Citation
Response of Subterranean Termites to Neighboring Populations of Inter and Intraspecific Colonies after Baiting

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Title:
Response of Subterranean Termites to Neighboring Populations of Inter and Intraspecific Colonies after Baiting
Creator:
Bernard, Sarah J
Place of Publication:
[Gainesville, Fla.]
Florida
Publisher:
University of Florida
Publication Date:
Language:
english
Physical Description:
1 online resource (63 p.)

Thesis/Dissertation Information

Degree:
Master's ( M.S.)
Degree Grantor:
University of Florida
Degree Disciplines:
Entomology and Nematology
Committee Chair:
SU,NAN-YAO
Committee Co-Chair:
SCHEFFRAHN,RUDOLF H
Committee Members:
KERN,WILLIAM H,JR
Graduation Date:
5/2/2015

Subjects

Subjects / Keywords:
Agonism ( jstor )
Bioassay ( jstor )
Colonies ( jstor )
Death ( jstor )
Foraging ( jstor )
Sociobiology ( jstor )
Soldiers ( jstor )
Subterranean termites ( jstor )
Termites ( jstor )
Tunnels ( jstor )
Entomology and Nematology -- Dissertations, Academic -- UF
baiting -- intercolonial -- intracolonial -- isoptera -- noviflumuron -- reinvasion -- subterranean -- termite
Genre:
bibliography ( marcgt )
theses ( marcgt )
government publication (state, provincial, terriorial, dependent) ( marcgt )
born-digital ( sobekcm )
Electronic Thesis or Dissertation
Entomology and Nematology thesis, M.S.

Notes

Abstract:
Coptotermes formosanus Shiraki and Reticulitermes flavipes Kollar are sympatrically occurring and economically important subterranean termites in the Southeastern United States. Blocking of tunnels, unblocking of tunnels, and new tunnel construction was observed in intra and inter species experiments. Intraspecies experiments in two-dimensional arenas in which one population of C. formosanus was baited resulted in elimination of baited termites and subsequent reinvasion of territory. Territories held by unbaited termites increased significantly, nearly doubling after reinvasion. Intraspecies controls did not have a significant change in territory areas. In interspecies experiments, R. flavipes, retreated and blocked tunnels at choke points as C. formosanus advanced. R. flavipes reinvaded baited C. formosanus territory in one repetition. Reinvading termites of both species were eliminated by bait. ( en )
General Note:
In the series University of Florida Digital Collections.
General Note:
Includes vita.
Bibliography:
Includes bibliographical references.
Source of Description:
Description based on online resource; title from PDF title page.
Source of Description:
This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Thesis:
Thesis (M.S.)--University of Florida, 2015.
Local:
Adviser: SU,NAN-YAO.
Local:
Co-adviser: SCHEFFRAHN,RUDOLF H.
Statement of Responsibility:
by Sarah J Bernard.

Record Information

Source Institution:
UFRGP
Rights Management:
Copyright Bernard, Sarah J. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
Classification:
LD1780 2015 ( lcc )

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RESPONSE OF SUBTERRANEAN TERMITES TO NEIGHBORING POPULATIONS OF INTER AND INTRASPECIFIC COLONIES AFTER BAITING By SARAH J. BERNARD A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2015

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© 2015 Sarah J. Bernard

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To my mother, father, grandmother, and brother I could not have reached where I am today without your l ove, support, and understanding

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4 ACKNOWLEDGMENTS I would first like to thank and acknowledge Dr. Nan Yao Su for accepting me as his graduate student. Without his support, my work would not have been possible. His guidance in scientific literature has been an invaluable asset at the start of my entomology career. I would like to acknowledge my committee, Dr. Kern and Dr. Sheffrahn, for sharing their vast arrays of i nsect and termite knowledge with me. I would like to make special acknowledgment of Ron Pepin, without his innovations and designs in termite b ioassays and husbandry, much of the work in our I would like to thank my lab mates for offering their help and advice during this process, and would like to make spec ial note of Dr. Thomas Chouvenc and Caroline Efstathion for their help with reviewing my manuscripts, and Aaron Mullins for his help with field collection . I would also like to acknowledge Secret Woods and Anne Kolb nature centers for their cooperation in allowing us to collect termites for experiments on their grounds. Dr. Weste Osbrink also deserves my thanks and gratitude for usage of termite colonies originating from his laboratory in New Orleans, LA. And last but not least, to my family for being as su pportive as ever of all my decisions. To my father for always encouraging me, especially when it came to entomology, throughout my life. To my mother and brother, for believing I could and would do anything I set my mind to. Finally, t o both of my parents for supporting me financially as I find my way through numerous years of schooling.

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5 TABLE OF CONTENTS page ACKNOWLEDGMENTS ................................ ................................ ................................ .. 4 LIST OF TABLES ................................ ................................ ................................ ............ 7 LIST OF FIGURES ................................ ................................ ................................ .......... 8 LIST OF OBJECTS ................................ ................................ ................................ ....... 10 LIST OF DEFINITIONS ................................ ................................ ................................ . 11 ABSTRACT ................................ ................................ ................................ ................... 12 CHAPTER 1 THESIS INTRODUCTION ................................ ................................ ...................... 13 Economic Impact ................................ ................................ ................................ .... 13 Termite Management ................................ ................................ .............................. 13 History ................................ ................................ ................................ .............. 13 Baiting ................................ ................................ ................................ .............. 14 Territory ................................ ................................ ................................ .................. 15 Reinvasion ................................ ................................ ................................ ........ 15 Agonism ................................ ................................ ................................ ........... 15 Necrophobic Behavior ................................ ................................ ...................... 17 Objectives ................................ ................................ ................................ ............... 18 2 INTRASPECIES INTERACTIONS ................................ ................................ .......... 20 Introduction ................................ ................................ ................................ ............. 20 Materials and Methods ................................ ................................ ............................ 21 Termites ................................ ................................ ................................ ........... 21 Planar Foraging Arena Bioassay ................................ ................................ ...... 21 Procedure ................................ ................................ ................................ ......... 23 Data Collection ................................ ................................ ................................ . 24 Data Analysis ................................ ................................ ................................ ... 25 Results ................................ ................................ ................................ .................... 25 Decline of Termite Health ................................ ................................ ................. 25 Intraspecies Agonism ................................ ................................ ....................... 30 Blockages, Unblockages, and New Tunnel s ................................ .................... 33 Territory Shift ................................ ................................ ................................ .... 35 Death of Reinvaders ................................ ................................ ......................... 36 Addition al Observations ................................ ................................ .................... 38 Discussion ................................ ................................ ................................ .............. 38

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6 3 INTERSPECIES INTERACTIONS ................................ ................................ .......... 41 Introduction ................................ ................................ ................................ ............. 41 Materials and Methods ................................ ................................ ............................ 42 Termites ................................ ................................ ................................ ........... 42 Planar Foraging Arena Bioassay ................................ ................................ ...... 42 Baiting and Reinvasion Experiments ................................ ................................ 42 Data Collection ................................ ................................ ................................ . 42 Data Analysis ................................ ................................ ................................ ... 43 Results ................................ ................................ ................................ .................... 43 Decline of Termite Health ................................ ................................ ................. 43 In terspecies Agonism ................................ ................................ ....................... 47 Blockages, Unblockages, and New Tunnels ................................ .................... 49 Territory Shift ................................ ................................ ................................ .... 52 Discussion ................................ ................................ ................................ .............. 55 4 CONCLUSION ................................ ................................ ................................ ........ 57 LIST OF REFERENCES ................................ ................................ ............................... 58 BIOGRAPHICAL SKETCH ................................ ................................ ............................ 63

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7 LIST OF TABLES Table page 2 1 Phases of Decline in Intraspecies Baited Treatment Panels .............................. 27 2 2 Percent Cumulative Dead Observed in Intraspecies Control Panels .................. 27 2 3 Percent Cumulative Dead Observed in Intraspecies Treatment Panels ............. 27 2 4 Percent Cumulative Dead Observed in Intraspecies Paired Panels ................... 27 2 5 Average Intraspecies Blocks, Unblocks, New Tunnels per Control Panel .......... 34 2 6 Average Intraspecies Blocks, Unblocks, New Tunnels per Treatment Panel ..... 34 2 7 Average Intraspecies Blocks, Unblocks, New Tunnels for Paire d Panels .......... 34 2 8 Average Intraspecies Territory Areas Before and After Panel Connection ......... 36 3 1 Phases of Decline in Interspecies Baited Treatment Panels .............................. 45 3 2 Percent Cumulative Dead Observed in Interspecies Control Panels .................. 45 3 3 Percent Cumulative D ead Observed in Interspecies Treatment Panels ............. 45 3 4 Percent Cumulative Dead Observed in Interspecies Paired Panels ................... 45 3 5 Average Interspecies Blocks, Unblocks, New Tunnels per Control Panel .......... 50 3 6 Average Interspecies Blocks, Unblocks, New Tunnels per Treatment Panel ..... 50 3 7 Average Interspecies Blocks, Unblocks, New Tunnels for Paired Panels .......... 50 3 8 Average Interspecies Territory Areas Before and After Panel Connection ......... 52

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8 LIST OF FIGURES Figure page 1 1 Florida populations of Coptotermes formosanus mixing ................................ ..... 18 1 2 Mutual agonism between Coptotermes formosanus termites ............................. 18 1 3 Suicide cramming ................................ ................................ ............................... 18 2 1 Two panel bioassay design ................................ ................................ ................ 23 2 2 Observed dead over time in intraspecies panels ................................ ................ 28 2 3 Marbling. ................................ ................................ ................................ ............. 28 2 4 Intraspeci es observed marbled dead over time. ................................ ................. 29 2 5 Coptotermes formosanus abandon main chamber. ................................ ............ 29 2 6 Corner Blocking. ................................ ................................ ................................ . 30 2 7 Burials leading to blockages. ................................ ................................ .............. 31 2 8 Blocking and unblocking ................................ ................................ ..................... 31 2 9 Burial of dead by Coptotermes formosanus ................................ ....................... 32 2 10 New tunnels formed to bypass blockages in unbaited Coptotermes formosanus panel ................................ ................................ ............................... 32 2 11 New tunnels formed to bypass blockages in baited Coptotermes formosanus panel ................................ ................................ ................................ ................... 32 2 12 Intraspecies blockages, unblockages, and new tunnels over time in control panels. ................................ ................................ ................................ ................ 34 2 13 Int raspecies blockages, unblockages, and new tunnels over time in treatment panels. ................................ ................................ ................................ ................ 35 2 14 Dead observed over time in reinvading C. formosanus foraging population after feeding on bait ................................ ................................ ............................ 37 2 15 Observed dead marbled reinvading C. formosanus over time. ........................... 37 2 16 Fungi growing on bait. ................................ ................................ ........................ 38 3 1 Observed dead over time in interspecies panels ................................ ................ 45

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9 3 2 Observed dead during the reinvasion of a baited Coptotermes formosanus panel by neighboring Reticulitermes flavipes ................................ ...................... 46 3 3 Interspecies observed marbled dead over time ................................ .................. 46 3 4 Typical encounter between Coptotermes formosanus and Reticulitermes flavipes ................................ ................................ ................................ ............... 48 3 5 Choke points in Reticulitermes flavipes control panels ................................ ....... 48 3 6 Boundary walls between species in control panels ................................ ............. 49 3 7 Presoldier production in Reticuliterme s flavipes ................................ ................. 49 3 8 Interspecies blockages over time in control panels ................................ ............ 51 3 9 Interspecies blockages over time in baited panels ................................ ............. 51 3 10 Differences in gallery layout between Coptotermes formosanus and Reticulitermes flavipes ................................ ................................ ........................ 53 3 11 Invasion of Reticulitermes flavipes by Coptotermes formosanus in control panels ................................ ................................ ................................ ................. 53 3 12 Interspecies trial before connection ................................ ................................ .... 54 3 13 Interspecies trial after connection and reinvasio n ................................ ............... 54

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10 LIST OF OBJECTS Object page 1 1 Two Florida C. formosanus colonies encounter (.mp4 file 180MB) .................... 19 1 2 Two New Orleans C. formosanus colonies encounter (.mp4 7MB) .................... 19 3 1 R. flavipes soldiers defend, workers retreat from C. formosanus (.mp4 file 50.8MB) ................................ ................................ ................................ .............. 49 3 2 R. flavipes retreats from injured C. formosanus (.mp4 file 29.7MB) ................... 49

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11 LIST OF DEFINITIONS Blockage Occlusion of the tunnel. Most often occurs when a termite dies and substrate is moved to cover the body of the deceased. Choke point A restriction found in Reticulitermes flavipes tunnels that may be occluded by substrate. Does not occur from a burial of a deceased termite. Foraging population A group of termites collected from the same colony, consisting of workers and soldiers, with the exclusion of reproductives. Marbling Believed to be caused by the build up of uric acid in the fat body of unhealthy, stressed termites. The body of the termite is observed as more opaque, which contrasts strongly with dark gut contents forming a crescent shape in the abdomen. New tunnel A n ew passage created around or in avoidance of a blockage. Suicide cramming Upon an aggressive interception between two colonies, multiple termites crowd together, wedging themselves within the tunnel, completely blocking the passage and killing themselves in the process. Unblock Tunnel obstruction is removed.

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12 Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science RESPONSE OF SUBTERRANEAN TERMITES TO NEIGHBORING POPULATIONS OF INTER AND INTRASPECIFIC COLONIES AFTER BAITING By Sarah J. Bernard M ay 2015 Chair: Nan Yao Su Major: Entomology and Nematology Coptotermes formosanus Shiraki and Reticulitermes flavipes Kollar are sympatrically occurring and economically important subterranean termites in the South e astern United States. Blocking of tunnels, unblocking of tunnels, and new tunnel construction was observed in intra and inter species experiments. Intraspecie s experiments in two dimensional arenas in which one population of C. formosanus was baited resulted in elimination of baited termites and subsequent reinvasion of territory. Territories held by unbaited termites increased significantly, nearly doubling af ter reinvasion. Intraspecies controls did not have a significant change in territory areas. In interspecies experiments, R. flavipes , retreated and blocked tunnels at choke points as C. formosanus advanced. R. flavipes reinvaded baited C. formosanus territ ory in one repetition. Reinvading termites of both species were eliminated by bait.

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13 CHAPTER 1 THESIS INTRODUCTION Economic I mpact Termites (Isoptera) ar e important structural pests, it was estimated that in the United States, termite control alone cost about $1.5 billion in 1994 {Su, 2000 #45} , and an estimated $40 billion is spent annually on termite control and repairs globally ( Rust and Su 2012 ) . About 80% of this cost is due to subterranean termites ( Su 1994 , Su and Scheffrahn 2000 , Rust and Su 2012 ) such as Coptot ermes formosanus Shiraki in the Southeast ern part of the United States and Hawaii, and Reticulitermes flavipes Kollar which is found throughout the continental United States ( Su and Tamashiro 1987 , Su and Scheffrahn 1990 ) . These s pecies are highly destructive and are often found infesting wooden structures. In areas of high termite pressure, such as New Orleans, s tructural repair s may cost more than five times the cost of preventative measures and control ( Su and Scheffrahn 2000 ) . Termite M anagem ent History Control of subterranean termites once depended on liquid termiticide barriers ( Su and Scheffrahn 1990 , Su 1991 ) ; from the ear ly 1900s, chemical barriers have been used in pre construction of buildings to protect them from future infestations ( Randall and Doody 1934 ) . Between the 1950s and 1980s , c yclodienes were u sed extensively because they were relatively inexpensive, effective, and persisted in the environment . These chemicals were withdrawn in the mid 1980s because of environmental persistence and concern s about bioaccumulation in the environment ( Su and Scheffrahn 1990 , 1998 ) . Currently, the leading liquid termiticide treatments use fipronil

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14 or imidacloprid as active ingredients, although concerns ov er whether or not these methods are able to effectively control entire termite colon ies have been raised ( Su 2005 ) . Termites killed by liquid termiticides o ften cause other colony members to avoid a treated area, this is called secondary repellency . Thi s is not a desirable effect in baiting strategies since termites must come into contact with and share toxicants with the rest of the colony. Baiting The growing concern over pesticides in our environment has led to innovations in termite control; a bait containing insect growth regulators can safely be used in situations in which liquid termitici des cannot ( Cabrera and Thoms 2006 ) . It has been established that the use of a bait ing system is effective in suppressing and eliminating field colonies of subterranean termites ( Su et al. 1991b , Su 1994 , Su et al. 1998 , Getty et al. 2000 , Grace and Su 2001 ) . Th e active ingredient in an effective baiting strategy must be slow acting, non repellent, and lethal time has to be dose independent ( Jones 1984 , Su et al. 1985 , Haverty et al. 1989 , Su and Scheffrahn 1989 , 1998 ) . These properties allow for the spread of active ingredient throughout the colony by trophallaxis, or regurgitation of food and bait material. Insect growth regulators such as c hitin synthesis inhibitors (CSIs) are currently used in commercial baits, and have been documented to eliminate subterranean termite colonies on many occasions ( Su 1994 , Su et al. 1998 , Getty et al. 2000 , Grace and Su 2001 , Messenger et al. 2005 ) . Noviflumuron is one of the current CSI s being used in baiting systems; the toxicant has exhibited a greater speed of action and efficacy than its predecessor, hexaflumuron ( Karr et al. 2004 , Rust and Su 2012 ) .

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15 Territory Reinvasion O ne concern of baiting technology is the possibility that previous ly treated areas will become reinfested by nearby colonies of subterranean termites ( Grace et al. 1996 , Getty et al. 2000 , Vargo 2003 , Messenger et al. 2005 ) . In the field, elimination of a C. formosanus colony and subsequent occupation of former territory by another subterranean termite colony has been documented ( Grace and Su 2001 , Messenger et al. 2005 ) . Questions have been raise d about the sp eed of reinvasion of territories. S ubterranean termites can reclaim an area rapidly in regions of high termite density ( Messenger et al. 2005 , Mullins et al. 2011 ) . Due to the cryptic na ture of subterranean termites, it is difficult to obse rve this behavior in the field. Indirect means of monitoring, such as m ark recapture studies , have found termites from a nearby marked colony appear in monitoring stations that were previously inactive after the elimination of a colony ( Getty et al. 2000 , Messenger et al. 2005 ) . Older colonies may grow to include territory once its neighboring colony is eliminated , or smaller coloni es may take the o pportunity to grow and claim newly available territory . Reticulitermes sp . have also been found inhabiting monitoring stations which were once populated by C. formosanus ( Messenger et al. 2005 ) . Knowledge of how a colony responds to a bait ed or eliminated neighboring colony would provide insight on how best to manage areas of known termite infestation. Agonism Laboratory observation. Laboratory studies on agonistic behavior and inter and intra colonial aggression have shown varying results. Studies dealing with agonism between different populations of C. formosanus have been previously observed using

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16 petri dish bioassays ( Su and Haverty 1991 , Cornelius and Osbrink 2003 ) but these conditions are unnatural and do not simulate termite tunnel structure s . More recent studies have been conducted with the use of planar arenas ( Li et al. 2010 , Chouvenc et al. 2011 ) which are more similar to field conditions than petri dish bioassays, while still allowing observation of termites . Interspecies agonism. In laboratory foraging a renas, different species of subterranean termites separate d themselves by building barriers ( Li et al. 2010 ) and in some cases blocked tunnels with their bodies. I t has been suggested that inter colonial aggression may be a mode of determining foraging territories amongst neighboring colonies ( Levings and Adams 1984 , Jones 1993 , Messenger and Su 2005b , Li et al. 2010 ) . Intraspecies agonism. S ome colonies of the same species may mix and subsequently co habitate in the same territory ( Su and Haverty 1991 , Messenger and Su 2005b ) . Florida colonies of C. formosanus were found to have no agonistic response with each other , most likely due to the low genetic diversity of the introduced population ( Su and Haverty 1991 ) . In Florida, colony fusion has been docum ented in C . formosanus ( Su and Scheffrahn 1988a ) and colonies have been found sharing the same tunnels and monitoring stations. Preliminary laboratory experiments confirmed that foraging populations of C. formosanus from Florida mix ed and fuse d with no instances of aggression observed between workers and soldiers (Figure 1 1, Object 1 1), which supports the field result found in Broward County, Florida by Su and Scheffrahn (1988b) .

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17 In contrast, colony mixing has not been observed in New Orleans colonies ( Husseneder et al. 2003 , Messenger and Su 2005b ) . This is b ecause C. formo sanus colonies in New Orleans most likely arose from multiple introductions to the port city . These colonies exhibit a scenario in which there is high termite pressure, and one . This makes the area, and areas like it, difficult to manage. Preliminary experiments found that New Orleans populations of C. formosanus had mutually agonistic responses (Figure 1 2, 1 3, Object 1 2) in which fighting and boundaries between populations w ere created, which may resemble the natural condition of neighboring field colonies ( Li et al. 2010 ) . For this reason, New Orleans populations of C. formosanus were chosen for intraspecies experiments. Necrophobic Behavior It has been noted that termites may avoid and retreat from tunnels in which aggressive encounters have occurred ( Messenger and Su 2005b ) . This is thought to be because of dead and or decaying termite bodies left in the tunnels ( Messenger and Su 2005b , Su 2005 , L i et al. 2010 ) . It has been suggested that decaying cadavers may have an impact on termite tunneling behavior and they may be a site of substrate deposition ( Su et al. 1982 , Li et al. 2010 ) . A voidance may be caused by a colony odor , such as from the chemical oleic acid, that wil l eventually dissipate after the colony is eli minated ( Messenger and Su 2005b ) . More insight into avoidance behavior associated with deca ying cadavers may be useful in this regard ( Li et al. 2010 ) . Despite this avoidance behavior, termites have been observed removing blockages from tunnels, as well as creating new tunnels, bypassing blockages ( Li et al. 2010 ) .

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18 Objectives To study the behavioral response s of C. formosanus and R. flavipes to neighboring baited C. formosanus colon ies . Figure 1 1 . Florida populations of Coptotermes formosanus mixing. No agonism occurred. Figure 1 2 . Mutual agonism between Coptotermes formosanus termites . W orkers and soldiers are both aggressive, biting at unrelated individuals. Figure 1 3 . Suicide cramming. Two C. formosanus foraging populations encounter, results in death and blockage of tunnel.

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19 Object 1 1. Two Florida C. formosanus colonies encounter (.mp4 file 180MB) Object 1 2. Two New Orleans C. formosanus colonies encounter (.mp4 7MB)

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20 CHAPTER 2 INTRASPECIES INTERACTIONS Introduction The use of a slow acting toxicant in the field is important for reduction of the colony population ( Su and Scheffrahn 1993 , Su 1994 , Su et al. 1998 ) . According to Su (2005) , laboratory population s of termites began to die two weeks aft er feeding on 0.5% noviflumuron bait (Recruit HD, Dow AgroSciences) and were eliminated by week seven . It was documented that termites of neighboring colonies may begin appearing in territories of treated colonies before they were completely eliminated , sugge sting that ( Messenger et al. 2005 ) . In the following experiment, I hypo thesized that termites would 1) invade a neighboring territory immediately upon encountering the weakened termites , 2) they would not invade a panel with accumulated dead due to decaying termite cadavers , 3) new tunnels would be created more than unblockages 4) and territories of untreated control populations would not change . In this chapter we explored the effects of baiting with noviflumuron o n neighboring unrelated foraging populations of the same species. We observed the decline of termite health, intraspecies agonism, the fate of blockages in tunnels, shifts in territory, and reinvasion of an eliminated foraging population by neighboring populations .

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21 Materials and Methods Termites New Orleans C. formosanus were used in this bioassay after observations from the preliminary study resulted in mutual agonism without full elimination of either foraging population . Coptotermes formosanus individuals were collected from New Orleans, L A laboratory colonies maintained as described by Su (2013) . C olonies were lab reared from swarm captured reproductives, and were at least 5 years old . T ermites from different colonies were extracted as needed by use of a moistened roll of corrugated cardboa rd placed inside the enc losure ( La Fage et al. 1983 ) . Foraging populations were marked with 0.1% Nile Blue A solution (Fisher Scientific, Pittsburgh, PA) (wt:wt) , so that different coloni es could be easily identifie d. Groups of about 300 termites were placed in glass petri dishes with 90mm filter paper rounds (Whatman, UK) moistened with DI water or Nile Blue solution for 5 days prior to the start of the experiment to allow the dye to be stor ed in the fat body ( Su et al. 1991a , 1993 , Su and Scheffrahn 1996 ) . Planar Foraging Arena Bioassay P lanar arenas similar to those described by Li et al. (2010) and Chouvenc et al. (2011) were used for this experiment (Figure 2 1) . Arenas were constructed ou t of 2 sheets of clear acrylic (60 x 60 x 0.6 cm thickness) with laminate strips of 5cm width and 0.15 cm thickness placed around the margins between the two pieces to form a frame. Screws held t he two pieces of clear acrylic sheets on e ither side of the laminate strips to form a livable area of 100 square cm with a thi ckness of 0.15 cm ( Li et al. 2010 ) . 1 cm d iameter holes were drilled into the top acrylic panel in each of the four corners of the arena to allow for attachments of a chamber (8.5 cm height, 6.5 cm diam eter ) which

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22 was the site of introduction , and connection ports. Four 1 cm diameter holes were drilled in the center of the arena to accommodate for the attachment of a second chamber, which allows access to either a food or bait source. The center chamber had a 2.5 cm by 2.5 cm square sponge (Ocello, 3M, St. Paul, MN) attached to the chamber cover by way of a nail and hot glue. The sponge was moistened with 10 ml DI water, which was sufficient to maintain humidity throughout the length of the experiment. A 2.5 cm diameter sheet of clear acrylic was secured between the two panels in the center of the arena to maintain equal spacing, and d id not obscure the holes allowing access to the chamber. Connection ports were made of clear acrylic , cut to the desired shape with a band saw. Three connection ports were secured to the remaining corners of the arena setup with washers and hex nuts . Sifted (35 100 mm sieves) multipurpose sand ( Sakrete , Charlotte, NC ) was washed with deionized water (DI) , sterilized by oven at 100 º C for at least 48 h ours , and cooled to room temperature before use. Sand was funneled into the void between the two acrylic panels. Sand was moistened with DI water by way of the access holes in the center of the arena. Two of the panels described were set side by side and comprised one bioassay. Stoppers were placed at the ends of the connection ports, provisioned with a piece of wood, to encourage termites to leave the connec tions open until connected. Introduction and food chambers were provisioned with c ut spruce wood sticks (6 cm x 1 cm) which had been soaked in DI water for at least 24 hours. Chambers d esignated for baiting had their wood replaced with a 2 cm cubed block of 0.5% noviflumuron bait once the te rmites encountered the chamber (Figur e 2 1) .

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23 Figure 2 1 . T wo panel b ioas say design. Arenas made from clear acrylic sheets and laminate strips. Total foraging area of each panel 100 cm 2 filled with sterile sand. Each panel had one introduction chamber at one corner, and one feeding chamber in the center of the arena. Procedure Dyed and undyed C. fo r mosanus foraging populations were added to separate panels described in the two dimensiona l foraging bioassay section. F our pairs of control panels and four pairs of panels in which one panel was baited, for a total of 16 arenas , were used for this experiment. One thousand termites consisting of 900 workers and 100 s oldiers ( Haverty 1979 ) were added to each panel , and were allowed to tunnel throughout the ir panel of introduction until they reac hed the central feeding chamber, which contained wooden sticks, and corner connection ports . Controls were then connected by connection ports with 50 cm lengths of 4 mm ID x 6 mm OD Tygon® tubing (Saint Gobain Performance Plastics, Northboro, MA ) with 2.5 cm lengths of 6 mm ID x 9 mm OD clear vinyl tubing (Watts, North Andover, MA) , about six days after their introduction. Ba ited panels had their wood replaced with bait after termites

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24 encountered the feeding chamber about six days after introduction. Termit es in baited panels w ere monitored for symptoms of intoxication, which may include ataxia and sluggishness ( Karr et al. 2004 ) , death , and marbling . D aily counts of observed dead and marbled individuals were recorded. Marbling was defined by an opaque white body with a visible dark contrast of the gut (Fig 2 4 ). Panels with bait were connected when roughly 25% cumulative population mortality and a large spike in deaths had been observed , on average 33 days after introduction . Data Collection Control and treatment groups were replicated four times to determine tota l cumulative dead observed, cumulative marbled termite s dead , blockages, unblockages, new tu nne ls, and territory areas. Panels to determine t otal dead and marbled dead termites of the reinvading foraging population were replicated three times . Panels were checked twice a day for dead worker termites , marbled dead worker termites, blockages, unblo ckages, and new tunnels . Dead worker termites were counted by marking on top of the clear acrylic sheet with a permanent marker by circling cadavers to ensure they would not be counted more than once. Blockages, unblockages, and new tunnels were also noted on top of the acrylic panel along with the da te on which they were observed . Data were taken for 50 days in control and treatment panels. Reinvading termites contin ued to be observed for deaths for an additional 20 days after the elimination of baited termites . To estimate territory areas occupied, b ack lit photos of panels were taken the day of conn ection and a month after connection. Photos were taken with a Samsung Galaxy 4s ( Samsung, Ridgefield Pa rk, NJ) .

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25 Data Analysis Variables . Number of dead worker termites and marbled dead worker termites, blockages, unblockages, and new tunnels observed per day were recorded and a cumulative count was obtained over the course of 50 days . Cumulative counts over time were averaged for rep licates and graphed. The cumulative number of dead termites observed before panel connection and the cumulative number of dead termites observed at the end of the experime nt were analyzed using a T test with a significance l evel of 0.05 (Microsoft Excel 2013, Microsoft Corp., Redmond, WA ). Panels were either treated singly (per panel, baited or unbaited) and compared or taken together as a pair of panels (control and treatment) during analysis. Territory area. Territory areas were obtained b y analyzing panel photos with GNU Image Manipulation Program ( GIMP 2.8.14 ) soft ware to estimate tunnel area before and one month after panels were connected . Territory was considered occupied if termites were found alive within a tunnel and tunnels connected to it without blockages present. The difference in t erritory areas were analyzed by a T test with a significance level of 0.05. Results Decline of Termite Health Dead observed . The number of dead termites i n baited panels increased over time compared to either control panels or unbaited panels (Figure 2 2) . On average, the first dead from baiting were observed around nine days after termites were found in the baiting chamber (Table 2 1) . Termite deaths reached their peak arou nd day 1 8. Termites die d until fe w were observed in tunnels and most remaining live termites were in the introduction chamber or the bait chamber, around day 26. At this time, panels

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26 were connected and neighboring unbaited termites invaded the moribund termites p anel ( Table 2 1 ). There was no significant difference in the number of dead observed in control panels (Table 2 2). Significantly more baited termites were observed dead in baited treatment panels as compared to unbaited treatment panels (Table 2 3). Signi ficantly more termites were observed dead in treatment panels as compared to control panels (Table 2 4). Marbling . The bodies of termites transformed from shiny and near translucent to opaque white, with a clearly contrasting darkened gut , causi ng a marbli ng effect (Figure 2 3 ). Marbled workers only appeared in baited colonies, on average, 21.5 days after baiting (Table 2 1 ) . Observation of marbled individuals increased until all remaining termites obse rved were marbled, on average 35 days after baiting (Table 2 1 ) . These marbled individuals persisted for a number of days before they were eliminated by a neighboring foraging population . Marbled in dividuals did not appear in control groups or in unbaited groups ( Figure 2 4 ). Abandoning main chamber. A la rge chamber or enlarged tunnel was found in C. formosanus galleries in all repetitions . Termites were observed dying in this large primary chamber, most likely when attempting to molt. When termites began to become affected by bait, group members cannibali zed dying and or dead termites found in and around this chamber. Termites then buried dead individuals in this chamber. Lastly, a large spike in deaths occurred before termites abandoned the chamber without further burial of dead, and then termites relocat ed to another area of the panel. New chambers

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27 were sometimes observed being created by enlarging a tunnel after abandoning the first cham ber (Figure 2 5 ). Repair and defense . Baited populations often did not block corner s of arena s after an invasion attemp t was made by a neighboring foraging group . Healthy populations completely filled the corner connector and the surrounding area with sand whe n an invasion attempt was made (Figure 2 6 ). Table 2 1. Phases of Decline in Intraspecies Baited Treatment Panels Number of Days First Dead Observed 9.00 ± 0.91 Death Spike 18.25 ± 2.29 First Marbled Observed 21.50 ± 1.94 Number of Deaths Slow 26.00 ± 3.19 All Remaining Marbled 34.75 ± 1.44 First Marbled Dead 34.75 ± 2.02 Baited Eliminated 43.00 ± 1.58 Table 2 2. Percent Cumulative Dead Observed in Intraspecies Control Panels n = 4 Control 1 Control 2 T P Sig. At connection 0.00 ± 0.00 0.00 ± 0.00 End 3.31 ± 0.89 3.63 ± 1.76 0.34 0.377 ns Table 2 3. Percent Cumulative Dead Observed in Intraspecies Treatment Panels n = 4 Unbaited C .f . Baited C.f . T P Sig. At Connection 0.08 ± 0.03 33.50 ± 5.85 5.70 0.0 05 s End 2.57 ± 0.57 44.76 ± 4.64 8.46 0.0 02 s Table 2 4. Percent Cumulative Dead Observed in Intraspecies Paired Panels Control Treatment T P Sig. At Connection 0.00 ± 0.00 33.58 ± 5.84 5.75 0.005 s End 6.94 ± 2.62 47.32 ± 4.34 7.68 0.002 s

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28 Figure 2 2. Observed dead over time in intraspecies panels. Few dead were observed in control and unbaited panels, the number of dead oberved in baited treatment panels increased over time. A B Figure 2 3. Marbling. A) Healthy, unaffected termites. B) Affected termites with opaque appearance.

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29 Figure 2 4 . Intraspecies observed marbled dead over time. Dead marbled C optotermes formosanus were observed in baited treatment panels. Marbled did not appear at all in control or unbaited panels . A B Figure 2 5. C optotermes formosanus abandon main chamber. A) Chamber with active termites at left. B) Termites bury dead in main chamber, leave and enlarge tunnel for new chamber (asterisk).

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30 A B Figure 2 6 . Co rner Blocking. A) The corner of a control C optotermes formosanus panel before an invasion attempt b y neighboring foraging group was made. B) After invasion attempt, a healthy population fill ed this corner with sand. Intraspecies Agonism F oraging populations responded aggressively towards each other, with soldiers and workers biting at unrelated individuals (Figure 1 2 ). Upon an aggressive interception between two populations, many termites crowded together, wedged themselves within the tunnel, and completely blocked the passage which killed them in the process (Figure 1 3) . T his has been previously observed as an extreme form of ( Messenger and Su 2005b , Li et al. 2010 ) . This behavior le d to a large deposition of substrate creating a blocka ge of the tunnel. Foraging populations were observed making invasion attempts and neither group retreated upon the encounter . D ead termites beca me the site of substrate deposition, which led to a cordoning off of the area (Figure 2 7 ) . Foraging populations were observed reinvading tunnels in neighboring panels . Groups unblock ed a tunnel by removing the substrate from the area of a blo ckage (Figure 2 8 ). Invading groups were observed pushing dead termites and substrate against the walls of tunnels when they reinvade d a panel and encounter ed unburied

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31 dead (Figure 2 9 ) . Invaders buried cadavers left in the p anel, but often these burials did not lead to blockages, maintaining the usabilit y of the tunnel. Excavation of n ew tunnels were observed (Figure 2 10, 2 11 ) . A B Figure 2 7 . Burial s leading to blockages. A) T unnels before invasion of the panel . B) After invasion, dead were buried (asterisks) , leading to a blockage in the tunnel. A B Figure 2 8 . Blocking and unblocking. A) Tunnel blocked as a result of agonism between two C optotermes formosanus foraging populations . B) Unblocking of tunnel by C. formosanus (asterisks) .

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32 A B Figure 2 9 . Burial of dead by Coptotermes formosanus . A ) Dead left by invading foraging population . B ) Invading group buried dead with substrate but left the tunnel usabl e. Invading termites d id not consume unrelated dead. A B Figure 2 10 . New tunnel s formed to bypass blockage s in unbaited Coptotermes formosanus panel . A) Blocked tunnel passage. B) Ne w tunnel (asterisk) to bypass blockage . A B Figure 2 11 . New tunnels formed to bypass blockages in baited Coptotermes formosanus panel . A ) Blockages formed around dead from baiting. B) New tunnel (asterisk) created bypass blockage s (blue) in baited panel.

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33 Blockages, Unblockages , and New T unnels Controls. There were no significant difference s in variables between control panel 1 and control panel 2 (Table 2 5 ) . Blockages , u nblocking of tunnels , and new tunnel creation increased over time after panels were connected (Figure 2 1 2 ) . Healthy populations encounter ed each other, a fight occur red , and then a blockage at the site of encounter wa s formed. Health y termites did this multiple times until blockages formed a boundary between the populations at the corner or i n the tubing that connected panels . Baited p anels. As compared with unbaited panels , the number of blockages and unblockages were significantly higher in baited panels. There was no significant difference in new tunnel creation bet ween unbaited or ba ited panels (Table 2 6 ). B lockages started appearing before panels were connected and continued to increase after panels were connected. Unblock ages were observed after panels were connected (Figure 2 1 3 ) . In baited panels , termites were obs erved burying their dead and caused b lockages in their own panel before panels were connected. Blockages inc reased after connection, due to invasion by neighboring termites . After panel connection, unblock ages were observed by invading termites i n the baited termites f ormer galleries . Control vs. treatment panels. As compared with control panels, treatment panels had significantly more blockages and unblockages . The number of new tunnels created was not significantly different between the control and treatment (Table 2 7 ) .

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34 Tab le 2 5 . Average Intraspecies Blocks, Unblocks, New Tunnels per Control Panel n = 4 Control 1 Control 2 T P Sig. Blocks 1.50 ± 0.50 7.00 ± 3.54 1.3 9 0.130 ns Unblocks 0.25 ± 0.29 1.75 ± 1.18 1.13 0.170 ns New Tunnels 0.25 ± 0.29 0.50 ± 0.29 1 .00 0.195 ns Tab le 2 6 . Average Intraspecies Blocks, Unblocks, New Tunnels per Treatment Panel n = 4 Unbaited C .f . Baited C.f . T P Sig. Blocks 5.50 ± 3.66 21.00 ± 3.56 2.4 4 0.046 s Unblocks 1.75 ± 1.04 10.75 ± 2.50 3.2 2 0.024 s New Tunnels 0.00 ± 0.00 1.75 ± 1.04 1. 70 0.094 ns Tab le 2 7 . Average Intraspecies Blocks, Unblocks, New Tunnels for Paired Panels Control Treatment T P Sig. Blocks 8.75 ± 3.33 26.50 ± 4.77 2.7 1 0.03 7 s Unblocks 2.0 0 ± 1.08 12.5 0 ± 2.53 4.04 0.01 4 s New Tunnels 1.00 ± 0.58 1.75 ± 1.03 0.6 8 0.27 4 ns Figure 2 1 2 . Intraspecies blockages , unblockages, and new tunnels over time in control panels.

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35 Figure 2 1 3 . Intraspecies blockages , unblockages, and new tunnels over time in treatment panels . Territory S hift There was not a significant difference between tunnel areas oc cupied before and after foraging populations encountered each other in control intraspecies panels (Table 2 8 ). A significant difference was found between tunnel areas occupied by the un baited foraging population before and after they encount ered a neighboring baited group . The area of the territory roughly doubled in size when a healthy group en countered the neighboring group in decline (Table 2 8 ) . Conversely, the baited territory was reduced to zero, which was significantly different from t he beginning territory area (Table 2 8 ) . Baited foraging populations were eliminated within the first week of panel connection , though reinvasion did not always oc cur by the neighboring foraging population immediately. B ut b y one

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36 month after connection , all baited foraging population territories were reinvaded by neighboring termites . I nvasion was initiated by both healthy, unbaited termites as well as un healthy, ba ited termites , although unhealthy termites were unsuccessful at invasion . Table 2 8 . Average Intraspecies Territory Areas Before and After Panel Connection Area (cm ²) n Before After T P Sig. Control 1 4 374.33 ± 42.56 317.82 ± 29.80 1.39 4 0.12 9 ns Control 2 4 386.70 ± 40.60 465.48 ± 20.44 1.47 3 0.11 9 ns Unbaited C .f. 4 547.44 ± 52.61 998.80 ± 110.76 7.350 0.00 3 s Baited C.f. 4 466.98 ± 53.85 0.00 ± 0.00 8.671 0.00 2 s Mean Area ( cm² ) ± S E Death o f R e i nvaders Secondary kill. It took about 43 days on average from termite baiti ng to elimination of the foraging population . L ive termites were no longer observed , or an invading foraging population killed the remaining termites. As termites reinvaded th e area, they unblocked tunnels , a nd continued to use old gallery passages. Passa ges leading to bait were found by the invading termites , and feeding on bait ensued. L ive termites were no longer observed in the invading population 94 days after the beginning of the experiment on average, w hich was 51 days after they begun to (Figure 2 1 4 ) . Dead marbled reinvading termites were observed starting at day 80 , which was 37 days after invaders begun feeding on bait ( Figure 2 1 5 ).

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37 Figure 2 1 4 . Dead o bserved over time in reinvading C. formosanus foraging population after feeding on bait. T hree baited foraging populations had been reinvaded by neighboring C. formosanus , dead of reinvading termites over time was observed. Figure 2 1 5 . Observed d ead marbled reinvading C. formosanus over ti me. Marbling was observed in reinvading termites .

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38 Additional Observations Fungi. Black colored fungi begun to grow on old er bait towards the end of experiments . Molding bait has been observed in the field, and there is no detriment currently a ssociated with this occurrence . Molding bait was examined and samples were taken for plating (Figure 2 1 6 ) . Figure 2 1 6 . Fungi growing on bait. Discussion Termite health. In the current study, foraging populations of baited C. formosanus congregated and died in an enlarged chamber within their galleries , which is similar to a finding in extended foraging experiments where baited termites died in panels which had higher termite density ( G. Kakkar, personal communicatio n) . This

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39 may be because termites require assistance to molt, or must acquire new gut bacteria soon after molting from other colony members by participating in proctodeal trophallaxis. B aited foraging populations did not fill and block corners after confron tation s as thoroughly as unbaited, healthy termites did. This may be similar to colonies in declining health that fail to repair their c arton nest after being damaged. Marbling . It has been noted that termites collected from bait stations in the field may thought to be caused by a buildup of uric acid ( Li et al. 2010 , Xing et al. 2014 ) . This was observed in bioassays with baited termites . Termites began to look less translucent, individuals with a gut that starkly contrast s with the fat body were observed , on average, 23 days after termites were found on bait. As termites age, they molt less frequently ( Alibert and Martoja 1976 , Grassé 1982 ) which would allow for build up or uric acid in the fat bo dy by way of bait and dead nest mate consumption, without immediate death . Marbled individuals were observed 5 days after a spike in dead were observed. These marbled termites may be a result of i ncurring higher levels of stress after a large die off, or more dead and dying being cannibalized by remaining termites . Considering the length of time it takes for marbling to be easily observable, it is likely that these affected individuals were older members of the foraging population and thus molted inf reque ntly. T he ratio of marbled termites to non marbled termites had increased over time as non marbled termites die d , with marbled termites only being observed dead towards the end of experiments .

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40 Baiting and r einvasion . Although baited termites were eliminate d in roughly 43 days , and reinvading termites were eliminated by bait after an additional 51 days, t his study is not necessarily indicative of how long it might take to eliminate a colony or reinvading colony . Several fac tors of this experiment were artificial and may have impacted termite health or behavior. Points at which termites could encounter were very controlled; in a natural environment there may be several points at which two colonies may interface. This may increase the opportunities for in vasion or reinvasion, although even in our restricted bioassays this occurred in a matter of days after panel connection. The length of time and amount of confrontations experienced in these arenas may have also put added stress on termites that may have a ffected their health or behavior. Despite these concerns, this study clearly shows the elimination of termites and then reinvading termites by baiting. This study confirmed reinvading termites use the same galleries and monitori ng stations as the former in habitants, a s suggested previously by Messenger et al. ( 2005 ) . This supports maintaining a baiting program as a means of continued area wide suppression in areas of high termite density .

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41 CHAPTER 3 INTERSPECIES INTERACTIONS Introduction Florida col onies of C optotermes formosanus have been found to have no agonistic response with each other , most likely due to the low genetic diversity of the introduced population ( Su and Haverty 1991 ) and the fusing of Florida C. formosanus colonies has been docu mented ( Su and Scheffrahn 1988a ) . However, agonism between C. formosanus and R eticulitermes spp. have been observed. Colony agonism may be a necessary mechani sm for maintaining territories between colonies ( Li et al. 2010 ) . Field studies ascertained that multiple termite species are able to cohabitate in neighboring territories for years at a time ( Deheer and Vargo 2004 , Messenger and Su 2005a ) . I t has also been observed that in the event of elimination by baiting, t erritories may shift to absorb a neighboring territory ( Vargo 2003 ) . Reinvasion of C. formosanus territories after elimination has been documented in Florida as well as other locations such as Hawaii and New Orleans ( Messenger et al. 2005 , Mullins et al. 2011 ) . In some cases, reinvasion of C. formosanus territories by Reticuli termes sp . ha ve been observed ( Messenger et al. 2005 ) . I hypothesized that 1) in unbaited controls, C. formosanus would invade R eticulitermes flavipes and take over their territory, 2) baited C. formosanus would be too weak to invade neighboring R. flavipes , and 3) R. flavipes would invade C. formosanus after termites are eliminated . In this chapter we explored the effects of baiting with noviflumuron on neighboring foraging populations of different species of termites . We observed the

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42 dec line of termite health, inter species agonism, the fate of blockages in tunnels, shifts in territor y, and reinvasion of an eliminated foraging population Materials and Methods Termites Foraging populations of R. flavipes and C. formosanus were collected from Secret Woods and Anne Kolb Nature Center s in Broward County, Florida , respectively. Termites were collected in the manner described by Su and Scheffrahn (1986) . Fie ld collection of termites occurred between September, 2013 and November, 2014 as necessary. Soldiers available from foraging populations in Florida colonies differed in abundance from soldiers available in New Orleans foraging populations, and therefore th e ratio of soldiers to workers had to be modified for this experiment. Planar Foraging Arena Bioassay Set up of the bioassay is the same as the modified intraspecie s bio assay outlined in Chap t er 2 . Baiting and Reinvasion E xperiments Arenas were provisioned with foraging populations of either 970 C. formosanus workers (undifferentiated larvae of at least the third insta r) and 30 soldiers ( Haverty 1979 ) or 1000 R. flavipes workers and 10 sol diers ( Ha verty and Howard 1981 ) and were left to tunnel for at least 72 hours. The same data were recorded and procedures followed as documented in Chapter 2 for control and treated panels . Only C. formosanus was baited in interspecific experiments. Data Collection Co ntrol panels were replicated four times . Treatment panels were attempted five times, however C. formosanus died u nexpectedly in two replications , so only data from

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43 three replications were used . Because reinvasion of the baited territory only h appened once, there were no replications for dead over time of reinvading R. flavipes in C. formosanus panels. Panels were checked twice a day for dead worker termites, dead marbled worker termites, blockages, unblockages, and new tunnels in the same mann er as explained in Chapter 2 for intraspecies experiments . Data were collected for 60 days in control and treatment panels. R. flavipes continued to be observed for deaths and reinvasion for an additional 15 days after C. formosanus had been eliminated by bait . Beginning and final t erritory areas were photographed in the same manner as expl ained in Chapter 2 for intraspecies experiments . Data Analysis Variables . Number of dead worker termites and marbled dead worker termites, blocka ges, unblockages, and new tunnels observed per day were recorded and a cumulative count was obtained over the course of 60 days. Cumulative counts over time were averaged in replications and graphed. Analysis by T test of observed dead before connection an d at the end of the experiment, blockages, unblockage s, and new tunnels were performed as explained in Chapter 2 for intraspecies data analysis. Territory area. Territory areas were analyzed in the same manner as explained in Chapter 2 for intraspecies dat a analysis . Results Decline of Termite Health Dead observed . Very few dead were observed in either control panels or unbaited panels , however the number of dead termites in baited treatment panels increased over time (Figure 3 1) . On average, the first dead fro m baiting were observed

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44 around six days after termites were found in the baiting chamber (Table 3 1) . Termite deaths reached their peak arou nd day 15 . Termites died until few were observed in tunnels and most remaining live termites were in the int roduction chamber or the bait chamber, around day 2 7 . At this time, treatment panels were connected and Table 3 1 ). There was no significant difference between the number of deaths in C. formosanus and R. flavipes in control panels (Table 3 2 ). There was a significantly higher number of deaths in C. formosanus observed as compared to R. flavipes deaths in baited panels (Table 3 3 ). The number of observed dead C. formosanus i ncreased in baited panels while few dead R. flavipes were observed ( Figure 3 1 ) . There was a significantly higher number of dead in paired treatment panels as compa red to paired control panels (Table 3 4 ). It took on average 54 days for C. formosanus to be eliminated by bait ( Table 3 1) . In one instance, R. flavipes invaded a neighboring C. formosanus panel of moribund termites . As the remaining C. formosanus died off, R. flavipes moved into the panel, began feedi ng on bait, and subsequently were elim inated (Figure 3 2 ). In the remaining replicates, R. flavipes did not attempt to leave their panels of origin before or after neighboring C. formosanus elimination. Marbling. Marbled termites were never observed in control panels or in unbaited treatment panels. In baited treatment panels, C. formosanus marbled began to appear, on average, 24 days after baiting and dead marbled were first observed 42 days after baiting (Table 3 1, Figure 3 3) .

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45 Table 3 1 . Phases of Decline in Interspecies Baited Treatment Panels Number of Days First Dead Observed 6.00 ± 0.58 Death Spike 15.33 ± 0.67 Number of Deaths Slow 27.00 ± 1.73 First Marbled Observed 24.00 ± 2.00 All Remaining Marbled 40.33 ± 0.88 First Marbled Dead 41.67 ± 0.88 Baited Eliminated 53.67 ± 1.33 *Marbled termites were never observed in unbaited panels or controls Tab le 3 2 . Percent Cumulative Dead Observed in Interspecies Control Panels n = 4 Control R.f . Control C.f . T P Sig. At C onnection 0.00 ± 0.00 0.00 ± 0.00 End 2.80 ± 1.37 0.45 ± 0.25 1.94 0.074 ns Tab le 3 3 . Percent Cumulative Dead Observed in Interspecies Treatment Panels n = 3 Unbaited R.f . Baited C.f . T P Sig. At Connection 0.00 ± 0.00 41.45 ± 4.72 8.78 0.0 06 s End 0.80 ± 0.27 45.63 ± 3.50 13.75 0.0 03 s Tab le 3 4 . Percent Cumulative Dead Observed in Interspecies Paired Panels Control Treatment T P Sig. At Connection 0.00 ± 0.00 41.45 ± 4. 72 10.49 6.79121E 05 s End 3.25 ± 1.55 45.90 ± 3. 72 11.82 3.80814E 05 s Figure 3 1 . Observed dead over time in interspecies pa nels. Control and panels had few observed dead over time. Coptotermes formosanus dead observed increased over time in treatment panels .

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46 Figure 3 2. Observed dead during the reinvasion of a baited Coptotermes formosanus panel by neighboring Reticulitermes flavipes . On day 50, R. flavipes invaded neighboring baited C. formosanus panel and subsequently was eliminated after feeding on bait. Though only 115 R. flavipes out of 1000 were observed dead, at the end of the experiment no live termites were observed. Figure 3 3 . Interspecies observed marbled dead over time . Dead marbled C optotermes formosanus were observed in baited treatment panels. Marbled termites were not observed i n control or unbaited panels .

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47 Interspecies Agonism Controls. Both baited and unbaited C. formosanus attempted invasion r egardless of their condition . Upon confrontation, R. flavipes soldiers defend ed area s while workers retreat ed and block ed passages (Figure 3 4 , 3 5 , Object 3 1, 3 2) . Boundaries also formed between species, which were maintained for a period of time, but were eventually breached (Figur e 3 6 ). I n control panels there were very few deaths during c onfrontations . However , R. flavipes was most often displaced or eliminated by C. formosanus once they were no longer able to relocate within panels . In one replicate, n ew presoldiers were observed in R. flavipes as soon as one day after invasion by C. formosanus (Figure 3 7 ). B aited C. formosanus . In treatment panels , baited C. formosanus continued to attempt invasion of R. flavipes panels. R. flavipes retreated and blocked choke points, as in the control panels (Figure 3 4, 3 5) . Bait continued to kill C. formosanus colonies and advancement into R. flavipes colonies ceased . In two out of three panels, R. flavipes populations did not attempt to invade C. formosanus territory even after elimination by bait .

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48 Figure 3 4 . Typical encounter between C optotermes formosanus and Reticulitermes flavipes . C. formosanus invade d R. flavipes territory from the right. R. flavipes retreat ed behind choke points, moving away fr om confrontation, until there was a small buffer zone. C. f ormosanus continue d to dig new tunnels that sometimes intersect ed R. flavipes tunnels. Occasionally new boundary walls form ed , s oldiers were observed on either side . Figure 3 5 . Choke points in R eticulitermes flavipes control panel . A) Open choke point. B) Blocked choke points.

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49 Figure 3 6 . Boundary walls between species in control panels . Walls were reinforced at borders between R eticulitermes flavipes and Coptotermes formosanus territories. Soldiers of both species were observed at these walls before being breached. F igure 3 7 . Presoldier production in R eticulitermes flavipes . R. flavipes molting to presoldiers one day after invasion by C optotermes formosanus . Object 3 1. R. flavipes soldiers defend, workers retreat from C. formosanus (.mp4 file 50.8MB) Object 3 2. R. flavipes retre ats from injured C. formosanus (.mp4 file 29.7MB) Block ages, Unblockages , and New T unnels In control panels, blockages increased as C. formosanus invaded R. flavipes panels (Figure 3 8 ) . Blockages and unblockages in R. flavipes panels were significantly

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50 higher than C. formosanus in control panels (Table 3 5 ) . There was no significant difference between new tunnels in either control panel (Table 3 5 ) . In contrast, baited panels ha d less blockages overall due to lack of R. flavipes movement into neighboring moribund C. formosanus panels (Figure 3 9 ) . There were no significant difference s between blockages , unblockages, or new tunnels in baited C. formosanus and unbaited R. flavipes panels (Table 3 6 ). There were no significant differences between blockages, unblockages, or new tunnels between the control and treatment paired panels (Table 3 7 ). Tab le 3 5 . Average Inter species Blocks, Unblock s , New Tunnels per Control Panel n = 4 Control R.f. Control C.f. T P Sig. Blocks 15.50 ± 2.96 1.50 ± 0.87 4.71 0.009 s Unblocks 4.25 ± 1.31 0.00 ± 0.00 3.23 0.024 s New Tunnels 1.50 ± 0.87 0.00 ± 0.00 1.73 0.091 ns Tab le 3 6 . Average Inter species Blocks, Unblocks, New Tunnels per Treatment Panel n = 3 Unbaited R .f . Baited C.f . T P Sig. Blocks 5.00 ± 5.00 5.67 ± 3.84 0.3 8 0.371 ns Unblocks 2.33 ± 2.33 1.33 ± 0.67 0.48 0.339 ns New Tunnels 0.33 ± 0.33 0.00 ± 0.00 1 .00 0.211 ns Tab le 3 7 . Average Inter species Blocks, Unblocks, New Tunnels for Paired Panels Control Treatment T P Sig. Blocks 16.75 ± 3.64 11.00 ± 3.61 1.09 0.16 2 ns Unblocks 4.25 ± 1.31 3.66 ± 1.67 0.2 8 0.39 6 ns New Tunnels 1.5 0 ± 0.87 0.33 ± 0.33 1. 10 0.161 ns

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51 Figure 3 8 . Interspecies blockages over time in control panels. Blockages, unblockages , and new tunnels increase d after panels were connected on day 5. Figure 3 9 . Interspecies blockages over time in treatment panels . B lockages began before connection. Unblock age s did not occur until after panels were connected on day 43 . F ew new tunnels were observed .

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52 Territory S hift Gallery l ayout. Wider C. formosanus tunnels led to an enlarged portion of a tunnel or a large open chamber in which many termites were observed. Thinner tunnels of R. flavipes were observed to have multiple restrictions between th em, leading to many small chambers at wh ich small groups of termites were found (Fig ure 3 1 0 ) . Control s . In unbaited controls, C. formosanus invaded and reduced R. flavipes territory (Figure 3 11, Table 3 8). R. flavipes moved behind restrictions in tunnels C . formosanus advanced through the tunnels, unblocking or creating new tunnels. R. flavipes continued to block choke points and retreat until they were unable to relocate further. R. flavipes was then forced to confront the invading C. formosanu s, which attacked and caused blockages. Groups of R. flavipes were divided until their populations were no longer functional ( Fig ure 3 11). Baited panel s. In baited colonies of C. formosanus , R. flavipes territory remained unchanged or slightly reduced (Table 3 8). In two out of three trials, R. flavipes did not attempt to invade neighboring territory. In one bioassay, R. flavipes attempted to invade a C. formosanus panel and succeeded. In this panel, R. flavipes absorbed the former C. formosanus territory, and began feeding off the bait within the panel, R. flavipes subsequently was eliminated (Figure 3 13). Table 3 8 . Average Int e rspecies Territory Are as Before and After Panel Connection Area (cm ²) n Before After T P Sig. Control C.f. 4 310.00 ± 35.79 418.12 ± 60.24 3.76 0.016 s Control R.f . 4 114.02 ± 12.44 47.61 ± 24.16 5.4 8 0.00 6 s Baited C.f. 3 343.01 ± 30.98 0.00 ± 0.00 11.07 0.004 s Unbaited R.f. 3 145.64 ± 20.61 239.89 ± 117.14 0.6 9 0.28 2 ns Mean Area ( cm²) ± SE

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53 A B Fig ure 3 1 0 . Differences in g allery layout between Coptotermes formosanus and Reticulitermes flavipes . A) C . formosanus excavated thick tunnels with an enlarged chamber . B) R . flavipes dug thin tunnels with multiple restrictions throughout. A B A ABBBBB Figure 3 1 1 . Invasion of R eticulitermes flavipes by C optotermes formosanus in control panels . A) R. flavipes territory ( green ) , before connection to C. formosanus . B) H ealthy C. formosanus ( red ) invade R. flavipes . After 3 weeks, R. flavipes is confined to top left corner of panel, unable to retreat further.

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54 A B Figure 3 1 2 . Interspecies trial before connection. A) Baited C optot ermes formosanus territory ( red ) before c ompletely eliminated by baiting . B) Reticulitermes flavipes territory ( green ) before panels were connected. A B Figure 3 1 3 . Interspecies trial after connection and reinvasion. A) Baited C optotermes formosanus has been eliminated. B) R eticulitermes flavipes (green) has re invaded and taken over C. formosanus former territory .

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55 Discussion The gallery structure of these species may lend to their behavior when interacting. Puche and Su (2001) suggested that the short, wide tunnels of C. formosanus and the longer, thinner tunnels of R. flavipes may be due to differing foraging strategie s. However, R. flavipes blocked these longer, thinner tunnels multiple times while retreating from C. formosanus , suggesting that their tunnels may take on a defensive purpose. Though C. formosanus were able to qu ickly reach R. flavipes in large numbers within their wider tunnels, they were not able to attack large numbers of R. flavipes , as many of their population retreated into their thinner tunnels, blocking them as they moved. This is supported by the relative ly few dead observed upon intersection of tunnels, as compared with intraspecies confrontations within C. formosanus . It should be noted that in a field situation, R. flavipes may have the opportunity to continue to avoid confrontation , as this bioassay li mits movement . In one replicate , R. flavipes reinvaded baited C. formosanus territory, which supports the field observation of R. flavipes entering territories of C. formosanus after their elimination (Messenger et al. 2005) . In the other two rep licates in which R. flavipes did not attempt to invade the neighboring panel even after C. formosanus was weakened by bait , it is possible that tunnel volume and termite density was not great enough to warrant movement into other territories ( Su and Lee 2009 ) . Reticulitermes spp. have been observed reinvading monitoring stations once other Reticulitermes spp. have b een eliminated (Getty, 2000, Vargo 2003), so it is likely that with further replication, we might see R. flavipes regularly reinvade C. formosanus as observed in this study. The subsequent elimination of R. flavipes after entering a former

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56 C. formosanus ba ited panel suggests that control can be maintained over an area by continuing a baiting program.

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57 CHAPTER 4 CONCLUSION Intraspecies experiments showed C. formosanus alway s reinvade neighboring territories . They use d old galleries of eliminated termites and feed on bait left behind, and subsequently wer e eliminated themselves. R. flavipes interspecies intera ctions differ in that they retreat ed from confrontation with C. formosanus when possible. R. flavipes may invade a neighboring weakened C. formosanus population, although more replications would show a clearer picture of these behaviors. Regardless, reinvading R. flavipes were eliminated by bait left in C. formosanus galleries. This supports maintaining a baiting program in areas of high term ite pressure for management and control of C. formosanus and R. flavipes infestations .

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58 LIST OF REFERENCES Alibert, J., and R. Martoja. 1976. Bioaccumulation minérale et purique chez les termites. Insect . Soc . 23: 49 73. Cabrera, B. J., and E. M. Thoms. 2006. Versatility of baits containing noviflumuron for control of structural infestations of Formosan subterranean termites (Isoptera: Rhinotermitidae). Fl a. Entomol . 89: 20 31. Chouvenc, T., P. Bardunias, H. F. Li, M. L. Elliott, and N. Y. Su. 2011. Planar arenas for use in laboratory bioassay studies of subterranean termites (Rhinotermitidae). Fl a. Entomol . 94: 817 826. Cornelius, M. L., and W. L. Osbrink. 2003. Agonistic interactions between colonies of the Formosan subterranean termi te (Isoptera: Rhinotermitidae) in New Orleans, Louisiana. Environ . E ntomol . 32: 1002 1009. Deheer, C. J., and E. L. Vargo. 2004. Colony genetic organization and colony fusion in the termite Reticulitermes flavipes as revealed by foraging patterns over time and space. Mol . Ecol . 13: 431 441. Getty, G. M., M. I. Haverty, K. A. Copren, and V. R. Lewis. 2000. Response of Reticuliterme s spp. (Isoptera: Rhinotermiti dae) in Northern California to b aiting with Hexaflumuron with Sentricon Termite Colony Elimination System. J . Econ . Entomol . 93: 1498 1507. Grace, J., C. Tome, T. Shelton, R. Oshiro, and J. Yates. 1996. Baiting studies and consideration with Coptotermes formosanus (Isoptera: Rhinotermitidae) in Hawaii. Sociobiology 28: 511 520. Grace, J. K., and N. Y. Su. 2001. Evidence supporting the use of termite baiting systems for long term structural protection (I soptera). Sociobiology 37: 301 310. Grassé, P. 1982. Termitologia: Anatomie, Physiologie, Reproduction des Termites. Vol I. Masson, Paris. Haverty, M. 1979. Soldier production and maintenance of soldier proportions in laboratory experimental groups of Co ptotermes formosanus Shiraki. Insect . Soc . 26: 69 84. Haverty, M., and R. Howard. 1981. Production of soldiers and maintenance of soldier proportions by laboratory experimental groups of Reticulitermes flavipes (Kollar) and Reticulitermes virginicus (Bank s)(Isoptera: Rhinotermitidae). Insect . Soc . 28: 32 39.

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59 Haverty, M. I., N. Y. Su, M. Tamashiro, and R. Yamamoto. 1989. Concentration dependent presoldier induction and feeding deterrency: potential of two insect growth regulators for remedial control of t he Formosan subterranean termite (Isoptera: Rhinotermitidae). J . Econ. Entomol. 82: 1370 1374. Husseneder, C., J. Grace, M. Messenger, E. Vargo, and N. Y. Su. 2003. Describing the spatial and social organization of Formosan subterranean termite colonies i n Armstrong Park, New Orleans. Sociobiology 41: 61 66. Jones, S. C. 1984. Evaluation of two insect growth regulators for the bait block method of subterranean termite (Isoptera: Rhinotermitidae) control. J . Econ. Entomol. 77: 1086 1091. Jones, S. C. 1993. Field observations of intercolony aggression and territory changes in Heterotermes aureus (Iso ptera: Rhinotermitidae). J. I nsect B ehav . 6: 225 236. Karr, L. L., J. J. Sheets, J. E. King, and J. E. Dripps. 2004. Laboratory p erformance and p harmacokin etics of the benzoylphenylurea noviflumuron in eastern subterranean t ermites (Isoptera: Rhinotermitidae). J . Econ . Entomol . 97: 593 600. La Fage, J., N. Su, M. Jones, and G. Esenther. 1983. A rapid method for collecting large numbers of subterranean termi tes from w ood . Sociobiology 7. Levings, S. C., and E. S. Adams. 1984. Intra and interspecific territoriality in Nasutitermes (Isoptera: Termitidae) in a Panamanian mangrove forest . J . Anim . Ecol . 705 714. Li, H. F., R. L. Yang, and N. Y. Su. 2010. Interspecific competition and territory defense mechanisms of Coptotermes formosanus and Coptotermes gestroi (Isoptera: Rhinotermitidae). Environ . E ntomol . 39: 1601 1607. Messenger, M., and N. Y. Su. 2005a. Colony characteristics and seasonal activity of the Formosan subterranean termite (Isoptera: Rhinotermitidae) in Louis Armstrong Park, New Orleans, Louisiana. J . Entomol. Sci. Messenger, M. T., and N. Y. Su. 2005b. Agonistic be havior between colonies of the F ormosan subterranean termite (Isoptera: Rhinotermitidae) from Louis Armstrong Park, New Orleans, Louisiana. Sociobiology 45: 331 345. Messenger, M. T., N. Y . Su, C. Husseneder, and J. K. Grace. 2005. Elimination and reinvasion s tudies with Coptoterme s formosanus (Isoptera: R hinotermitidae) in Louisiana. J . Econ. Entomol. 98: 916 929.

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60 Mullins, A. J., N. Y. Su, and C. Owens. 2011. Reinvasion and colony e xpansion of Coptotermes formosanus (Isoptera: Rhinotermitidae) after areawide e limination. J . Econ . Entomol . 104: 1687 1697. Puche, H., and N. Y . Su. 2001. Tunnel formation by Reticulitermes flavipes and Coptotermes formosanus (Isoptera: Rhinotermitidae) in response to wood in sand. J . E con . Entomol. 94: 1398 1404. Randall, M., and T. C. Doody. 1934. P oison dusts. I. Treatments with poisonous dusts. Termites and termite control. Univ. Calif. Press, Berkeley 17: 463 476. Rust, M. K., and N. Y. Su. 2012. Managing social insects of urban importance. Ann . R ev . E ntomol . 57: 355 375. Su, N. Y. 1991. Evaluation of bait toxicants for suppression of subterranean termite populations. Sociobiology 19: 1 220. Su, N. Y. 1994. Field evaluation of a hexaflumuron bait for population suppression of subterranean termites (Isoptera: Rhinotermitidae). J . Econ . En tomol . 87: 389 397. Su, N. Y. 2005. Response of the Formosan subterranean t ermites (Isoptera: Rhinotermitidae) to baits or nonrepellent termiticides in e xt ended foraging a renas. J . Econ. Entomol. 98: 2143 2152. Su, N. Y. 2013. Estimating population size of large laboratory colonies of the Formosan subterranean termite using the capture probability equilibrium. J . Econ. Entomol. 106: 2442 2447. Su, N. Y., and M. Tamashiro. 1987. An overview of the Formosan subterranean termite (Isoptera: Rhinotermitidae) in the world. Research extension series College of Tropical Agriculture and Human Resources, University of Hawaii, Cooperative Extension Service (USA). Su, N. Y., and R. H. Scheffrahn. 1988a. Intra and interspecific competition of the Formosan and the eastern subterranean termite: Evidence from field observations (Isoptera: Rhin otermitidae). Sociobiology . Su, N. Y., and R. H. Scheffrahn. 1988b. Foraging population and territory of the Formosan subterranean termite(Isoptera: Rhi notermitidae) in an urban environment. Sociobiology 14: 353 360. Su, N. Y., and R. H. Scheffrahn. 1989. Comparative effects of an insect growth regulator, S 31183, against the Formosan subterranean termite and eastern subterranean termite (Isoptera: Rhino termitidae). J . Econ. Entomol. 82: 1125 1129.

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61 Su, N. Y., and R. H. Scheffrahn. 1990. Economically important termites in the United States and their control. Sociobiology 17: 77 94. Su, N. Y., and M. I. Haverty. 1991. Agonistic behavior among colonies of t he Formosan subterranean termite, Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae), from Florida and Hawaii: Lack of correlation with cuticular hydrocarbon composition. J . Insect Beh a v . 4: 115 128. Su, N. Y., and R. H. Scheffrahn. 1993. Laborato ry evaluation of two chitin synthesis inhibitors, hexaflumuron and diflubenzuron, as bait toxicants against Formosan and eastern subterranean termites (Isoptera: Rhinotermitidae). J . Econ. Entomol. 86: 1453 1457. Su, N. Y., and R. H. Scheffrahn. 1996. Fate of subterranean termite colonies (Isoptera) after bait applications -an update and review. Sociobiology. Su, N. Y., and R. H. Scheffrahn. 1998. A review of subterranean termite control practices and prospects for integrated pest management programme s. Integrated Pest Management Rev . 3: 1 13. Su, N. Y., and R. H. Scheffrahn. 2000. Termites as pests of buildings, pp. 437 453, Termites: evolution, sociality, symbioses, ecology. Springer. Su, N. Y., and S. H. Lee. 2009. Tunnel volume regulation and gro up size of subterranean termites (Isoptera: Rhinotermitidae). Ann . Entomol . Soc . Am . 102: 1158 1164. Su, N. Y., M. Tamashiro, and M. I. Haverty. 1985. Effects of three insect growth regulators, feeding substrates, and colony origin on survival and presold ier production of the Formosan subterranean termite (Isoptera: Rhinotermitidae). J . Econ. Entomol. 78: 1259 1263. Su, N. Y., P. M. Ban, and R. H. Scheffrahn. 1991a. Evaluation of twelve dye markers for population studies of the eastern and Formosan subterranean termite (Isoptera: Rhinotermitidae). Sociobiology 19: 349 362. Su, N. Y., P. M. Ban, and R. H. Scheffrahn. 1991b. Suppression of foraging populations of the Formosan subterranean termite (Isoptera: Rhinotermitidae) by field applications of a slow acting toxicant bait. J . Econ. Entomol. 84: 1525 1531. Su, N. Y., P. M. Ban, and R. H. Scheffrahn. 1993. Foraging populations and territories of the eastern subterra nean termite (Isoptera: Rhinotermitidae) in southeastern Florida. Environ . E ntomol . 22: 1113 1117.

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62 Su, N. Y., J. D. Thomas, and R. H. Scheffrahn. 1998. Elimination of subterranean termite populations from the Statue of Liberty National Monument using a bait matrix containing an insect growth regulator, hexaflumuron. J . American Inst . Conserv . 37: 282 292. Su, N. Y., M. Tamashiro, J. R. Yates, and M. I. Haverty. 1982. Effect of behavior on the evaluation of insecticides for prevention of or remedial cont rol of the Formosan subterranean termite. J . Econ. Entomol. 75: 188 193. Su, N. Y., and R. H. Scheffrahn. 1986. A method to access, trap, and monitor field populations of the Formosan subterranean termite(Isoptera: Rhinotermitidae) in the urban environmen t. Sociobiology 12: 299 304. Vargo, E. L. 2003. Genetic structure of Reticulitermes flavipes and R. virginicus (Isoptera: Rhinotermitidae) colonies in an urban habitat and tracking of colonies following treatment with hexaflumuron bait. Environ . E ntomol . 32: 1271 1282. Xing, L., T. Chouvenc, and N. Y. Su. 2014. Behavioral and histological changes in the F ormosan subterranean t ermi te (Isoptera: Rhinotermitidae) induced by the chitin s ynt hesis inhibitor noviflumuron. J. Econ. Entom o l. 107: 741 747.

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63 BIOGRAPHICAL SKETCH As a child, Sarah would often be found peering into the grass with a butterfly net and an insect carrier in tow. the as no surprise to friends and family that she came to take an interest in e ntomology. Sarah grew up in Boca Raton, Florida, immersed in a culture of insects and science. She was encouraged by her father to be curious and to always continue learning. Her family owned and operated a pest contro l service business in her youth, and later the family took on a new role as a manufacturer of pest control products. After high s chool, Sarah continued on to earn her Bachelor of Science degree in b iology from Florida State University in Tallahassee before later joining the termite lab at the Ft. Lauderdale Research and Education Center in Davie. There she studied social insects and earned her Master of Science degree from the University of Florida w ith Dr. Nan Yao Su as her Major Professor. Sarah currently has a keen interest in medical entomology and is knowledgeable in urban entomology and insect identification .