Geomys Bursarius) Burrow Systems

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2021

Arthropod fauna of Plains pocket gopher (Geomys bursarius) burrow systems

Brett Lyle Gourley University of Northern Iowa

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BRETT GOURLEY

2021

ARTHROPOD FAUNA OF PLAINS POCKET GOPHER (GEOMYS BURSARIUS)

BURROW SYSTEMS

An Abstract of a Thesis

Submitted

in Partial Fulfillment

of the requirements for the Degree

Master of Science

Brett Lyle Gourley

University of Northern Iowa

May 2021

ABSTRACT

Pocket gophers are among the most asocial mammals, living a solitary existence except for mating and the rearing of offspring. They create expansive burrow systems underground while foraging for food and rarely surfacing above ground. As gophers burrow, excess soil is pushed to the surface or placed in older tunnels. This behavior creates a cave-like ecosystem where pocket gophers and associated arthropods live.

Few studies have been done on the arthropod fauna cohabitating with pocket gophers in these burrows. Most of these studies have focused on a family of arthropods over several sites. Many have focused on the southern United States, with few studies done in the Midwest (Minnesota, Nebraska, Wisconsin, Indiana).

I examined the arthropods living in Geomys bursarius burrow systems in northeast Iowa. I located a local farm with land in Conservation Reserve Program with a

Plains pocket gopher population. There were two prairies with G. bursarius, and samples were taken from burrow systems in both during late July 2015. At each burrow system, I placed a control cup above ground and 3 cups inside the burrow system as a pitfall trap with molasses as bait. In Prairie 1, five G. bursarius burrow systems were sampled, yielding a total of 20 sample cups per day for each of three days. In Prairie 2 three burrow systems were sampled with 12 sample cups per day for three days. In July

2016, more samples were taken from Prairie 1 with a focus on two pocket gopher burrows. The same method was used as the previous year, with samples collected for three days.

Through my study, I identified 19 species of macroinvetebrates associated with pocket gopher burrows. An additional eight macroinvertebrates were identified to genus level and nine more were identified to family level. In total, these macroinvertebrates belonged to 24 families, representing a diverse set of cohabitants within the burrows of pocket gophers. Several of the species identified were new local records for Delaware County: Ceuthophilus divergens, Ceuthophilus latens,

Anisodactylus ovularis, Harpalus erraticus, Glischrochilus quadrisignatus, Pericompsus ephippiatus, Foxella ignota, Dendrophilus sexstriatus, Geomydoecus geomydus. One species, Stictolinus flavipes, was the first documented in the state. Previously S. flavipes had been identified as far west as Illinois and Wisconsin (Klimaszewski et al. 2005).

Twelve species of macroinvertebrates were identified solely within (and not outside) the burrows. While some of these could have entered the opened burrows during the course of this experiment, each of these species may represent specialized burrow inhabitants.

ARTHROPOD FAUNA OF PLAINS POCKET GOPHER (GEOMYS BURSARIUS)

BURROW SYSTEMS

A Thesis

Submitted

in Partial Fulfillment

of the requirements for the Degree

Master of Science

Brett Lyle Gourley

University of Northern Iowa

May 2021

This Study by: Brett Lyle Gourley

Entitled: Arthropod Fauna of Plains Pocket Gopher (Geomys bursarius) Burrow Systems

has been approved as meeting the thesis requirement for the

Degree of Master of Science

______Date Dr. John Ophus, Chair, Thesis Committee

______Date Dr. James W. Demastes, Thesis Committee Member

______Date Dr. Theresa Spradling, Thesis Committee Member

______Date Dr. Jennifer Waldron, Dean, Graduate College

iii

ACKNOWLEDGEMENTS

I would like to thank my advisor Dr. John Ophus and my committee members Dr.

James W. Demastes and Dr. Theresa Spradling, for providing support throughout this entire process. I would like to thank Dr. Kenneth Elgersma for his support during this process as well. Also, I want to thank two students who helped me with the preservation and cleaning of specimen: Greg Gustafson and Husein Kurdic. Next, I want to thank my parents, brothers, Monyae Williamson, and friends for encouraging me over the last few years to finish this. Rick and Kerri Linderwell also deserve a thank you for allowing me to conduct this research on their property. Thank you to the University of Northern Iowa Department of Biology and the Graduate College for their financial support.

TABLE OF CONTENTS

PAGE

LIST OF TABLES ...... v

LIST OF FIGURES ...... vi

CHAPTER 1: INTRODUCTION ...... 1

CHAPTER 2: LITERATURE REVIEW ...... 4

Factors Contributing to Isolation ...... 5

Genetic Effect of Isolation...... 6

Coevolution ...... 7

Arthropods in Pocket Gopher Burrows...... 10

CHAPTER 3: METHODS ...... 13

Study Site ...... 13

Burrow System Preparation ...... 13

Arthropod Collection ...... 16

Arthropod Processing and Identification ...... 18

CHAPTER 4: RESULTS...... 19

CHAPTER 5: DISCUSSION ...... 25

REFERENCES ...... 32

APPENDIX ...... 40

LIST OF TABLES

TABLE PAGE

1. Comparison of parasites identified with pocket gophers...... 9

2. Year 1 pocket gopher mound GPS coordinates ...... 16

3. Year 2 pocket gopher mound GPS coordinates ...... 16

4. List of identified arthropods collected in control cups and burrow cups year 1

and year 2 ...... 20

5. Total number of specimens, classes, genera, and species found in each burrow

system ...... 22

LIST OF FIGURES

FIGURE PAGE

1. Map of pocket gopher locations year 1, prairie 1 ...... 15

2. Map of pocket gopher locations year 1, prairie 2 ...... 15

3. Map of pocket gopher locations year 2 ...... 16

4. C. divergens ...... 26

5. C. latens ...... 26

6. H. erraticus ...... 26

7. A. ovularis ...... 26

8. D. sexstriatus ...... 26

9. F. ignota ...... 26

10. G. geomydis ...... 26

11. S. flavipes ...... 29

CHAPTER 1

INTRODUCTION

Pocket gophers are unique among mammals in how asocial they are. They fight any intruders in their burrow systems, with mating being the exception. Zinnel and

Tester (1994) studied pocket gopher interactions and found gophers rarely came within

10 meters of other gophers unless their burrows ran adjacent to each other. Usually, pocket gophers use different behaviors such as vocalization, digging, and feces as signposts to surrounding gophers. In their study, Zinnel and Tester found gophers use these different behaviors as agonistic instead of affiliative to ensure there is enough food resources. Due to this, pocket gophers create unique environments with their burrow systems. Huntly and Inouye (1988) found that as the pocket gophers burrow and alter the soil and vegetation, they also affect other organisms. They found that, by changing the soil, pocket gophers increase plant diversity, resulting in an increase in animal diversity. Reichman and Seabloom (2002) further found that gopher burrows lead to substantial alterations of the soil, including nitrification, mineralization rates, bulk density, and moisture regimes, ultimately promoting diversity. These alterations create an opportunity for great biotic diversity in pocket gopher burrows.

Hubbell and Goff (1939) were the first to study the arthropod fauna of pocket gopher burrow systems. Through their research, 11 new obligate hypogean species of arthropods were discovered. Commonly, research has revolved around a specific family, e.g., Histeridae, Leiodidae, or Scarabaeidae (Connior et al. 2014; Kovarik et al.

2008; Kriska and Katovich 2005; Powell et al. 2017; Skelley and Woodruff 1991). Other examples include studies from Alabama (Skelley and Gordon 2001), Arkansas (Kovarik et al. 2008), Colorado (Miller and Ward 1960), Florida (Hubbell and Goff 1939; Skelley and

Woodruff 1991), Georgia (Skelley and Gordon 2001), Illinois (Peck and Skelley 2001),

Indiana (Powell et al. 2017), Louisiana (Peck and Skelley 2001), Mississippi (Peck and

Skelley 2001), Minnesota (Bartel and Gardner 2000) New Mexico (Peck and Skelley

2001), Oklahoma (McAllister et al. 2013; Peck and Skelley 2001), Texas (Peck and Skelley

2001), Wisconsin (Kriska and Katovich 2005). There is no evidence that work similar to this has been conducted in Iowa.

Relatively little is still known about pocket gophers and the arthropod fauna that lives in their burrow systems. Pocket gophers move above land as they move between populations, but distance does play an essential role with less gene flow over longer distances (Welborn and Light 2014). With increasing amounts of land being urbanized or converted to farmland, changes to vegetation, soil type, and quantity of ground litter restrict pocket gopher movement (Cortez et al. 2015).

Pocket gophers have low heterozygosity compared to other small mammals, a likely result of population isolation and genetic drift (Penney and Zimmerman 1976). As pocket gopher populations in Iowa become increasingly fragmented due to these human activities and their effects, pocket gopher populations may experience increasing population isolation and genetic drift. With pocket gophers showing low heterozygosity and occurring in isolated populations, arthropods dependent on pocket gophers or their

3 burrow systems may also exhibit this trait. Through this study, I identify the arthropod fauna living in Iowa pocket gopher, Geomys bursarius, burrow systems, compare the fauna between two different prairies (approximately 150 m apart), and compare the fauna identified in one of those prairies over a two-year period.

CHAPTER 2

LITERATURE REVIEW

Organisms that live in caves and other subterranean habitats offer an unique opportunity to study evolution since they often live life in isolation (Culver et al.

2006). Due to their isolation and small population size, genetic isolation and genetic drift may occur. Species are at various stages of adaptation to the subterranean ecosystem, with some species viewed as temporary inhabitants, others as having a robust hypogean, defined as subterranean, affinity, and the last group as obligatory hypogean. Because of this variability, biodiversity in subterranean ecosystems has been called a melting pot (Gibert and Deharveng 2002). Pocket gophers and other subterranean mammals share similarities with cave-dwelling organisms as they live most of their life beneath the surface and are only occasionally seen above ground. Within this cave-like ecosystem, several obligate species live in isolation while other species occur as temporary inhabitants or having a strong hypogean affinity. Several studies have focused on the arthropod fauna in pocket gopher burrows (Hubbell and Goff 1939;

Howell et al. 2016; Skelley and Gordon 2001; Peck and Skelley 2001; Connior et al.

2014).

In Iowa, there is one species of pocket gopher, Geomys bursarius. This species has a patchy distribution in prairies and grasslands from southern Canada throughout the midwest to Texas, west to New Mexico and east to Indiana (Connior 2011). It primarily feeds on the roots and stems of grasses and forbs. Individuals are solitary

5 coming into contact with other individuals only for mating purposes and defending territories.

Factors Contributing to Pocket Gopher Isolation

One of the features of Iowa prairies is that they are becoming increasingly fragmented as land is being converted into farmland and urban areas, ultimately causing more isolation for pocket gophers and other species. Modern farming practices lead to loss of habitat as the land is planted in monocultures and intensely managed. This fragmentation is leading to a wide array of effects on Iowa’s native species. Frog and toad species richness and abundance have been negatively impacted by urban development (Knutson et al. 1999). Butterflies are another example as they may show a strong response due to differences in vegetation, with edge permeability possibly being the most critical factor (Ries and Debinski 2001). Habitat fragmentation also affects mammal movement; as fragmentation increases for meadow voles, cotton rats, and deer mice, interpatch movement decreases (Diffendorfer et al. 1995). In addition to habitat fragmentation, the planting of monocultures has adversely affected pocket gophers as well, in part due to the removal of native plant species and soil erosion and compaction (Cortez et al. 2015). Anthropomorphic fragmentation, combined with pocket gophers’ naturally leading solitary, subterranean existence, results in an increased possibility for speciation.

Pocket gopher movements may be confined because individuals reuse burrow systems instead of moving to new areas (Williams and Baker 1976). This low vagility

6 may lead to more isolation and less gene flow for the pocket gophers, which could eventually lead to a high rate of subspeciation. This evolutionary isolation may also lead to unique species of arthropods living within specific populations of pocket gophers and other subterranean mammals.

Genetic Effect of Isolation

Due to the high possibility of natural biogeographic evolutionary experiments with pocket gopher populations, genetics of pocket gophers presents an intriguing evolutionary case study. Increasing barriers and distance reduces the gene flow between populations (Welborn and Light 2014). Gophers may travel above ground to the populations further away, albeit rarely, while closer populations interbreed and interact more frequently. With increasing habitat fragmentation occurring, pocket gopher interaction will decrease, genetically isolating populations and possibly leading to reproductive incompatibility (Hafner et al. 1987).

This may be further compounded by intensive farming practices, decreasing pocket gophers’ density meaning there would be even less gene flow between populations, possibly creating a bottleneck event. A high percentage of shared alleles between different species of pocket gophers (Selander et al. 1974) and low heterozygosity among pocket gopher species may indicate genetic drift or a bottleneck event having occurred (Penney and Zimmerman 1976). Due to this and the island type distribution of pocket gophers with small effective population size, evolutionary

7 isolation may occur, resulting in speciation of pocket gophers and the arthropods living in their burrows.

Coevolution

In order for coevolution to occur there needs to be an intimate relationship between two species that is demonstrably old (Hafner et al. 2000). Due to the life habits of subterranean mammals and the organisms coexisting with them, these organisms may be evolutionarily tied together. With both the subterranean mammals and the arthropods having small populations and patchy distributions, the chances of cospeciation are increased. This may be especially true with parasites of subterranean mammals as the parasites are dependent on the host (Hafner et al. 2000).

Interest in species interactions has led to studies on other subterranean mammals and their parasites, with new species of parasites being identified (Koudela et al. 2000; Gardner 1985). There are other factors which may determine the parasites found in subterranean mammal burrow systems. Helminth infections were limited by the mammal’s sex of the organism, feeding habits, individual age, level of activity or season (Lutermann and Bennett 2012; Rossin et al. 2010). In pocket gophers, the seasons affect helminth populations, as warmer soil temperatures increase the infectivity of parasite eggs and larvae (Gardner 1985). Increased interaction between pocket gopher individuals during their reproductive period will also increase individual chances of parasitism (Krasnov et al. 2005). The low vagility of pocket gophers prevents individuals from traversing long distances, leading to populations having the same

8 species of parasites, but restricting parasite gene flow over longer distances to other populations.

Parasite infestation of other mammals is influenced by the amount of space given. For example, large bat roosts allowed for more space to spread out and decreased parasite numbers (Dick et al. 2003). Parasite numbers in bats also increase with the continuous use of a roost (Zahn and Rupp 2004). In subterranean mammals, geographic isolation can lead to lower parasite infestation (Schaarf et al. 1997).

Genetic variation between pocket gophers and their chewing lice is about the same (Nadler et al. 1990) indicating genetic isolation for pocket gophers and their parasites. This also leads to the phylogenies of pocket gophers and their lice to be more similar than expected due to chance (Page 1996), meaning the louse phylogenetic tree is not independent of the pocket gopher phylogenetic tree (Hafner et al. 2000). As gene flow between the gophers is reduced due to bottlenecking, the founder effect, or their patchy distribution, the transfer of chewing lice and other parasites is reduced leading to conditions for speciation to occur.

Louse transmission may occur predominately through mother to offspring contact (Demastes et al. 2002), which is a possible contributor to a pattern of cospeciation (Demastes and Hafner 2003; Nadler et al. 1990). In addition to chewing lice, several other species of parasites have been identified living with pocket gophers

(Table 1).

Table 1: Comparison of parasites identified with pocket gophers Parasite Bartel Howell McAllister Miller Timm Tuszynski Gourley and et al. et al. and and and (2020) Gardner (2016) (2013) Ward Price Whitaker (2000) (1960) (1980) (1972) Opisocrostis bruneri X ------Foxella ignota X X X X - - X Dactylopsylla - - - X - - - percernis Thrassis petiolatus - - - X - - - Dactylopsylla ignota - - - - - X - Geomydoecus X - - X X X X geomydis Geomydoecus - - - - X - - illinoensis Geomydoecus - - - - X - - oklahomensis Geomydoecus - X - - X - - nebrathkensis Geomydoecus ewingi - - - - X - - Geomydoecus - - - - X - - heaneyi Geomydoecus spickai - - - - X - - Geomydoecus - - - - X - - subgeomydis Geomydoecus - - - X - - - thomyus Geomydoecus - - - X - - - chapini Geomydoecus - - - X - - - californicus Geomydoecus minor - - - X - - - Geomydoecus - - - X - - - geomydis- californicus Dermacentor X ------variabilis Ixodes sculptus - - - X - - - Ixodes kingi - - - X - - - Echinonyssus - X - - - - - geomydis Androlaelaps geomys - X - - - - - (table continues)

Parasite Bartel Howell McAllister Miller Timm Tuszynski Gourley and et al. et al. and and and (2020) Gardner (2016) (2013) Ward Price Whitaker (2000) (1960) (1980) (1972) Echinonyssus - X - - - - - geomydis Haemogamasus - - - X - - - ambulans Ischryopoda armatus - - - X - - - Hirstionyssus - - - X - X - geomydis Garmania ponorum - - - X - - - Haemolaelaps - - - X - - - geomys Aulaelaps stabularis - - - X - - - Androlaelaps geomys - - - - - X - Hirstionyssus - - - - - X - longichelae

Arthropods in Pocket Gopher Burrows

Aside from obligate parasite studies, there have been few studies done to observe the arthropod fauna of pocket gopher burrow systems. The most notable studies have been conducted by Hubbell and Goff (1939) in Florida and Kovarik et al.

(2008) in Nebraska. Hubbell and Goff (1939) did an in-depth observation of arthropod fauna in pocket gopher burrow systems, and their research methodology has been repeated multiple times, including the use of trapping both arthropods and gophers at the same time and the use of molasses as bait. Hubbell and Goff found nearly 80 species, with 15 of them believed to be obligate inquilines only living in the burrow systems. Around 55 species were classified as casual and considered able to live outside of the burrow system, even though the species spent most of its time in the

11 burrow. Eleven species were new to science. Kovarik et al. (2008) captured 12 species with 2 new to science species being identified.

In the Southeastern United States, Paul E. Skelley has conducted several studies specifically collecting different families of Coleoptera: Leiodidae and Scarabaeidae. From

Skelley and Gordon (2001), three new to science species of Aphodius were found in the burrows of pocket gophers in the southeastern United States. In working with carrion beetles, Peck and Skelley (2001) identified two new species in the southern United

States and expanded the range of other species. Skelley and Woodruff (1991) found five new species of Aphodius from Florida pocket gopher burrows. Additionally, Connior et al. (2014) found eight new state records of Coleoptera in pocket gopher burrows from

Arkansas.

In states neighboring Iowa, studies have focused on specific families associated with pocket gophers. Kriska and Katovich (2005) found six scarab species, with four being widespread throughout both pocket gopher subspecies. They also found fall to be the most productive time for collecting scarab beetles; May through July was less productive for collection because higher pocket gopher activity resulted in ineffective pitfall trapping as pocket gophers buried the scarab beetle traps.

A similar study was done in Indiana by Powell et al. (2017), examined Coleoptera associated with Geomys bursarius. In total, they found 25 species belonging to 8 families in the 885 individuals they examined. Among the nine most trapped species,

12 they looked at activity times with five species having one peak activity time throughout the year and the four other species exhibiting a bimodal distribution of activity peaks.

In Minnesota, Bartel and Gardner (2000) focused on the arthropods and helminth parasites found on Geomys bursarius. They examined 144 individuals from seven different localities. Twenty-five species were found with six of the species being new locality records. In addition, they found seven new host records out of the 25 species. Howell et al. (2016) also focused on the parasite species finding six living on pocket gophers in Nebraska.

CHAPTER 3

METHODS

Study Site

I conducted my research on a private farm adjacent to Backbone State Park in northwest Delaware County, Iowa, USA, 42.38070 N, 91.35105 W. There were two prairies I gathered my specimens from, and each of these prairies has a history of being crop farmed. The prairies have been placed in the Conservation Reserve Program (CRP) for over 25 years. Prairie 2 is approximately 1,100 feet east of Prairie 1. Using the web soil survey government website (Web Soil Survey 2019), Prairie 1 was determined to have Seaton silt loam for soil while prairie 2 has Lamont fine sandy loam.

Burrow System Preparation

In 2015 (year 1), samples were taken from both prairies by placing a control cup above ground and three cups inside the burrow system as a pitfall trap with molasses as bait. Prairie 1 had five burrow systems with a total of 20 sample cups per day taken over three days, with Prairie 2 having 3 burrow systems with 12 sample cups per day taken over the same three-day period. In 2016 (year 2), two burrow systems were sampled with Prairie 1 has specimen collected from two different pocket gopher burrow systems were sampled with 8 sample cups per day. The same cup placement as in 2015, with samples being collected for three days.

To determine the burrow systems to use, I looked for fresh piles of dirt from the pocket gophers without grass growing in it (Hubbell and Goff 1939). I then swiped the

14 mound of dirt away, so it was flat with the ground. Next, I felt the ground for a soft spot indicating a pocket gopher tunnel. I then dug to open the burrow system to a y- intersection. I placed cut mesh wire and bent it slightly to stick up into the tunnels for the burrows 1, 2, and 3, while gophers in 4 and 5 were trapped. The pocket gophers 1,

2, and 3 remained as they eluded the traps by pushing soil into the trap triggering it. In

2016, pocket gopher 1 was trapped while pocket gopher 2 remained. There was evidence of other pocket gopher activity in the burrow system of pocket gopher 1 as the burrow was backfilled. This may have been due to recolonization by another gopher, or there could have been another gopher living in the burrow system when I trapped the first one. Mesh wire was placed to help deter the gopher from destroying the pitfall traps as trapping several gophers was time-consuming, and gophers would replace the ones trapped. It is important to note there is no guarantee the cups were in the same burrow system as many are close to each other, and burrows may run several meters in length and width. Trapped gophers were placed in the UNI Mammal collection for teaching and research purposes.

Pocket Gopher 1 Pocket Gopher 2 Pocket Gopher 3 Pocket Gopher 4 Pocket Gopher 5

Figure 1: Map of pocket gopher locations year 1, prairie 1

Pocket Gopher 1 Pocket Gopher 2 Pocket Gopher 3 Figure 2: Map of pocket gopher locations year 1, prairie 2

Table 2: Year 1 pocket gopher mound GPS coordinates Prairie Pocket Gopher GPS Coordinates 1 1 42.63384 N, 91.58080 W 1 2 42.63385 N, 91.58076 W 1 3 42.63369 N, 91.58100 W 1 4 42.63358 N, 91.58114 W 1 5 42.63435 N, 91.57718 W 2 1 42.63458 N, 91.57733 W 2 2 42.63451 N, 91.57713 W 2 3 42.63435 N, 91.57718 W

Pocket Gopher 1 Pocket Gopher 2

Figure 3: Map of pocket gopher locations year 2

Table 3: Year 2 pocket gopher mound GPS coordinates Prairie Pocket Gopher GPS Coordinates 1 1 42.63363 N, 91.58006 W 1 2 42.63388 N, 91.58081 W

Arthropod Collection

My first arthropod collection occurred over the 17-19 and 24-26 of July 2015, with pocket gophers being trapped on July 18. The trapped gophers were taken back to

Dr. Demastes/Spradling lab where they were brushed, and samples were given back for identification. Collection occurred between 4 pm and 8 pm in both prairies. Pitfall traps were created by using 5-ounce plastic cups with lids and pouring approximately 10 ml of molasses to help lure and trap the arthropods. This method follows the procedure detailed by Hubbell and Goff (1939) as they found it to be more successful trapping arthropods with molasses baited traps than unbaited. Control cups were placed in the center of the three tunnel cups with GPS coordinates based off the control cups. All cups were dug and placed slightly below the ground with the sand placed back around them. Cut pieces of plywood were placed over the top of the burrow with dirt and grass chunks placed on top of these to help secure the boards in place and to try to discourage other animals from disturbing the study and protect the traps from other outside disturbances. The following day, I collected the cups and replaced them with a new cup. Once taken out, the cups were marked using the following notation, 1= prairie, 1=pocket gopher, 1=mound/cup, 1=day. Then the new cups were placed in the same spot and covered up using plywood, which was then covered with grass and dirt.

The specimen cup was placed in a cooler and taken back to the entomology lab. In year

2, pocket gophers were trapped on July 22 and 23. Arthropods were collected on July

23, 24, 25, between 4 pm and 8 pm with the same process of digging into the burrow, covering it, and taking the cups back to the lab.

Arthropod Processing and Identification

Once back to the lab, the arthropod samples were separated from the molasses and dirt using filter paper and water. After the samples were thoroughly cleaned and separated from the molasses and soil, they were stored in 5-ounce cups with 70% ethanol. It was a goal to identify all samples to the species level. However, due to damage from the collection procedure, some specimens could not be identified past the family- or genus-level (Table 4). Fifty-two percent of the specimens were identified to the species level with the use of available references and a high-power dissecting microscope.

CHAPTER 4

RESULTS

In total, five classes were identified: Arachnida, Insecta, Entognatha, Isopoda,

Diplopoda. From these classes, 13 orders were identified. Three of these orders were omitted from my analysis because they were not found in any experimental cups and, thus, were not burrow inhabitants. My analysis also excluded the two classes, Isopoda and Diplopoda, as specimen from these classes were only found in the control cups.

Out of the remaining ten orders, 24 families were identified. Class Arachnida consisted of two orders Araneae and Ixodida, with four families. Class Insecta had seven orders:

Coleoptera, Diptera, Hemiptera, Hymenoptera, Orthoptera, Phthiraptera, and

Siphonaptera with 18 families. Lastly, class Entognatha consisted of one order

Entomobryomorpha with one family. From this point, specimens were identified to family, genus, or species. I was unable to identify some specimens to genus or species possibly due to damage from the process, gophers damaging cup and cup contents, or other organisms eating some or most of the cup’s contents. The identified species and genera are listed below by order in the following format: class, order, family, scientific name (if found). For a more detailed table of specimens collected with location and year, see Appendix.

Table 4: List of identified arthropods collected in control cups and burrow cups year 1 and year 2 Class Order Family Genus Species

Arachnida Araneae Lycosidae Tigrosa helluo

Arachnida Araneae Trachelidae Trachelas tranquillus

Arachnida Araneae Salticidae Phidippus putnami

Arachnida Ixodida Ixodidae - -

Insecta Coleoptera Carabidae Blemus discus

Insecta Coleoptera Carabidae Pterostichus melanarius

Insecta Coleoptera Carabidae Pericompsus ephippiatus

Insecta Coleoptera Carabidae Anisodactylus ovularius

Insecta Coleoptera Carabidae Cicindela repanda

Insecta Coleoptera Carabidae Micratopus -

Insecta Coleoptera Carabidae Harpalus erraticus

Insecta Coleoptera Carabidae - -

Insecta Coleoptera Carabidae Bembidion -

Insecta Coleoptera Curculionidae Otiorhynchus sulcatus

Insecta Coleoptera Histeridae Dendrophilia sexstriatus

Insecta Coleoptera Nitidulidae Glischrochilus quadrisignatus

Insecta Coleoptera Staphylinidae Stictolinus flavipes

Insecta Coleoptera Staphylinidae Syntomium - Megalopsidiinae Insecta Coleoptera Staphylinidae Lathropinus - Pinophilinae Insecta Coleoptera Staphylinidae Stictocranius - Euaesthetinae (table continues)

Class Order Family Genus Species

Insecta Diptera Anthomyiidae - -

Insecta Diptera Bombyliidae - -

Insecta Diptera Hippobiscidae - -

Insecta Diptera Muscidae Musca domestica

Insecta Diptera Mycetophilidae - -

Insecta Diptera Phoridae Puliciphora -

Insecta Diptera Tachinidae - -

Insecta Hemiptera - - -

Insecta Hymenoptera Formicidae Formica -

Insecta Hymenoptera Sphecidae - -

Insecta Orthoptera Rhaphidophorid Ceuthophilus divergens ae Insecta Orthoptera Rhaphidophorid Ceuthophilus latens ae Insecta Orthoptera Gryllidae Gryllus veletis

Insecta Phthiraptera Trichodectidae Geomydoecus geomydus

Insecta Siphonaptera Ceratophyllidae Foxella ignota

Entognatha Entomobryomor Isotomidae Isotoma - pha

Note. Specimen were identified to the lowest classification with dashes indicating I was unable to identify further for various reasons, including damage to the specimen.

This study design did not yield enough data for statistical analyses of trends.

However, the species diversity reported here provides insight into the species

22 composition of the cohabitants of G. bursarius burrows in Iowa, and the quantitative data produced here provide an initial assessment of total abundance (Table 5) and species-specific abundance that can inform the design of future studies.

Table 5: Total number of specimen-, class-, genera- and species- found in each burrow system Year 1 Prairie 1 Total N Class Genera Species 1 1398 3 10 7 2 431 2 14 11 3 849 3 12 7 4 449 2 12 8 5 309 3 12 9 Prairie 2 1 373 2 11 8 2 199 2 10 7 3 205 3 9 7 Year 2 Prairie 1 1 550 2 8 5 2 609 2 12 6

Twelve species were identified exclusively within burrows. Three of these were in the order Araneae: Tigrosa helluo, Trachelas tranquillas, and Phidippus putnami. Four more species were from the family Carabidae: Blemus discus, Pericompsus ephippiatus,

Anisodactylus ovularius, and Harpalus erraticus. There was one species of histerid beetle

(family Histeridae), Dendrophilia sexstriatus, and there was one species from the family

Nitulidae, Glischrochilus quadrisignatus. There was one species each in the family

Staphylinidae and Ceratophyllidae, Stictolinus flavipes and Foxella ignota. The obligate

23 ectoparasite Geomydoecus geomydis (family Trichodectidae) was found exclusively within burrows.

Eight species were identified in both prairies, including four species in the order

Coleoptera: P. melanarius, D. sexstriatus, O. sulcatus, and S. flavipes. One species in the family Muscidae, M. domestica, was recovered in both prairies. Two species in the order Orthoptera, C. latens and G. veletis, and one species of the family Trichodectidae,

G. geomydis, were found at both prairies. Nine more species were identified exclusively in Prairie 1. These included two species of the Order Aranae, T. helluo and T. tranquillas, and five species of the Order Coleoptera, B. discus, P. ephippiatus, A. ovularius, C. repanda, and G. quadrisignatus. In addition, one species of the family

Rhaphidophoridae, C. divergens, and one species of the Family Ceratophyllidae, F. ignota, were found only in Prairie 1. Two species, P. putnami and H. erraticus, were identified exclusively in Prairie 2.

Year 1 data included 11 species specific to that year, including three species of the order Araneae species: T. helluo, T. tranquillas, and P. putnami. Four species in the family Carabidae, B. discus, A. ovularius, C. repanda, H. erraticus, were only found in

Year 1. One species in the family Curulionidae, O. sulcatus, and one species in the family

Nitidulidae, G. quadrisignatus, were only found in Year 1, as was a species in the family

Staphylinidae, S. flavipes. There was one species of the family Rhaphidophoridae species: C. divergens, and one Ceratophyllidae species: F. ignota. Year 2 had one species specific to that year: P. ephippiatus. Six species were commonly identified between

24 both years, including two species in the family Coleoptera, P. melanarius and D. sexstriatus, one species in the family Muscidae, M. domestica, and two species in the order Orthoptera, C. Latens and G. veletis. The obligate ectoparasite, G. geomydis, also was recovered in both year 1 and year 2.

CHAPTER 5

DISCUSSION

Pocket gopher burrow systems provide habitat for several different species of arthropods. There were 24 families found in the burrow systems. In the 24 families, 19 species were identified with eight more specimens identified to the genus level and nine being identified to family. For various reasons, damaged during cleaning and storing, due to the gophers or by other organisms, some of the specimen we were not possible to identify to the genus or species level. It is important to note that, due to the collection process, some of the identified species may have originated outside of the burrow.

Seven species are common to Iowa and are in state records multiple times.

These species include: Tigrosa helluo (wolf spider), Trachelas tranquillus (broad-faced sac spider), Phidippus putnami (jumping spider), Gryllus veletis (spring field cricket),

Musca domestica (housefly), Cicindela repanda (bronzed tiger beetle) and Glischrochilus quadrisignatus (four-spotted sap beetle). In addition, three species found are invasive species found in the area already: Blemus discus (ground beetle), Pterostichus melanarius (common black ground beetle), and Otiorhynchus sulcatus (black vine weevil).

Figure 4: C. divergens Figure 5: C. latens

Figure 6: H. erraticus (head missing) Figure 7: A. ovularis

Figure 8: D. sexstriatus Figure 9: F. ignota

Figure 10: G. geomydis

Through this study, we were able to confirm and extend the ranges of several other species living in Iowa. This study seems to be the first one done on arthropods living in Geomys bursarius burrow systems in Iowa, and as such several of these species have not been associated with pocket gophers in the state but have been in surrounding states. Both camel cricket species, Ceuthophilus divergens and Ceuthophilus latens, have been found in Iowa (Knutson and Jaques 1935), but we were able to confirm and add them to the Delaware county fauna. The preferred habitat of Ceuthophilus divergens is dry, open, or grassy woodlands, while the Ceuthophilus latens has been found in a wide array of habitats including open grasslands (Bland 2003). The ground beetle, Anisodactylus ovularis, represents a new specimen for Delaware County, Iowa, as it has been noted of already living in Iowa and has been collected along fencerows

(Esau 1968). Anisodactylus ovularis is a species which has not been found with pocket gophers in Iowa previously. Another specimen added to Delaware County was Harpalus erraticus (ground beetle), it has been identified in nearby counties Johnson and

Buchanan (El-Moursy 1958). Similarly, Glischrochilus quadrisignatus (four-spotted sap beetle) was added as a specimen from Delaware county, it has previously been identified in Story county (Bugguide). Also, the ground beetle, Pericompsus ephippiatus

(Bugguide), was added to the Delaware County fauna as it has few records in the state.

Foxella ignota (flea) has been found in the state, but this is a new record of the species adding it to Delaware County and confirming its associating with pocket gophers in the state (McAllister et al. 2013; Howell et al. 2016). Dendrophilus sexstriatus (hister beetle)

28 has previously been found in Henry and Monroe counties (Hatch 1938), here D. sexstriatus is added to Delaware County. Geomydoecus geomydus is a species of louse that has been identified around Ames, Iowa (Timm and Price 1980). This species of louse is found parasitizing pocket gophers in much of the western United States (Timm and Price 1980). This study confirms Geomydoecus geomydus also inhabits northeastern Iowa and is new for Delaware County records.

One species found in the burrow systems is a new state record. Stictolinus flavipes a rove beetle was a species not found in the state as it has been a species with a range further north and east as it has been found in Canada and in the United States from Connecticut and New Hampshire to Wisconsin and Illinois (Klimaszewski et al.

2005). Specimen have been collected in forests (Klimaszewski et al. 2005) but not in burrow systems such as this. Typically, S. flavipes is found in various types of litter or decaying organic matter (Brunke et al. 2011). Other species of subfamily Staphylininae, such as Heterothops marmotae and Bisnius lautus, have been found in mammal nests and underground burrows (Webster et al. 2012). Hubbell and Goff (1939) found two staphylinid specimens, which they thought were predatory, in their gopher burrows.

These specimens were near the genus Atheta. Stictolinus flavipes is likely a general predator searching for easy prey in the cups as it was going through the burrows. Since

I did not find any specimen of S. flavipes above ground and all were in the burrow systems, the species is probably a casual hypogean species going between above ground and the tunnels. This is an extension of its current range to include Iowa.

Figure 11: S. flavipes

Out of the 12 species found exclusively in the burrows, two are obligate hypogeans: Foxella ignota and Geomydoecus geomydis. These species depend on the pocket gophers and were brushed from coats of the gophers. The other 10 species may be casual hypogean as several of these species, excluding S. flavipes, have been found in the state already outside of pocket gopher burrow systems. These species include:

Tigrosa helluo, Trachelas tranquillas, Phidippus putnami, Blemus discus, Pericompsus ephippiatus, Anisodactylus ovularius, Harpalus erraticus, Dendrophilia sexstriatus,

Glischrochilus quadrisignatus, and Stictolinus flavipes.

Prairie 2 was a smaller prairie with a smaller population of pocket gophers limiting sampling which may be why there were only a few species found exclusively in that prairie compared to Prairie 1. A similar pattern took place between years as sampling in year one was more extensive than sampling in year two.

In conclusion, ten species were confirmed to live in Iowa with new records for

Delaware county. One species is a new record for Iowa that expands the known range of the species. More research is needed to fully catalog all of the arthropod species living with pocket gophers in Iowa. As I used molasses for bait and did my study during

July, and different arthropods will be attracted to other baits and have different periods of peak activity (Kriska and Katovich 2005). Changing the sampling method or sampling times could result in finding new species and new localities for species to add to our knowledge of the natural world.

Contrary to my expectations at the beginning of this study, there were no new species identified through my research. Other studies such as Hubbell and Goff (1939) did identify new species by exploring the unique habitat of pocket gopher burrows. This could be due to multiple reasons such as time of year trapped considering arthropods activity varies throughout the year. Location may have also been a factor as the gopher population where my research was conducted was in CRP land, which had been previously farmed. Five specimens were damaged through the capturing and cleaning process, which also could have been a factor in no new species being identified. To determine if these were factors in identifying new species, I believe other locations throughout Iowa would need to be examined with a range of land types from natural prairie to previously farmed land.

Several species ranges were confirmed or expanded, which aligns with my expectations and studies done by Kriska and Katovich (2005), Connior et al. (2014), and others. Several of the species were confirmed in Delaware County as part of their range. The known distribution of one species, Stictolinus flavipes, was expanded to include Iowa as part of its range.

Further research is needed to confirm and determine other species living in pocket gopher burrow systems. Using different bait in the pitfall traps may also help to attract different species altering results and possibly discovering new species due to

Geomys bursarius isolation.

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APPENDIX

Year 1 Year 2 Prairie 1 Prairie 1 Prairie 2 Prairie 2 Prairie 1 Prairie 1 Controls Tunnels Controls Tunnels Controls Tunnels Burr Sam Burr Sam Burr Sam Burr Sam Burr Sam Burr Sam ow ple ow ple ow ple ow ple ow ple ow ple Size Size Size Size Size Size (n) (n) (n) (n) (n) (n) Tigrosa B 1 0 B 1 0 B 1 0 B1 0 B 1 0 B 1 0 helluo B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 1 B 3 0 B 3 0 - - - - B 4 0 B 4 0 ------B 5 0 B 5 0 ------Trachelas B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 tranquillas B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 0 ------B 5 0 B 5 1 ------Phidippus B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 putnami B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 1 - - - - B 4 0 B 4 0 ------B 5 0 B 5 0 ------Ixodidae B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 0 - - # 1 B 4 0 B 4 0 ------B 5 0 B 5 0 ------Blemus B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 discus B 2 0 B 2 1 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 0 ------B 5 0 B 5 0 ------Pterostich B 1 1 B 1 0 B 1 1 B 1 1 B 1 1 B 1 0 us B 2 1 B 2 0 B 2 0 B 2 1 B 2 1 B 2 1 melanariu B 3 0 B 3 0 B 3 2 B 3 0 - - - - s B 4 1 B 4 0 ------B 5 4 B 5 0 ------Pericomps B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 us B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 1 B 3 0 B 3 0 B 3 0 B 3 0 - - - -

Year 1 Year 2 Prairie 1 Prairie 1 Prairie 2 Prairie 2 Prairie 1 Prairie 1 Controls Tunnels Controls Tunnels Controls Tunnels Burr Sam Burr Sam Burr Sam Burr Sam Burr Sam Burr Sam ow ple ow ple ow ple ow ple ow ple ow ple Size Size Size Size Size Size (n) (n) (n) (n) (n) (n) ephippiat B 4 0 B 4 0 ------us B 5 0 B 5 0 ------Bembidio B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 n sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 2 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 0 ------B 5 0 B 5 0 ------Anisodact B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 ylus B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 ovularius B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 1 ------B 5 0 B 5 1 ------Cicindela B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 repanda B 2 0 B 2 1 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 0 ------B 5 1 B 5 0 ------Micratopu B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 s sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 1 ------B 5 0 B 5 0 ------Harpalus B 1 0 B 1 0 B 1 0 B 1 1 B 1 0 B 1 0 erraticus B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 0 ------B 5 0 B 5 0 ------Carabidae B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 1 ------B 5 0 B 5 1 ------Otiorhync B 1 0 B 1 1 B 1 0 B 1 0 B 1 0 B 1 0 hus B 2 0 B 2 2 B 2 1 B 2 0 B 2 0 B 2 0 sulcatus B 3 0 B 3 0 B 3 0 B 3 0 - - - -

Year 1 Year 2 Prairie 1 Prairie 1 Prairie 2 Prairie 2 Prairie 1 Prairie 1 Controls Tunnels Controls Tunnels Controls Tunnels Burr Sam Burr Sam Burr Sam Burr Sam Burr Sam Burr Sam ow ple ow ple ow ple ow ple ow ple ow ple Size Size Size Size Size Size (n) (n) (n) (n) (n) (n) B 4 4 B 4 0 ------B 5 0 B 5 0 ------Dendrophi B 1 0 B 1 7 B 1 0 B 1 4 B 1 0 B 1 2 lia B 2 0 B 2 15 B 2 0 B 2 3 B 2 0 B 2 1 sexstriatus B 3 0 B 3 1 B 3 0 B 3 1 - - - - B 4 0 B 4 3 ------B 5 0 B 5 15 ------Glischroch B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 ilus B 2 0 B 2 1 B 2 0 B 2 0 B 2 0 B 2 0 quadrisign B 3 0 B 3 0 B 3 0 B 3 0 - - - - atus B 4 0 B 4 0 ------B 5 0 B 5 0 ------Stictolinus B 1 0 B 1 1 B 1 0 B 1 1 B 1 0 B 1 0 flavipes B 2 0 B 2 3 B 2 0 B 2 1 B 2 0 B 2 0 B 3 0 B 3 3 B 3 0 B 3 1 - - - - B 4 0 B 4 1 ------B 5 0 B 5 1 ------Syntomiu B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 m sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 1 B 3 0 B 3 0 - - - - B 4 0 B 4 0 ------B 5 0 B 5 0 ------Stictocrani B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 us sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 1 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 0 ------B 5 0 B 5 0 ------Lathropin B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 us sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 1 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 0 ------B 5 0 B 5 0 ------Anthomyii B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 dae sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 1 B 3 0 B 3 0 - - - -

Year 1 Year 2 Prairie 1 Prairie 1 Prairie 2 Prairie 2 Prairie 1 Prairie 1 Controls Tunnels Controls Tunnels Controls Tunnels Burr Sam Burr Sam Burr Sam Burr Sam Burr Sam Burr Sam ow ple ow ple ow ple ow ple ow ple ow ple Size Size Size Size Size Size (n) (n) (n) (n) (n) (n) B 4 0 B 4 0 ------B 5 0 B 5 0 ------Bombyliid B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 ae sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 1 ------B 5 0 B 5 0 ------Hippobisci B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 dae sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 1 ------B 5 0 B 5 0 ------Musca B 1 0 B 1 2 B 1 0 B 1 0 B 1 2 B 1 0 domestica B 2 0 B 2 0 B 2 1 B 2 0 B 2 0 B 2 1 B 3 2 B 3 0 B 3 0 B 3 1 - - - - B 4 1 B 4 0 ------B 5 0 B 5 0 ------Mycetoph B 1 0 B 1 0 B 1 0 B 1 1 B 1 0 B 1 0 ilidae sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 0 ------B 5 0 B 5 0 ------Puliciphor B 1 0 B 1 2 B 1 3 B 1 42 B 1 3 B 1 6 a sp. B 2 15 B 2 5 B 2 2 B 2 2 B 2 10 B 2 1 B 3 0 B 3 13 B 3 0 B 3 0 - - - - B 4 4 B 4 14 ------B 5 2 B 5 3 ------Tachinida B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 e sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 1 - - - - B 4 0 B 4 0 ------B 5 0 B 5 0 ------Hemiptera B 1 0 B 1 8 B 1 3 B 1 4 B 1 1 B 1 1 nymph B 2 1 B 2 9 B 2 1 B 2 1 B 2 1 B 2 5 B 3 4 B 3 11 B 3 0 B 3 0 - - - -

Year 1 Year 2 Prairie 1 Prairie 1 Prairie 2 Prairie 2 Prairie 1 Prairie 1 Controls Tunnels Controls Tunnels Controls Tunnels Burr Sam Burr Sam Burr Sam Burr Sam Burr Sam Burr Sam ow ple ow ple ow ple ow ple ow ple ow ple Size Size Size Size Size Size (n) (n) (n) (n) (n) (n) B 4 13 B 4 3 ------B 5 3 B 5 1 ------Formica B 1 442 B 1 868 B 1 55 B 1 212 B 1 263 B 1 219 sp. B 2 79 B 2 213 B 2 63 B 2 114 B 2 6 B 2 547 B 3 46 B 3 678 B 3 101 B 3 83 - - - - B 4 170 B 4 163 ------B 5 45 B 5 181 ------Sphecidae B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 sp. B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 0 ------B 5 0 B 5 1 ------Ceuthophi B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 lus B 2 0 B 2 1 B 2 0 B 2 0 B 2 0 B 2 0 divergens B 3 1 B 3 8 B 3 0 B 3 0 - - - - B 4 15 B 4 4 ------B 5 0 B 5 0 ------Ceuthophi B 1 0 B 1 12 B 1 0 B 1 4 B 1 0 B 1 38 lus latens B 2 1 B 2 38 B 2 0 B 2 2 B 2 0 B 2 3 B 3 0 B 3 40 B 3 1 B 3 1 - - - - B 4 0 B 4 24 ------B 5 2 B 5 2 ------Gryllus B 1 0 B 1 0 B 1 0 B 1 1 B 1 0 B 1 0 veletis B 2 0 B 2 2 B 2 0 B 2 0 B 2 0 B 2 1 B 3 0 B 3 0 B 3 0 B 3 0 - - - - B 4 0 B 4 0 ------B 5 0 B 5 0 ------Foxella B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 B 1 0 ignota B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 2 0 B 3 0 B 3 0 B 3 0 B 3 0 - - # 1 B 4 0 B 4 0 ------B 5 0 B 5 1 ------Isotoma B 1 0 B 1 45 B 1 20 B 1 18 B 1 0 B 1 8 sp. B 2 2 B 2 32 B 2 1 B 2 3 B 2 1 B 2 21 B 3 0 B 3 33 B 3 0 B 3 9 - - - -