5 1 1 10 Myrmecinae 2 Aphaenogaster fulva KAU-08-0610-10 2 2 2 5.5 1 1 23 Myrmecinae 2 Aphaenogaster fulva KAU-08-0706-16 2 2 2 5.5 1 1 34 Myrmecinae 2 Aphaenogaster fulva KAU-08-0815-1 2 2 2 5.5 1 1 60 Myrmecinae 2 Aphaenogaster fulva KAU-09-0613-17 2 2 2 5.5 1 1 41 Myrmecinae 2 Aphaenogaster fulva KAU-09-0613-21a 2 2 2 5.5 1 1 24 Myrmecinae 2 Aphaenogaster fulva KAU-09-0707-5 2 2 2 5.5 1 1 68 Myrmecinae 2 Aphaenogaster fulva KAU-09-0707-6 2 2 2 5.5 1 1 77 Myrmecinae 2 Aphaenogaster fulva KAU-09-0721-11 2 2 2 5.5 1 1 62 Myrmecinae 2 Aphaenogaster fulva KAU-09-0721-12 2 2 2 5.5 1 1 41 Myrmecinae 2 Aphaenogaster fulva KAU-09-0721-14 2 2 2 5.5 1 1 51 Myrmecinae 2 Aphaenogaster fulva KAU-09-0721-15 2 2 2 5.5 1 1 53 Myrmecinae 2 Aphaenogaster fulva KAU-09-0721-9 2 2 2 5.5 1 1 40 Myrmecinae 2 Aphaenogaster picea complex KAU-08-0531-4 2 2 2 5.1 1 1 69 Myrmecinae 2 Aphaenogaster picea complex KAU-08-0610-7 2 2 2 5.1 1 1 31 214 Myrmecinae 2 Aphaenogaster picea complex KAU-08-0829-5 2 2 2 5.1 1 1 30

Myrmecinae 2 Aphaenogaster picea complex KAU-08-0829-9 2 2 2 5.1 1 1 30 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0424-17 2 2 2 5.1 1 1 24 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0613-18 2 2 2 5.1 1 1 31 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0613-20 2 2 2 5.1 1 1 33 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0613-32 2 2 2 5.1 1 1 27 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0725-10 2 2 2 5.1 1 1 49 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0725-11 2 2 2 5.1 1 1 57 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0725-13 2 2 2 5.1 1 1 61 Continued Table 38. Raw data for Predictors used in Statistical Analyses. Codes used for predictors are as follows: Subfamily- 1=Formicinae, 2=Myrmecinae, 3=Dolichoderinae, 4=Ponerinae, 5=Amblyoponerinae. Parasitic: 1= Parasitic, 2=non parasitic. Lasius: 1=Lasius, 2=Not Lasius. Colony size: 1=small, 2=moderate, 3=large. Habitat: 1=woods, 2=open. Nest substrate: 1=wood, 2=soil.

Table 38 continued

SUB PARA COLONY HOST NEST TOTAL SUB FAMILY FAM GENUS SPECIES FIELD # SITIC LASIUS SIZE SIZE HABITAT SUBSTRATE ANTS Myrmecinae 2 Aphaenogaster picea complex KAU-09-0725-2 2 2 2 5.1 1 1 81 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0725-3 2 2 2 5.1 1 2 38 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0725-4 2 2 2 5.1 1 1 37 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0725-5 2 2 2 5.1 1 1 45 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0725-7 2 2 2 5.1 1 1 51 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0725-8 2 2 2 5.1 1 1 89 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0725-9 2 2 2 5.1 1 1 56 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0824-6 2 2 2 5.1 1 1 32 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0910-11 2 2 2 5.1 1 1 36 Myrmecinae 2 Aphaenogaster picea complex KAU-09-0910-12 2 2 2 5.1 1 1 30

215 Myrmecinae 2 Aphaenogaster rudis complex KAU-08-0706-20 2 2 2 5.15 1 1 34 Myrmecinae 2 Aphaenogaster rudis complex KAU-08-0706-31 2 2 2 5.15 1 1 36

Myrmecinae 2 Aphaenogaster rudis complex KAU-08-0706-8 2 2 2 5.15 1 1 30 Myrmecinae 2 Aphaenogaster rudis complex KAU-09-0424-19 2 2 2 5.15 1 1 31 Myrmecinae 2 Aphaenogaster rudis complex KAU-09-0424-6 2 2 2 5.15 1 1 20 Myrmecinae 2 Aphaenogaster rudis complex KAU-09-0613-11 2 2 2 5.15 1 1 40 Myrmecinae 2 Aphaenogaster rudis complex KAU-09-0613-15 2 2 2 5.15 1 1 40 Myrmecinae 2 Aphaenogaster rudis complex KAU-09-0613-24 2 2 2 5.15 1 1 32 Myrmecinae 2 Aphaenogaster rudis complex KAU-09-0613-25 2 2 2 5.15 1 1 29 Myrmecinae 2 Aphaenogaster rudis complex KAU-09-0613-31 2 2 2 5.15 1 1 37 Myrmecinae 2 Aphaenogaster rudis complex KAU-09-0613-4 2 2 2 5.15 1 1 33 Myrmecinae 2 Aphaenogaster rudis complex KAU-09-0613-45 2 2 2 5.15 1 1 56 Continued

Table 38 continued SUB PARA COLONY HOST NEST TOTAL SUBFAMILY FAM GENUS SPECIES FIELD # SITIC LASIUS SIZE SIZE HABITAT SUBSTRATE ANTS Myrmecinae 2 Aphaenogaster rudis complex KAU-09-0721-8 2 2 2 5.15 1 1 50 Myrmecinae 2 Aphaenogaster tennessensis KAU-08-0610-1 1 2 3 5.75 1 1 67 Myrmecinae 2 Aphaenogaster tennessensis KAU-08-0706-11 1 2 3 5.75 1 1 37 Myrmecinae 2 Aphaenogaster tennessensis KAU-08-0812-3 1 2 3 5.75 1 1 30 Myrmecinae 2 Aphaenogaster tennessensis KAU-09-0707-8 1 2 3 5.75 1 1 62 Myrmecinae 2 Aphaenogaster tennessensis KAU-09-0910-13 1 2 3 5.75 1 1 30 Formicinae 1 Brachymyrmex depilis KAU-09-0725-6 2 2 1 1.3 1 2 40 Formicinae 1 Brachymyrmex depilis KAU-09-0824-4 2 2 1 1.3 1 2 22 Formicinae 1 Camponotus chromaiodes KAU-08-0424-3 2 2 3 9.2 1 1 58 Formicinae 1 Camponotus chromaiodes KAU-09-0613-10 2 2 3 9.2 1 1 33 Formicinae 1 Camponotus chromaiodes KAU-09-0910-8 2 2 3 9.2 1 1 69 216 Formicinae 1 Camponotus nearcticus KAU-08-0812-1 2 2 1 5.25 1 1 22

Formicinae 1 Camponotus subbarbatus KAU-09-0720-1 2 2 2 5.1 2 2 52 Formicinae 1 Camponotus subbarbatus KAU-09-0910-10a 2 2 2 5.1 1 1 21 Myrmecinae 2 Crematogaster cerasi KAU-09-0522-1 2 2 2 3.35 2 1 72 Myrmecinae 2 Crematogaster cerasi KAU-09-0522-11 2 2 2 3.35 2 1 51 Myrmecinae 2 Crematogaster cerasi KAU-09-0721-5 2 2 2 3.35 1 1 47 Myrmecinae 2 Crematogaster cerasi KAU-09-0910-14 2 2 2 3.35 1 1 55 Myrmecinae 2 Crematogaster lineolata KAU-08-0829-6 2 2 2 3.5 1 1 31 Myrmecinae 2 Crematogaster lineolata KAU-08-0829-7 2 2 2 3.5 1 1 30 Formicinae 1 Formica exectoides KAU-08-0718-2 1 2 3 6.7 2 2 30 Formicinae 1 Formica exectoides KAU-09-0613-35 1 2 3 6.7 2 2 24 Formicinae 1 Formica exectoides KAU-09-0613-36a 1 2 3 6.7 2 2 35 Formicinae 1 Formica exectoides KAU-09-0613-37 1 2 3 6.7 2 2 40 Continued

Table 38 continued SUB PARA COLONY HOST NEST TOTAL SUBFAMILY FAM GENUS SPECIES FIELD # SITIC LASIUS SIZE SIZE HABITAT SUBSTRATE ANTS Formicinae 1 Formica exectoides KAU-09-0613-38 1 2 3 6.7 2 2 30 Formicinae 1 Formica exectoides KAU-09-0725-14 1 2 3 6.7 2 2 30 Formicinae 1 Formica exectoides KAU-09-0725-15 1 2 3 6.7 2 2 48 Formicinae 1 Formica exectoides KAU-09-0909-27 1 2 3 6.7 2 2 40 Formicinae 1 Formica exectoides KAU-09-0909-28 1 2 3 6.7 2 2 40 Formicinae 1 Formica exectoides KAU-09-0909-29 1 2 3 6.7 2 2 40 Formicinae 1 Formica integra KAU-09-0516-6a 2 2 3 6.9 1 2 89 Formicinae 1 Formica nitidiventris KAU-09-0824-17a 2 2 3 5.65 2 2 37 Formicinae 1 Formica subsericea KAU-09-0824-14a 2 2 3 6.4 2 2 22 Formicinae 1 Formica subsericea KAU-09-0824-18 2 2 3 6.4 2 2 42 Formicinae 1 Formica subsericea KAU-09-0909-1 2 2 3 6.4 2 2 55

217 Formicinae 1 Lasius alienus KAU-08-0610-5 2 1 3 3.15 1 1 146

Formicinae 1 Lasius alienus KAU-08-0611-3 2 1 3 3.15 1 1 22 Formicinae 1 Lasius alienus KAU-08-0611-4 2 1 3 3.15 1 1 51 Formicinae 1 Lasius alienus KAU-08-0706-15 2 1 3 3.15 1 1 33 Formicinae 1 Lasius alienus KAU-08-0829-11 2 1 3 3.15 1 1 33 Formicinae 1 Lasius alienus KAU-08-0903-10 2 1 3 3.15 1 1 32 Formicinae 1 Lasius alienus KAU-08-0903-12 2 1 3 3.15 1 1 80 Formicinae 1 Lasius alienus KAU-09-0424-15 2 1 3 3.15 1 1 37 Formicinae 1 Lasius alienus KAU-09-0424-18 2 1 3 3.15 1 1 24 Formicinae 1 Lasius alienus KAU-09-0612-2 2 1 3 3.15 2 2 31 Formicinae 1 Lasius alienus KAU-09-0613-8 2 1 3 3.15 1 1 32 Formicinae 1 Lasius alienus KAU-09-0707-2 2 1 3 3.15 1 1 37 Formicinae 1 Lasius alienus KAU-09-0824-2 2 1 3 3.15 2 1 60 Formicinae 1 Lasius alienus KAU-09-0910-9 2 1 3 3.15 1 2 39 Continued

Table 38 continued SUB PARA COLONY NEST TOTAL SUBFAMILY FAM GENUS SPECIES FIELD # SITIC LASIUS SIZE HOST SIZE HABITAT SUBSTRATE ANTS Formicinae 1 Lasius nearcticus KAU-08-0828-3 2 1 1 3.25 1 1 145 Formicinae 1 Lasius neoniger KAU-08-0829-13 2 1 3 3.2 2 2 96 Formicinae 1 Lasius neoniger KAU-08-0916-1 2 1 3 3.2 2 2 45 Formicinae 1 Lasius neoniger KAU-09-0424-1a 2 1 3 3.2 2 2 36 Formicinae 1 Lasius neoniger KAU-09-0525-2 2 1 3 3.2 2 2 78 Formicinae 1 Lasius neoniger KAU-09-0613-29 2 1 3 3.2 2 2 46 Formicinae 1 Lasius neoniger KAU-09-0613-30 2 1 3 3.2 2 2 20 Formicinae 1 Lasius neoniger KAU-09-0824-13 2 1 3 3.2 2 2 40 Formicinae 1 Lasius neoniger KAU-09-0824-23a 2 1 3 3.2 2 2 39 Formicinae 1 Lasius neoniger KAU-09-0909-15 2 1 3 3.2 2 2 21 Formicinae 1 Lasius speculiventris KAU-09-0725-1 1 1 3 4.85 1 2 56

218 Formicinae 1 Lasius umbratus KAU-08-0611-2 1 1 3 4.4 1 1 25

Formicinae 1 Lasius umbratus KAU-08-0829-8a &b 1 1 3 4.4 2 1 99 Formicinae 1 Lasius umbratus KAU-09-0613-2 1 1 3 4.4 1 2 38 Formicinae 1 Lasius umbratus KAU-09-0613-22 1 1 3 4.4 1 1 36 Formicinae 1 Lasius umbratus KAU-09-0613-5 1 1 3 4.4 1 1 22 Formicinae 1 Lasius umbratus KAU-09-0909-11 1 1 3 4.4 1 1 75 Formicinae 1 Lasius umbratus KAU-09-0909-9a 1 1 3 4.4 1 2 40 Formicinae 1 Lasius umbratus KAU-09-0910-2 1 1 3 4.4 1 1 34 Formicinae 1 Lasius umbratus KAU-09-0910-4 1 1 3 4.4 1 1 97 Formicinae 1 Lasius umbratus KAU-09-0910-5 1 1 3 4.4 1 2 37 Formicinae 1 Lasius umbratus KAU-09-0910-7 1 1 3 4.4 1 1 74 Formicinae 1 Lasius claviger KAU-08-0529-3 1 1 3 4.1 1 1 34 Formicinae 1 Lasius claviger KAU-09-0721-10 1 1 3 4.1 1 1 68 Formicinae 1 Lasius claviger KAU-09-0909-12 1 1 3 4.1 1 2 73 Continued

Table 38 continued SUB PARA COLONY HOST NEST TOTAL SUBFAMILY FAM GENUS SPECIES FIELD # SITIC LASIUS SIZE SIZE HABITAT SUBSTRATE ANTS Myrmecinae 2 Monomorium minimum KAU-09-0516-5 2 2 3 1.75 1 1 61 Myrmecinae 2 Monomorium minimum KAU-09-0909-22 2 2 3 1.75 2 2 33 Myrmecinae 2 Myrmecina americana KAU-09-0909-9b 2 2 1 3.35 1 2 10 Myrmecinae 2 Myrmica americana KAU-09-0424-4 2 2 1 5.35 2 2 22 Myrmecinae 2 Myrmica punctiventris KAU-09-0707-10 2 2 1 4.95 1 1 13 Myrmecinae 2 Myrmica punctiventris KAU-09-0707-11 2 2 1 4.65 1 1 11 Formicinae 1 Paratrechina faisonensis KAU-08-0722-1 2 2 1 2.2 2 2 30 Formicinae 1 Paratrechina faisonensis KAU-09-0522-2 2 2 1 2.2 2 2 32 Formicinae 1 Paratrechina faisonensis KAU-09-0522-3a 2 2 1 2.2 2 2 32 Formicinae 1 Paratrechina parvula KAU-09-0909-21a 2 2 3 2.2 2 2 70 Myrmecinae 2 Pheidole bicarinata KAU-09-0909-19 2 2 1 2.575 2 2 29 219 Ponerinae 4 Ponera pennsylvanica KAU-08-0706-10 2 2 1 3.4 1 1 10

Ponerinae 4 Ponera pennsylvanica KAU-08-0829-10 2 2 1 3.4 1 1 13 Ponerinae 4 Ponera pennsylvanica KAU-09-0522-9 2 2 1 3.4 2 2 11 Ponerinae 4 Ponera pennsylvanica KAU-09-0613-13 2 2 1 3.4 1 1 17 Ponerinae 4 Ponera pennsylvanica KAU-09-0613-19 2 2 1 3.4 1 1 10 Ponerinae 4 Ponera pennsylvanica KAU-09-0725-12 2 2 1 3.4 1 1 12 Formicinae 1 Prenolepis imparis KAU-10-0316-1 2 2 2 3.6 2 2 72 Formicinae 1 Prenolepis imparis KAU-10-0327-1 2 2 2 3.6 2 2 85 Myrmecinae 2 Pyramica ohioensis KAU-08-0706-7 2 2 1 2.375 1 1 10 Myrmecinae 2 Pyramica ohioensis KAU-08-0903-3 2 2 1 2.375 1 1 43 Myrmecinae 2 Pyramica reflexa KAU-09-0522-6 2 2 1 2.175 2 2 13 Myrmecinae 2 Solenopsis molesta KAU-09-0516-7 2 2 3 1.65 2 2 85 Myrmecinae 2 Solenopsis molesta KAU-09-0516-8b 2 2 3 1.65 2 2 90 Myrmecinae 2 Solenopsis molesta KAU-09-0516-9 2 2 3 1.65 2 2 61 Myrmecinae 2 Solenopsis molesta KAU-09-0612-3 2 2 3 1.65 2 2 46 Continued

Table 38 continued SUB PARA COLONY HOST NEST TOTAL SUBFAMILY FAM GENUS SPECIES FIELD # SITIC LASIUS SIZE SIZE HABITAT SUBSTRATE ANTS Myrmecinae 2 Solenopsis molesta KAU-09-0613-12 2 2 3 1.65 1 1 23 Myrmecinae 2 Solenopsis molesta KAU-09-0613-36b 2 2 3 1.65 1 2 51 Myrmecinae 2 Solenopsis molesta KAU-09-0721-2 2 2 3 1.65 2 2 59 Myrmecinae 2 Solenopsis molesta KAU-09-0824-14b 2 2 3 1.65 2 2 71 Myrmecinae 2 Solenopsis molesta KAU-09-0824-15b 2 2 3 1.65 2 2 66 Myrmecinae 2 Solenopsis molesta KAU-09-0824-17b 2 2 3 1.65 2 2 119 Myrmecinae 2 Solenopsis molesta KAU-09-0824-19 2 2 3 1.65 2 2 44 Myrmecinae 2 Solenopsis molesta KAU-09-0824-23b 2 2 3 1.65 2 2 36 Myrmecinae 2 Solenopsis molesta KAU-09-0909-21b 2 2 3 1.65 2 2 49 Dolichoderinae 3 Tapinoma sessile KAU-09-0511-1 2 2 2 2.8 2 2 54 Dolichoderinae 3 Tapinoma sessile KAU-09-0518-1 2 2 2 2.8 2 2 73 220 Dolichoderinae 3 Tapinoma sessile KAU-09-0612-5 2 2 2 2.8 2 2 30

Dolichoderinae 3 Tapinoma sessile KAU-09-0613-1 2 2 2 2.8 2 2 50 Dolichoderinae 3 Tapinoma sessile KAU-09-0613-9 2 2 2 2.8 1 2 24 Dolichoderinae 3 Tapinoma sessile KAU-09-0909-16 2 2 2 2.8 2 2 22 Dolichoderinae 3 Tapinoma sessile KAU-09-0909-17 2 2 2 2.8 2 2 48 Dolichoderinae 3 Tapinoma sessile KAU-09-0909-18 2 2 2 2.8 2 2 51 Dolichoderinae 3 Tapinoma sessile KAU-09-0909-2 2 2 2 2.8 2 2 42 Myrmecinae 2 Tetramorium caespitum KAU-08-0609-1 2 2 3 3.55 2 2 20 Myrmecinae 2 Tetramorium caespitum KAU-08-0611-1 2 2 3 3.55 2 2 118 Myrmecinae 2 Tetramorium caespitum KAU-08-0620-1 2 2 3 3.55 2 2 76 Myrmecinae 2 Tetramorium caespitum KAU-08-0625-1 2 2 3 3.55 2 2 30 Myrmecinae 2 Tetramorium caespitum KAU-08-0625-2 2 2 3 3.55 2 2 44 Myrmecinae 2 Tetramorium caespitum KAU-08-0707-1a 2 2 3 3.55 2 2 157 Myrmecinae 2 Tetramorium caespitum KAU-08-0707-3a 2 2 3 3.55 2 2 380 Continued

Table 38 continued SUB PARA COLONY HOST NEST TOTAL SUBFAMILY FAM GENUS SPECIES FIELD # SITIC LASIUS SIZE SIZE HABITAT SUBSTRATE ANTS Myrmecinae 2 Tetramorium caespitum KAU-09-0608-1 2 2 3 3.55 2 2 124 Myrmecinae 2 Tetramorium caespitum KAU-09-0824-15a 2 2 3 3.55 2 2 82 Myrmecinae 2 Tetramorium caespitum KAU-09-0824-8 2 2 3 3.55 2 2 77 Myrmecinae 2 Tetramorium caespitum KAU-09-0824-9 2 2 3 3.55 2 2 58

221

TOTAL PREV TOTAL PREV W TOTAL PREV F TOTAL PREV M ANT GENUS ANT SPECIES FIELD # ANTS WITH W WITH F WITH M WITH Amblyopone pallipes KAU-08-0828-2 10 0.00 5 0.00 2 0.00 3 0.00 Aphaenogaster fulva KAU-08-0610-10 23 0.61 21 0.67 2 0.00 0 n/a Aphaenogaster fulva KAU-08-0706-16 34 0.35 34 0.35 0 n/a 0 n/a Aphaenogaster fulva KAU-08-0815-1 60 0.27 37 0.32 0 n/a 23 0.17 Aphaenogaster fulva KAU-09-0613-17 41 0.15 41 0.15 0 n/a 0 n/a Aphaenogaster fulva KAU-09-0613-21a 24 0.46 24 0.46 0 n/a 0 n/a Aphaenogaster fulva KAU-09-0707-5 68 0.09 68 0.09 0 n/a 0 n/a Aphaenogaster fulva KAU-09-0707-6 77 0.14 50 0.16 8 0.13 19 0.11 Aphaenogaster fulva KAU-09-0721-11 62 0.48 39 0.23 18 1.00 5 0.60 Aphaenogaster fulva KAU-09-0721-12 41 0.07 41 0.07 0 n/a 0 n/a Aphaenogaster fulva KAU-09-0721-14 51 0.35 35 0.46 2 1.00 14 0.00 Aphaenogaster fulva KAU-09-0721-15 53 0.43 39 0.23 13 1.00 1 1.00

222 Aphaenogaster fulva KAU-09-0721-9 40 0.20 40 0.20 0 n/a 0 n/a

Aphaenogaster picea complex KAU-08-0531-4 69 0.09 69 0.09 0 n/a 0 n/a Aphaenogaster picea complex KAU-08-0610-7 31 0.10 30 0.10 1 0.00 0 n/a Aphaenogaster picea complex KAU-08-0829-5 30 0.00 30 0.00 0 n/a 0 n/a Aphaenogaster picea complex KAU-08-0829-9 30 0.33 30 0.33 0 n/a 0 n/a Aphaenogaster picea complex KAU-09-0424-17 24 0.29 24 0.29 0 n/a 0 n/a Aphaenogaster picea complex KAU-09-0613-18 31 0.32 31 0.32 0 n/a 0 n/a Aphaenogaster picea complex KAU-09-0613-20 33 0.36 33 0.36 0 n/a 0 n/a Aphaenogaster picea complex KAU-09-0613-32 27 0.30 27 0.30 0 n/a 0 n/a Aphaenogaster picea complex KAU-09-0725-10 49 0.37 49 0.37 0 n/a 0 n/a Aphaenogaster picea complex KAU-09-0725-11 57 0.19 38 0.11 17 0.41 2 0.00 Continued Table 39. Raw data for Prevalence. Ant species used in statistical analyses of prevalence are shown. "W" designates worker, "F" designates female alate, "M" designates male alate. Mean (M), Standard deviation (SD), and number (N) are calculated in the last 3 rows of the table. “n/a” is used when no castes of a type were found in the nest.

Table 39 continued TOTAL PREV TOTAL PREV W TOTAL PREV F TOTAL PREV M ANT GENUS ANT SPECIES FIELD # ANTS WITH W WITH F WITH M WITH Aphaenogaster picea complex KAU-09-0725-13 61 0.15 60 0.13 1 1.00 0 n/a Aphaenogaster picea complex KAU-09-0725-2 81 0.09 45 0.11 0 n/a 36 0.06 Aphaenogaster picea complex KAU-09-0725-3 38 0.05 38 0.05 0 n/a 0 n/a Aphaenogaster picea complex KAU-09-0725-4 37 0.03 37 0.03 0 n/a 0 n/a Aphaenogaster picea complex KAU-09-0725-5 45 0.13 45 0.13 0 n/a 0 n/a Aphaenogaster picea complex KAU-09-0725-7 51 0.02 51 0.02 0 n/a 0 n/a Aphaenogaster picea complex KAU-09-0725-8 89 0.18 54 0.19 7 0.57 28 0.07 Aphaenogaster picea complex KAU-09-0725-9 56 0.11 50 0.04 6 0.67 0 n/a Aphaenogaster picea complex KAU-09-0824-6 32 0.16 32 0.16 0 n/a 0 n/a Aphaenogaster picea complex KAU-09-0910-11 36 0.11 36 0.11 0 n/a 0 n/a Aphaenogaster picea complex KAU-09-0910-12 30 0.10 30 0.10 0 n/a 0 n/a 223 Aphaenogaster rudiscomplex KAU-08-0706-20 34 0.44 34 0.44 0 n/a 0 n/a

Aphaenogaster rudiscomplex KAU-08-0706-31 36 0.11 35 0.11 0 n/a 0 n/a Aphaenogaster rudiscomplex KAU-08-0706-8 30 0.13 30 0.13 0 n/a 0 n/a Aphaenogaster rudiscomplex KAU-09-0424-19 31 0.81 31 0.81 0 n/a 0 n/a Aphaenogaster rudiscomplex KAU-09-0424-6 20 0.00 20 0.00 0 n/a 0 n/a Aphaenogaster rudiscomplex KAU-09-0613-11 40 0.05 40 0.05 0 n/a 0 n/a Aphaenogaster rudiscomplex KAU-09-0613-15 40 0.30 40 0.30 0 n/a 0 n/a Aphaenogaster rudiscomplex KAU-09-0613-24 32 0.44 32 0.44 0 n/a 0 n/a Aphaenogaster rudiscomplex KAU-09-0613-25 29 0.45 29 0.45 0 n/a 0 n/a Aphaenogaster rudiscomplex KAU-09-0613-31 37 0.76 21 0.71 7 1.00 9 0.67 Aphaenogaster rudiscomplex KAU-09-0613-4 33 0.39 33 0.39 0 n/a 0 n/a Aphaenogaster rudiscomplex KAU-09-0613-45 56 0.32 41 0.37 0 n/a 15 0.20 Aphaenogaster rudiscomplex KAU-09-0721-8 50 0.20 50 0.20 0 n/a 0 n/a Aphaenogaster tennessensis KAU-08-0610-1 67 0.63 34 0.56 33 0.70 0 n/a Continued

Table 39 continued TOTAL PREV TOTAL PREV W TOTAL PREV F TOTAL PREV M ANT GENUS ANT SPECIES FIELD # ANTS WITH W WITH F WITH M WITH Aphaenogaster tennessensis KAU-08-0706-11 37 0.49 35 0.51 0 n/a 2 0.00 Aphaenogaster tennessensis KAU-08-0812-3 30 0.70 30 0.70 0 n/a 0 n/a Aphaenogaster tennessensis KAU-09-0707-8 62 0.10 30 0.17 0 n/a 32 0.03 Aphaenogaster tennessensis KAU-09-0910-13 30 0.07 30 0.07 0 n/a 0 n/a Brachymyrmex depilis KAU-09-0725-6 40 0.00 40 0.00 0 n/a 0 n/a Brachymyrmex depilis KAU-09-0824-4 22 0.00 21 0.00 0 n/a 1 0.00 Camponotus chromaiodes KAU-08-0424-3 58 0.43 0 n/a 57 0.44 1 0.00 Camponotus chromaiodes KAU-09-0613-10 33 1.00 33 1.00 0 n/a 0 n/a Camponotus chromaiodes KAU-09-0910-8 69 0.61 50 0.58 14 0.93 5 0.00 Camponotus nearcticus KAU-08-0812-1 22 0.05 10 0.00 11 0.09 1 0.00 Camponotus subbarbatus KAU-09-0720-1 52 0.00 52 0.00 0 n/a 0 n/a 224 Camponotus subbarbatus KAU-09-0910-10a 21 0.05 21 0.05 0 n/a 0 n/a

Crematogaster cerasi KAU-09-0522-1 72 0.07 72 0.07 0 n/a 0 n/a Crematogaster cerasi KAU-09-0522-11 51 0.00 51 0.00 0 n/a 0 n/a Crematogaster cerasi KAU-09-0721-5 47 0.53 47 0.53 0 n/a 0 n/a Crematogaster cerasi KAU-09-0910-14 55 0.05 55 0.05 0 n/a 0 n/a Crematogaster lineolata KAU-08-0829-6 31 0.06 31 0.06 0 n/a 0 n/a Crematogaster lineolata KAU-08-0829-7 30 0.03 30 0.03 0 n/a 0 n/a Formica exectoides KAU-08-0718-2 30 0.00 30 0.00 0 n/a 0 n/a Formica exectoides KAU-09-0613-35 24 1.00 24 1.00 0 n/a 0 n/a Formica exectoides KAU-09-0613-36a 35 0.97 35 0.97 0 n/a 0 n/a Formica exectoides KAU-09-0613-37 40 0.90 40 0.90 0 n/a 0 n/a Formica exectoides KAU-09-0613-38 30 1.00 30 1.00 0 n/a 0 n/a Formica exectoides KAU-09-0725-14 30 0.67 30 0.67 0 n/a 0 n/a Formica exectoides KAU-09-0725-15 48 0.44 48 0.44 0 n/a 0 n/a Continued

Table 39 continued TOTAL PREV TOTAL PREV W TOTAL PREV F TOTAL PREV M ANT GENUS ANT SPECIES FIELD # ANTS WITH W WITH F WITH M WITH Formica exectoides KAU-09-0909-27 40 0.00 40 0.00 0 n/a 0 n/a Formica exectoides KAU-09-0909-28 40 0.00 40 0.00 0 n/a 0 n/a Formica exectoides KAU-09-0909-29 40 0.00 40 0.00 0 n/a 0 n/a Formica integra KAU-09-0516-6a 89 0.56 89 0.56 0 n/a 0 n/a Formica nitidiventris KAU-09-0824-17a 37 0.59 37 0.59 0 n/a 0 n/a Formica subsericea KAU-09-0824-14a 22 0.27 22 0.27 0 n/a 0 n/a Formica subsericea KAU-09-0824-18 42 0.88 42 0.88 0 n/a 0 n/a Formica subsericea KAU-09-0909-1 55 0.07 55 0.07 0 n/a 0 n/a Lasius alienus KAU-08-0610-5 146 0.03 146 0.03 0 n/a 0 n/a Lasius alienus KAU-08-0611-3 22 0.59 21 0.62 0 n/a 1 0.00 Lasius alienus KAU-08-0611-4 51 0.88 51 0.88 0 n/a 0 n/a 225 Lasius alienus KAU-08-0706-15 33 0.27 33 0.27 0 n/a 0 n/a

Lasius alienus KAU-08-0829-11 33 1.00 33 1.00 0 n/a 0 n/a Lasius alienus KAU-08-0903-10 32 0.31 32 0.31 0 n/a 0 n/a Lasius alienus KAU-08-0903-12 80 0.01 80 0.01 0 n/a 0 n/a Lasius alienus KAU-09-0424-15 37 0.92 37 0.92 0 n/a 0 n/a Lasius alienus KAU-09-0424-18 24 0.71 24 0.71 0 n/a 0 n/a Lasius alienus KAU-09-0612-2 31 0.65 31 0.65 0 n/a 0 n/a Lasius alienus KAU-09-0613-8 32 0.28 23 0.39 0 n/a 9 0.00 Lasius alienus KAU-09-0707-2 37 0.19 37 0.19 0 n/a 0 n/a Lasius alienus KAU-09-0824-2 60 0.50 60 0.50 0 n/a 0 n/a Lasius alienus KAU-09-0910-9 39 0.41 39 0.41 0 n/a 0 n/a Lasius nearcticus KAU-08-0828-3 145 0.17 51 0.06 45 0.22 49 0.24 Lasius neoniger KAU-08-0829-13 96 0.36 16 0.25 41 0.73 39 0.03 Lasius neoniger KAU-08-0916-1 45 0.29 45 0.29 0 n/a 0 n/a Lasius neoniger KAU-09-0424-1a 36 0.17 36 0.17 0 n/a 0 n/a Continued

Table 39 continued TOTAL PREV TOTAL PREV W TOTAL PREV F TOTAL PREV M ANT GENUS ANT SPECIES FIELD # ANTS WITH W WITH F WITH M WITH Lasius neoniger KAU-09-0525-2 78 0.13 78 0.13 0 n/a 0 n/a Lasius neoniger KAU-09-0613-29 46 0.13 44 0.14 0 n/a 2 0.00 Lasius neoniger KAU-09-0613-30 20 0.50 18 0.50 0 n/a 2 0.50 Lasius neoniger KAU-09-0824-13 40 0.10 40 0.10 0 n/a 0 n/a Lasius neoniger KAU-09-0824-23a 39 0.08 39 0.08 0 n/a 0 n/a Lasius neoniger KAU-09-0909-15 21 0.33 21 0.33 0 n/a 0 n/a Lasius speculiventris KAU-09-0725-1 56 0.57 55 0.58 0 n/a 1 0.00 Lasius umbratus KAU-08-0611-2 25 1.00 25 1.00 0 n/a 0 n/a Lasius umbratus KAU-08-0829-8a &b 99 0.63 65 0.86 3 0.67 28 0.14 Lasius umbratus KAU-09-0613-2 38 0.97 38 0.97 0 n/a 0 n/a Lasius umbratus KAU-09-0613-22 36 0.89 36 0.89 0 n/a 0 n/a 226 Lasius umbratus KAU-09-0613-5 22 1.00 22 1.00 0 n/a 0 n/a

Lasius umbratus KAU-09-0909-11 75 0.55 31 0.87 44 0.32 0 n/a Lasius umbratus KAU-09-0909-9a 40 0.63 40 0.63 0 n/a 0 n/a Lasius umbratus KAU-09-0910-2 34 0.91 34 0.91 0 n/a 0 n/a Lasius umbratus KAU-09-0910-4 97 0.46 37 0.65 29 0.48 31 0.23 Lasius umbratus KAU-09-0910-5 37 0.70 37 0.70 0 n/a 0 n/a Lasius umbratus KAU-09-0910-7 74 0.53 37 0.68 17 0.53 20 0.25 Lasius (A.) claviger KAU-08-0529-3 34 0.74 34 0.74 0 n/a 0 n/a Lasius (A.) claviger KAU-09-0721-10 68 0.71 68 0.71 0 n/a 0 n/a Lasius (A.) claviger KAU-09-0909-12 73 0.29 46 0.39 0 n/a 27 0.11 Monomorium minimum KAU-09-0516-5 61 0.00 61 0.00 0 n/a 0 n/a Monomorium minimum KAU-09-0909-22 33 0.00 33 0.00 0 n/a 0 n/a Myrmecina americana KAU-09-0909-9b 10 0.40 9 0.44 0 n/a 1 0.00 Myrmica americana KAU-09-0424-4 22 0.41 22 0.41 0 n/a 0 n/a Myrmica punctiventris KAU-09-0707-10 13 0.00 13 0.00 0 n/a 0 n/a Continued

Table 39 continued TOTAL PREV TOTAL PREV W TOTAL PREV F TOTAL PREV M ANT GENUS ANT SPECIES FIELD # ANTS WITH W WITH F WITH M WITH Myrmica punctiventris KAU-09-0707-11 11 0.00 11 0.00 0 n/a 0 n/a Paratrechina faisonensis KAU-08-0722-1 30 0.00 30 0.00 0 n/a 0 n/a Paratrechina faisonensis KAU-09-0522-2 32 0.00 32 0.00 0 n/a 0 n/a Paratrechina faisonensis KAU-09-0522-3a 32 0.00 32 0.00 0 n/a 0 n/a Paratrechina parvula KAU-09-0909-21a 70 0.00 67 0.00 3 0.00 0 n/a Pheidole bicarinata KAU-09-0909-19 29 0.00 29 0.00 0 n/a 0 n/a Ponera pennsylvanica KAU-08-0706-10 10 0.00 10 0.00 0 n/a 0 n/a Ponera pennsylvanica KAU-08-0829-10 13 0.00 13 0.00 0 n/a 0 n/a Ponera pennsylvanica KAU-09-0522-9 11 0.00 10 0.00 0 n/a 0 n/a Ponera pennsylvanica KAU-09-0613-13 17 0.00 17 0.00 0 n/a 0 n/a Ponera pennsylvanica KAU-09-0613-19 10 0.00 10 0.00 0 n/a 0 n/a 227 Ponera pennsylvanica KAU-09-0725-12 12 0.17 12 0.17 0 n/a 0 n/a

Prenolepis imparis KAU-10-0316-1 72 0.26 36 0.14 0 n/a 36 0.39 Prenolepis imparis KAU-10-0327-1 85 0.11 63 0.13 4 0.00 18 0.06 Pyramica ohioensis KAU-08-0706-7 10 0.00 8 0.00 0 n/a 0 n/a Pyramica ohioensis KAU-08-0903-3 43 0.02 32 0.00 11 0.09 0 n/a Pyramica reflexa KAU-09-0522-6 13 0.00 13 0.00 0 n/a 0 n/a Solenopsis molesta KAU-09-0516-7 85 0.02 85 0.02 0 n/a 0 n/a Solenopsis molesta KAU-09-0516-8b 90 0.00 90 0.00 0 n/a 0 n/a Solenopsis molesta KAU-09-0516-9 61 0.00 61 0.00 0 n/a 0 n/a Solenopsis molesta KAU-09-0612-3 46 0.02 46 0.02 0 n/a 0 n/a Solenopsis molesta KAU-09-0613-12 23 0.09 23 0.09 0 n/a 0 n/a Solenopsis molesta KAU-09-0613-36b 51 0.10 51 0.10 0 n/a 0 n/a Solenopsis molesta KAU-09-0721-2 59 0.03 59 0.03 0 n/a 0 n/a Solenopsis molesta KAU-09-0824-14b 71 0.00 71 0.00 0 n/a 0 n/a Solenopsis molesta KAU-09-0824-15b 66 0.05 46 0.02 20 0.10 0 n/a Continued

Table 39 continued TOTAL PREV TOTAL PREV W TOTAL PREV F TOTAL PREV M ANT GENUS ANT SPECIES FIELD # ANTS WITH W WITH F WITH M WITH Solenopsis molesta KAU-09-0824-17b 119 0.02 119 0.02 0 n/a 0 n/a Solenopsis molesta KAU-09-0824-19 44 0.05 44 0.05 0 n/a 0 n/a Solenopsis molesta KAU-09-0824-23b 36 0.25 34 0.24 2 0.50 0 n/a Solenopsis molesta KAU-09-0909-21b 49 0.02 49 0.02 0 n/a 0 n/a Tapinoma sessile KAU-09-0511-1 54 0.00 53 0.00 0 n/a 0 n/a Tapinoma sessile KAU-09-0518-1 73 0.00 61 0.00 0 n/a 0 n/a Tapinoma sessile KAU-09-0612-5 30 0.00 30 0.00 0 n/a 0 n/a Tapinoma sessile KAU-09-0613-1 50 0.06 50 0.06 0 n/a 0 n/a Tapinoma sessile KAU-09-0613-9 24 0.00 24 0.00 0 n/a 0 n/a Tapinoma sessile KAU-09-0909-16 22 0.00 22 0.00 0 n/a 0 n/a Tapinoma sessile KAU-09-0909-17 48 0.00 45 0.00 0 n/a 3 0.00 228 Tapinoma sessile KAU-09-0909-18 51 0.00 45 0.00 0 n/a 6 0.00

Tapinoma sessile KAU-09-0909-2 42 0.00 42 0.00 0 n/a 0 n/a Tetramorium caespitum KAU-08-0609-1 20 0.00 0 n/a 0 n/a 20 0.00 Tetramorium caespitum KAU-08-0611-1 118 0.00 50 0.00 10 0.00 58 0.00 Tetramorium caespitum KAU-08-0620-1 76 0.04 0 n/a 6 0.33 4 0.25 Tetramorium caespitum KAU-08-0625-1 30 0.07 0 n/a 5 0.20 2 0.50 Tetramorium caespitum KAU-08-0625-2 44 0.57 31 0.39 3 1.00 10 1.00 Tetramorium caespitum KAU-08-0707-1a 157 0.08 30 0.13 80 0.05 45 0.11 Tetramorium caespitum KAU-08-0707-3a 380 0.13 351 0.12 7 0.43 22 0.09 Tetramorium caespitum KAU-09-0608-1 124 0.39 87 0.13 12 1.00 25 1.00 Tetramorium caespitum KAU-09-0824-15a 82 0.04 82 0.04 0 n/a 0 n/a Tetramorium caespitum KAU-09-0824-8 77 0.06 77 0.06 0 n/a 0 n/a Tetramorium caespitum KAU-09-0824-9 58 0.05 58 0.05 0 n/a 0 n/a MEAN 48.29 0.27 40.81 0.27 3.09 0.46 3.75 0.19 STDEV 36.39 0.30 31.22 0.31 10.19 0.37 10.08 0.29 N 8450 175 7142 173 541 34 656 41 Ω